Wachendorfia

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Wachendorfia
Wachendorfia thyrsiflora Flipphi 6.jpg
Wachendorfia thyrsiflora
Wachendorfia thyrsiflora Flipphi 5.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Commelinales
Family: Haemodoraceae
Subfamily: Haemodoroideae
Genus: Wachendorfia
Burm., 1757, not Loefl. 1758 (Commelinaceae)
species
Synonyms [1]
  • PediloniaC.Presl

Wachendorfia is a genus of perennial herbaceous plants that is assigned to the bloodroot family. The plants have a perennial rootstock with red sap. From the rootstock emerge lance- or line-shaped, sometime sickle-shaped, pleated, simple leaves set in a fan, that are flattened to create a left and right surface rather than an upper and lower surface. The leaves die when the seeds are shed in three of the species, and are perennial in one species.

Contents

The rootstock also produces flowering stems annually that carry a panicle of zygomorphic, yellow or yellowish flowers in two distinct forms, one with the style and one stamen bent to the right and two stamens to the left, and the other vice versa. The fruit opens with three valves and each contains a single, hairy seed. All species only occur in the fynbos biome in the Cape Provinces of South Africa. [1]

Description

Pleated leaves on the stem of W. thyrsiflora Wachendorfia thyrsiflora Flipphi 10.jpg
Pleated leaves on the stem of W. thyrsiflora
W. thyrsiflora seeds Wachendorfia thyrsiflora seeds, by Omar Hoftun.jpg
W. thyrsiflora seeds

Wachendorfia is a genus of perennial herbaceous plants of 10–250 cm (0.33–8.20 ft) high when flowering, which emerge from a fleshy, bright red-coloured, rounded, egg-shaped to cylindrical underground rootstock. Three species become dormant for the dry, hot summer, surviving with their rootstocks, but one species (W. thyrsiflora) is evergreen. Flowering stems emerge from the bud at the tip of the rootstock. Each of the three side buds may grow into a new rootstock. The roots are thin and adventitious, clustering around the nodes between the old and the new rhizomes.

The leaf bases form a tunic around the old rhizome. The leaves together form a fan-like structure, are 0.1–0.9 m (0.33–2.95 ft) long, erect or spreading, simple and entire, line- to lance-shaped, sometimes sickle-shaped, have a firm texture, are flattened sideways as to produce left and right surfaces, not upper and lower surfaces and are always pleated along their length. The leaves are green or yellowish green, in W. thyrsiflora and W. brachyandra always hairless, but those in W. multiflora and W. paniculata may be hairless or roughly hairy. The base of the lower leaves form a sheath around the base of the inflorescence stem. Three leaves grow from the rootstock, and a variable number emerges from the stem. The lower pair of stem leaves are opposite and envelop the stem at their base; higher stem leaves are arranged in a spiral and do not envelop the stem. Wachendorfia has so-called paracytic stomata. [1]

The inflorescence stem of 0.1–2.5 m (0.33–8.20 ft) high dies down each year after flowering. The inflorescence is a lax to dense deltoid or a dense cylindrical panicle, with a herbaceous, cylindrical to angular axis that is covered in short hairs which become glandular nearer to the tips. The bracts in the inflorescence have a pointy tip, are usually glandularly hairy, and are dry, brown and papery in three species and herbaceous in W. multiflora, erect or with their tips recurved. The perianth is pale apricot to pale or golden yellow, strongly zygomorphic and consists of six subequal, spreading, oblong tepals, set in two whorls. The lower three tepals are free, and the upper three are united at their base where they are adorned with small dark and light markings that apparently function as a nectar guide. Two open spur-like nectaries are present between the base of the outer upper tepals and the adjacent inner tepals. The outer tepals are hairy on the outside, especially the upper one.

There are three stamens opposite the inner tepals. Their filaments are free, line-shaped, curving downward from the base and upwards towards the tip. The lateral filaments are longer and carry a smaller anther than the lowest stamen. The lowest filament turns sideways in the opposite direction of the style. The anthers on top of each filament are oval to arrow-shaped, open with slits opposite the side where the filament is attached, over the length of both lobes. The pollen is boat-shaped, with one furrow. The style is thread-shaped, bent left or right consistently in any one plant and carries a minute head-like stigma. The ovary is attached above the attachment of the other floral parts; it consists of three compartments that each contain one ovule attached to the centre and develops into a softly hairy, mostly glandular, sharply three-lobed dry capsule that is wider than high. The seed is oval to globular, approximately 2 mm (0.079 in) in diameter, densely covered in hair, and brown in colour, at least in W. paniculata, the only species for which it has been recorded. [1]

Differences between the species

W. thyrsiflora is a large to very large, 0.6–2 m (2.0–6.6 ft) high, evergreen herb with golden yellow flowers that have diverging stamens and style of at least two thirds as long as the tepals, in a dense spiky inflorescence high above the leaves, and dry, brownish and membranous bracts recurved at the tip, with leaves mostly much wider than 1.5 cm (0.59 in). It is restricted to damp environments.

W. paniculata is a small to large, 0.1–0.9 m (0.33–2.95 ft), deciduous herb with apricot, yellow or orange flowers that have diverging stamens and style of at least two thirds as long as the tepals that are 13–31 mm (0.51–1.22 in) long and 4–16 mm (0.16–0.63 in) wide, in a lax to dense panicle with brown, dry and membranous bracts that recurve at the tip with age, with leaves narrower than 2 cm (0.79 in) and shorter than the inflorescence, and that may grow in dry and wetter circumstances.

W. brachyandra has apricot to pale yellow flowers in a lax panicle that is longer than the leaves, clustered stamens, which are, like the style, less than half the length of the tepals.

W. multiflora is a small plant of up to 25 cm (9.8 in) high, with leaves that are usually longer than the very short and dense inflorescence, with green, erect bracts, dull yellow, later purplish brown flowers with narrow tepals, 15–25 mm (0.59–0.98 in) long and 3–6 mm (0.12–0.24 in) wide. [1]

Taxonomy

The first description of a species of butterfly-lily was already published in 1700 by the English botanist Leonard Plukenet in his book Almagesti botanici mantissa. He named it Erythrobulbus. The description was probably based on a collection of W. thyrsiflora made by Henrik Bernard Oldenland between 1689 and 1697. In 1739, Johann Philipp Breyne described and illustrated what is undoubtedly a Wachendorfia, giving it the name Asphodelus latifolius in his book Prodromus fasciculi rariorum plantarum, but these names predate the start of Linnaean taxonomy in 1753 and are therefore invalid. Johannes Burman described both W. thyrsiflora and W. paniculata in his monography of the genus Wachendorfia in 1757. Pehr Löfling had already assigned the name Wachendorfia to a genus in the Commelinaceae, but his work was published only two years after his death in 1756 by Linnaeus, and therefore, Burman's name has priority, and Löfling's homonym was replaced by Callisia .

Carl Peter Thunberg, who visited the Cape from 1772 to 1775, collected additional specimens of Wachendorfia, and Carl Linnaeus the Younger described some of these as W. graminifolia in 1781. Thunberg renamed the specimens described by Linnaeus, creating the invalid name W. graminea. Richard Anthony Salisbury created the names W. elata, W. humilis, W. pallida and W. brevifolia. Thunberg distinguished W. hirsuta and W. tenella, both in his 1811 book Flora Capensis - sistens plantas promontorii Bonæ Spei Africes - secundum systema sexuale emendatum. William Herbert distinguished W. paniculata var. β in Curtis's Botanical Magazine vol. 53 in 1826, which was raised to the status of species by Robert Sweet, who called it W. herbertii. In 1829, Karel Bořivoj Presl described Pedilonia violacea. [1] Friedrich Wilhelm Klatt described Babiana multiflora in 1882. [2] In 1949, Winsome Fanny Barker described W. brachyandra and W. parviflora (a later synonym of W. multiflora).

In their 1992 revision of the genus Wachendorfia, Nick Helme and Hans Peter Linder conclude that four species can be distinguished: W. thyrsiflora, W. paniculata, W. brachyandra and W. parviflora. [1] John Manning and Peter Goldblatt recognised in 2000 that Babiana multiflora should be assigned to the genus Wachendorfia and so create the new combination W. multiflora. It has priority over its synonym W. parviflora because the original combination was published earlier. [3] W. elata is a synonym of W. thyrsiflora, and all others should be treated as synonyms for W. paniculata, a very variable species, which shows a continuous intergrading of characters. [1]

Names

The genus Wachendorfia is named in honor of Evert Jacob van Wachendorff, professor of botany and chemistry and later rector at the University of Utrecht in the 18th century. [4]

Phylogeny

Comparison of homologous DNA has increased the insight in the phylogenetic relationships between the genera in the Haemodoroideae subfamily, and between the species of Wachendorfia. The following trees represent those insights. [5]

  subfamily Haemodoroideae 

Ecology

The flowers of the species of Wachendorfia are enantiomorphic. The style is sometimes sharply deflected to the right, while in other plants it is bent to the left. In both morphs one of the three stamens is deflected to the same side as the style, whereas the remaining two curve in the opposite direction. This is thought to be a mechanism that enhances crosspollination and so boosts genetic diversity. Flowers of the species of Dilatris are also chirally dimorph, but here both left- and right-handed flowers are found on the same plants.

Two species, W. thyrsiflora and W. brachyandra, grow in permanently moist environments such as the banks of streams and seeps, whereas the other two, W. multiflora and W. paniculata, grow in locations that are dry, at least seasonally dry. All species flower during spring and early summer, but there are differences in flowering time. W. multiflora has its blooms in August and September. W. paniculata has a long flowering period, rather later at higher altitudes. W. thyrsiflora can often flower during summer, probably reflecting that it does not suffer from water-stress because it grows in permanently moist environments. [1]

Cultivation

Slightly frost hardy, Wachendorfias thrive outdoors in warm, near frost-free environments as an ornamental plant. Most require moist, well drained soils in a sunny position. Some are ideal for bog gardens. Propagation is via seed or division. [6]

Related Research Articles

<i>Babiana</i> Genus of flowering plants

Babiana is a genus of geophytes in the family Iridaceae with 93 recognized species as of March 2022. The leaves consist of a stalk and a blade that are at an angle to each other. The leaf blades are entire, laterally flattened and pleated, and often hairy. Each individual flower is subtended by two hairy or smooth bracts that are green in most species. The outer bract is often the largest of the two. In most species the bracts have a dry, brown tip, but in a few species it is entirely green or entirely dry when flowering or the outer bract is translucent and has a papery texture. The inner bract is forked or split all the way to its base. Each flower is without a pedicel, with six tepals that are merged at their base into a tube and form a perianth that is mirror-symmetrical in most species, with three anthers implanted where the perianth tube widens and that are, in almost every species, clustered at one side of the style. The style has three branches that widen towards the tip and the ovary is inferior. Flowers occur in almost every conceivable colour, many have markings on some of the tepals, and few star-symmetrical flowers have a centre that strongly contrasts with the free part of the perianth. The majority of these species are endemic to the west and southwest of South Africa, and southwestern Namibia, but one species occurs elsewhere in Namibia and South Africa and another species can be found in Botswana, Namibia, South Africa, Zambia and Zimbabwe. The genus name is derived from the Dutch word baviaan, referring to the Chacma baboon, Papio ursinus, that consumes the corms of plants in the genus. The genus is called bobbejaantjie in Afrikaans, meaning small baboon.

<i>Babiana ambigua</i> Species of flowering plant

Babiana ambigua is a species of plant in the Iridaceae. It is endemic to the Western Cape province of South Africa. It is a geophyte, that appears from an underground corm every year and grows to a height of 5–8 cm (2.0–3.1 in) or occasionally up to 16 cm (6.3 in). Its leaves are pleated, hairy, lance-shaped, reaching higher than the inflorescence. The fragrant, mirror-symmetric, blue to mauve flowers, consist of six tepals merged into a tube at their base, but with free lobes at the top. The lower lateral tepals have whitish markings accentuated by a more intense blue line along their margin. Each flower is supported by two green bracts sometimes with a brownish tip, and the inner bract is divided entirely to its base. There are three anthers crowded to the dorsal side of the perianth and a style divided in three branches on top of a smooth ovary. Flowers can be found from late July at sea level to the end of September at high altitude.

<i>Babiana nana</i> Species of flowering plant

Babiana nana is a species of geophyte of 6–15 cm (2.4–5.9 in) high that is assigned to the family Iridaceae. It has leaves that consist of a sheath and a blade that are at an angle with each other. The leaf blades are oval to almost line-shaped and have a left and right surface, rather than an upper and lower surface. The leaf blades are moderately pleated and covered in dense, soft hairs. The inflorescence contains two to six blue to violet or pale pink flowers adorned with white markings on the lower lip, and with three stamens crowding under the upper lip. Flowering occurs from late August to the end of September. The flowers emit a smell reminiscent of roses or violets.

<i>Dilatris</i> Genus of flowering plants

Dilatris is a genus of four species of evergreen perennial herbaceous plants of up to 60 cm (2.0 ft) high, that are assigned to the bloodroot family. The plants have hairless, line- to lance-shaped leaves set in a fan that emerges from a red or orange coloured rootstock. Six free tepals with some gland dots near their tips are present on the mauve or dirty yellow flowers' six free petals. The other two stamens are longer and spreading with smaller scarlet anthers, while the one stamen is short, upright, and has a large, yellow anther. The style is diverted from the centre opposite both longer stamens. The species only occur in the Western Cape and Northern Cape provinces of South Africa.

<i>Wachendorfia thyrsiflora</i> Species of flowering plant

Wachendorfia thyrsiflora, the marsh butterfly lily, is a plant species of 0.6–2.5 m (2.0–8.2 ft) high when flowering, that has been assigned to the bloodroot family. It is a large to very large evergreen perennial plant with an underground rootstock with clusters of roots produced at the nodes. The rootstock has a distinctive red colour that results from so-called arylphenalenone pigments. The sturdy, entire and broadly sword-shaped leaves have laterally flattened and pleated leaf blades. The golden yellow flowers are set a dense cylindrical panicle on a tall firm stalk. Flowering occurs from spring until mid-summer.

<i>Wachendorfia paniculata</i> Species of flowering plant

Wachendorfia paniculata is a species of plant of 10–90 cm (3.9–35.4 in) high, that emerges during the winter from an underground rootstock. It has entire, sword-shaped, mostly hairy, line- to lance-shaped, straight or sickle-shaped leaves, set in a fan at ground level with a lax to dense panicle consisting of pale apricot to yellow mirror-symmetric flowers with six tepals, three stamens and a undivided style that curves either to the right or left. The species is assigned to the bloodroot family. Flowering occurs between August and December at sea level, and until early February at high altitude, with a distinct peak from September to November. It can only be found in the Cape provinces of South Africa. Like other species of Wachendorfia, it is called butterfly lily in English and rooikanol or spinnekopblom in Afrikaans, and this species in particular is also called koffiepit in Afrikaans.

<i>Wachendorfia brachyandra</i> Species of flowering plant

Wachendorfia brachyandra is a small to large, 10–65 cm (3.9–25.6 in) high, winter-growing, perennial herbaceous plant that grows from a rootstock, and has been assigned to the bloodroot family. Its simple, entire, line- to lance-shaped leaves that are usually shorter than the stem, and have pleated, laterally flattened leaf blades af about 35 mm (1.4 in) wide, meaning that there are left and right surfaces rather than upper and lower. The inflorescence is a lax panicle and at the base of each flowerstalk is a dry, brown and papery bract, and those higher in the panicle have a recurved tip. The mirror-symmetrical pale apricot-yellow flowers consist of six tepals and are adorned with brown markings on the upper three tepals. There are three anthers that are clustered and about half as long as the tepals. Each individual flower only lasts one day. Flowering occurs from August to December. This species grows in the wild in the Western Cape province of South Africa only, and is much less common than its relatives W. paniculata and W. thyrsiflora. It is sometimes called short-stamen butterfly-lily in English.

<i>Babiana vanzijliae</i> Species of flowering plant

Babiana vanzijliae is a species of geophyte of 4–12 cm (1.6–4.7 in) high that is assigned to the family Iridaceae. It has leaves that consist of a sheath and a blade that are at an angle with each other. The leaf blades are narrow, sword- to lance-shaped and have a left and right surface, rather than an upper and lower surface. The leaf blades are pleated and covered in velvety hairs. The inflorescence contains three to five pale bluish mauve to yellow flowers, but the lower lateral tepals are yellow becoming pale around the edges, and with three stamens crowding under the upper lip. Flowering occurs from early August to the middle of September. The flowers emit a strong scent. B. vanzijliae grows along the Bokkeveld Escarpment near Nieuwoudtville in the Northern Cape province of South Africa.

<i>Babiana lobata</i> Species of flowering plant

Babiana lobata is a species of geophyte of 12–25 cm (4.7–9.8 in) high that is assigned to the family Iridaceae. It has leaves that consist of a sheath and a blade that are at a slight angle with each other. The leaf blades are narrow, sword- to lance-shaped and have a left and right surface, rather than an upper and lower surface. The leaf blades are slightly pleated and hairless. The inflorescence contains seven to twelve bluish mauve mirror-symmetrical flowers comprising six tepals, with the lower lateral tepals yellow sometimes flushed mauve at their tips, and with three stamens crowding under the dorsal tepal. Flowering occurs in July and August. The flowers emit a faint acrid-metallic scent. B. lobata grows in part of the Richtersveld in the Northern Cape province of South Africa.

<i>Babiana bainesii</i> Species of flowering plant

Babiana bainesii is a species of geophyte of 15–25 cm (5.9–9.8 in) high that is assigned to the family Iridaceae. It sometimes grows in tufts. The approximately upright leaf blades appear directly from the ground, are narrow, sword- to line-shaped and have a left and right surface, rather than an upper and lower surface, and far exceed the flowers in length. The leaf blades are pleated. The inflorescence stem is fully underground and often branched. It contains two to eight blue, violet or mauve mirror-symmetrical flowers comprising six tepals. B. bainesii has a wide distribution and occurs in Botswana, Namibia, South Africa, southernmost Zambia and Zimbabwe.

<i>Babiana villosa</i> Species of flowering plant

Babiana villosa is a species of geophyte of 10–20 cm (3.9–7.9 in) high that is assigned to the family Iridaceae. It has mauve-pink, purple or scarlet star-symmetrical wide chalice-shaped flowers with narrow tube, large, blackish or dark purple anthers, and velvety hairy, lance-shaped, laterally compressed leaves, set in a fan. Flowers occur during August and September. It grows between Malmesbury and Wellington in the Western Cape province of South Africa. It is commonly called red babiana in English and rooibobbejaantjie in Afrikaans.

<i>Babiana hirsuta</i> Species of flowering plant

Babiana hirsuta is a species of geophyte of 40–70 cm (16–28 in) high that is assigned to the family Iridaceae. It has many scarlet mirror-symmetrical flowers in a branched inflorescence with several short ascending branches. The flower has a narrow tube, and three large, blackish or dark purple anthers that extend beyond the dorsal tepal. The leaves are velvety hairy, lance-shaped, laterally compressed and set in a fan. It is an endemic species of South Africa that can be found along the west coast of the Northern and Western Cape provinces as far south as Saldanha. It is called red babiana in English, but that name is also applied to Babiana villosa, and strandlelie, sandlelie and rooihanekam in Afrikaans. Until 2008, the strandlelie was known as Babiana thunbergii.

<i>Babiana tubiflora</i> Species of flowering plant

Babiana tubiflora is a species of geophyte of 7–15 cm (2.8–5.9 in) high that is assigned to the family Iridaceae. It has whitish mirror-symmetrical flowers with a long narrow tube that split into six tepal lobes, have three stamens, and line- to lance-shaped, laterally compressed leaves. It is an endemic species of South Africa that can be found along the west and south coast of the Western Cape province. It flowers from August to early October.

<i>Babiana sambucina</i> Species of flowering plant

Babiana sambucina is a species of geophyte of 8–30 cm (3.1–11.8 in) high that is assigned to the family Iridaceae. It has dense spikes of blue to violet-coloured, often fragrant flowers. There are two subspecies, B. sambucina subsp. longibracteata is restricted to a small area in the Northern Cape, B. sambucina subsp. sambucina grows in the Western Cape and Eastern Cape provinces of South Africa. Flowers are present in August and September.

<i>Babiana brachystachys</i> Species of flowering plant

Babiana brachystachys is a species of geophyte of 20–30 cm (7.9–11.8 in) high that is assigned to the family Iridaceae. It has cream or pale pink, only slightly mirror-symmetrical flowers that are pink on the reverse with a long narrow tube that splits into six tepal lobes, three stamens, a style that divides in three branches opposite the tip of the anthers and line-shaped leaves that are circular in cross section. It is an endemic species of South Africa that can be found on sandy hills and inland dunes in the Northern Cape province. It flowers in September and October.

<i>Babiana scabrifolia</i> Species of flowering plant

Babiana scabrifolia is a perennial plant of 5–15 cm (2.0–5.9 in) high that annually forms leaves and flowers from an underground corm and is assigned to the Iris family. It produces relatively large, pale lilac or blue flowers slightly above the soil, and soft, lightly pleated leaves that reach beyond the flowers. These leaves are lance-shaped to oblong in adults, but line-shaped and twisted in non-flowering specimens. Flowering occurs from June to August. The species can be found in the Western Cape province of South Africa.

<i>Babiana hypogaea</i> Species of flowering plant

Babiana hypogaea is a perennial plant of about 5–8 cm (2.0–3.1 in) high that annually forms leaves and flowers from an underground corm that is assigned to the iris family. It has pale greenish yellow, buff or rarely white, mirro-symmetrical flowers and line- to sickle-shaped, slightly pleated, thinly hairy or hairless leaves that are held horizontally or inclined, that grows in the summer rainfall areas of southeastern Namibia and central and northern central South Africa. Flowers may be found between June and September and sometimes between December and May. It is called Bobbejaankalkoentjie, Bobbejaanuintjie or Ertappeluintjie in Afrikaans.

<i>Babiana mucronata</i> Species of flowering plant

Babiana mucronata is a perennial plant species that grows to about 5–18 cm (2.0–7.1 in) high and annually forms leaves and flowers from an underground corm. It is assigned to the iris family. It has a simple or branched, more or less upright spike of 3-12 dark to pale violet-blue, mirror-symmetrical flowers. Each flower consists of a perianth that is merged below into a funnel-shaped tube of 10–25 mm (0.39–0.98 in) long but splits into six unequal tepals. Three stamens are curved, crowded near the upper lip, and carry pale violet anthers. Flowers may be found between late July and September.

<i>Babiana torta</i> Species of flowering plant

Babiana torta is a species of geophyte of 7–15 cm (2.8–5.9 in) high that is assigned to the family Iridaceae. It has pale mirror-symmetrical flowers with a long narrow tube that split into six tepal lobes, have three stamens, and line- to lance-shaped, laterally compressed, undulating and coiled leaves. It is an endemic species of South Africa that can be found in the extreme north of the Western Cape province and the coastal mountains of the Northern Cape south of Namaqualand. It flowers from mid May to mid June.

<i>Babiana grandiflora</i> Species of flowering plant

Babiana grandiflora is a perennial plant of 8–14 cm (3.1–5.5 in) high that emerges at the start of every growing season from an underground corm. It has pleated sideways flattened leaves. It can be distinguished from other species of bobbejaantjie mainly by its very large, mirror-symmetric deep to pale bluish flowers near the ground. It also has very short filaments crowded near the dorsal tepal, resulting in the anthers being largely hidden within the floral cup of the funnel-shaped perianth. The perianth is entirely filled by the style in the bottom half of the tube. The flowers have a strong acrid-spicy, violet-like scent. Babiana grandiflora is native to the coastal plains of Namaqualand, South Africa. The continued survival of this species is considered being of least-concern.

References

  1. 1 2 3 4 5 6 7 8 9 Helme, N.A.; Linder, H.P. (1992). "Morphology, evolution and taxonomy of Wachendorfia (Haemodoraceae)" (PDF). Bothalia. 22 (1): 59–75. doi:10.4102/abc.v22i1.826.
  2. ↑ Royal Botanical Gardens Kew. "Babiana multiflora Klatt". Plants of the World Online.
  3. ↑ "Wachendorfia multiflora (Klatt) J.C.Manning & Goldblatt". The Plantlist.
  4. ↑ Alice Notten (2001). "Wachendorfia thyrsiflora". PlantZAfrica.com. South African National Biodiversity Institute.
  5. ↑ Hopper, Stephen D.; Smith, Rhian J.; Fay, Michael F.; Manning, John C.; Chase, Mark W. (2009). "Molecular phylogenetics of Haemodoraceae in the Greater Cape and Southwest Australian Floristic Regions". Molecular Phylogenetics and Evolution. 51 (1): 19–30. Bibcode:2009MolPE..51...19H. doi:10.1016/j.ympev.2008.11.015. PMID   19063982.
  6. ↑ Flora, The Gardeners Bible, ABC Publishing, Ultimo, NSW, Australia, 2005
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