''Heliophila'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

s in the family

Brassicaceae Brassicaceae () or (the older but equally valid) Cruciferae () is a medium-sized and economically important Family (biology), family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous pla ...

. Members of this genus are either annuals or perennials and some are popular as

ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...

s. Endemic to southern Africa, the majority of the approximately 80 species grow in South Africa, particularly the

Cape Floristic Region The Cape Floral Region is a floristic region located near the southern tip of South Africa. It is the only floristic region of the Cape Floristic Kingdom, and includes only one floristic province, known as the Cape Floristic Province. The Cap ...

, while a few extend into the

Namib Desert The Namib ( ; ) is a coastal desert in Southern Africa. According to the broadest definition, the Namib stretches for more than along the Atlantic coasts of Angola, Namibia, and northwest South Africa, extending southward from the Carunjamba Ri ...

Taxonomic history

The first recognised published description of ''Heliophila'' appears in the second edition of ''

Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genus, genera. It is the first work to consistently apply binomial nomenclature ...

'' by

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

, printed in 1763. He attributes his description, however, to

Nicolaas Laurens Burman Nicolaas Laurens Burman (27 December 1734 – 11 September 1793) was a Dutch botanist. He was the son of Johannes Burman (1707–1780). He succeeded his father to the chair of botany at the Athenaeum Illustre of Amsterdam., and at the Hortus Bot ...

, a Dutch botanist. The generic name ''Heliophila'' is Latin but originally derived from the

Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...

words ''hēlios'' (ἥλιος), meaning "sun", and ''phílos'' (φίλος), meaning "dear" or "beloved", and refers to the blooms opening in sun and closing at night or in overcast conditions, a habit of many of the species within the genus. In general, contention abounds regarding phylogenetic relationships in the family Brassicaceae. In the past, using morphological characters to establish tribes and genera was misleading and created dissension among

systematist Systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: phylogenetic trees, phylogenies). Phyl ...

s. The classification of species within Brassicaceae is gradually resolving through

molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

investigation, using

internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...

(ITS) data and other new-found sequencing knowledge. Previously there were six genera within the tribe Heliophileae, all endemic to southern Africa: ''Heliophila'', ''Cycloptychis'', ''Schlechteri'', ''Silicularia'', ''Thlaspeocarpa'', and ''Brachycarpaea''. The latter five genera contained among them only seven species, having been differentiated from the genus ''Heliophila'' mainly on morphological differences in the fruits they bore. Following extensive analysis of molecular evidence (nuclear (ITS) and plastid (trnL-F) DNA sequence data), Heliophileae was found to be monophyletic and the five small genera were reduced to synonymy with ''Heliophila''.

Distribution

''Heliophila'' is one of only two genera of Brassicaceae endemic to southern Africa. The majority of the species grow in the winter-rainfall Cape Floristic Region and the more arid

Namaqualand Namaqualand ( Khoikhoi: "Nama-kwa" meaning Nama Khoi people's land) is an arid region of Namibia and South Africa, extending along the west coast over and covering a total area of . It is divided by the lower course of the Orange River int ...

, while a smaller portion extend to the rest of South Africa, Namibia, Eswatini, and Lesotho. None of the members of this genus are listed on the

IUCN Red List of Threatened Species The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological spe ...

Description and habit

''Heliophila'' is regarded as one of the most diverse genera in the family Brassicaceae. Flower size, for example, varies greatly, with this genus containing both the largest of petals within the family (''H. juncea'') to the smallest (''H. pectinata''). There is a range of flower colour, with blue, white, and pink being the most common. Blue is an unusual colour for Brassicaceae, being known in only one other genus, the unrelated '' Solms-laubachia'' from the

Himalayas The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than list of h ...

. Within the genus are mainly

herb Herbs are a widely distributed and widespread group of plants, excluding vegetables, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal purposes, or for fragrances. Culinary use typically distingu ...

s and

subshrub A subshrub (Latin ''suffrutex'') or undershrub is either a small shrub (e.g. prostrate shrubs) or a perennial that is largely herbaceous but slightly woody at the base (e.g. garden pink and florist's chrysanthemum). The term is often interch ...

s, although

shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...

s and

liana A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...

s appear as well. They may be annual or perennial and the majority of the fruits produced by species in this genus are

dehiscent Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part. Structures that op ...

, not woody, and lack a carpophore. The plants are generally either

glabrous Glabrousness () is the technical term for a lack of hair, down, setae, trichomes, or other such covering. A glabrous surface may be a natural characteristic of all or part of a plant or animal, or be due to loss because of a physical condition, ...

or possess simple hairs. Leaves are variable; they are narrow or broad, but usually simple, and may be entire, lobed, or pinnatipartite. The

inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...

s are usually the

raceme A raceme () or racemoid is an unbranched, indeterminate growth, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are ...

type.

Species

Accepted species: * '' Heliophila acuminata'' (

Eckl. Christian Friedrich Ecklon (17 December 1795 – 1 December 1868) was a Danish botanical collector and apothecary. Ecklon is especially known for being an avid collector and researcher of plants in South Africa. Biography Ecklon was from Åbenr ...

& Zeyh.) Steud, 1840 * '' Heliophila adpressa'' O.E. Schulz * '' Heliophila affinis''

Sond. Otto Wilhelm Sonder (18 June 1812, Bad Oldesloe – 21 November 1881) was a German botanist and pharmacist. Life A native of Holstein, Sonder studied at Kiel University, where he sat pharmaceutical examinations in 1835, before becoming the prop ...

, 1846 * '' Heliophila africana'' ( L.) Marais, 1970 * '' Heliophila alpina'' Marais * '' Heliophila amplexicaulis'' L.f., 1782 * '' Heliophila arenaria'' Sond. * '' Heliophila arenosa''

Schltr. Friedrich Richard Rudolf Schlechter (16 October 1872 – 16 November 1925) was a German taxonomist, botanist, and author of several works on orchids. He went on botanical expeditions in Africa, Indonesia, New Guinea, South and Central America an ...

, 1899 * '' Heliophila brachycarpa''

Meisn. Carl Daniel Friedrich Meissner (1 November 1800 – 2 May 1874) was a Swiss botanist. Biography Born in Bern, Switzerland on 1 November 1800, he was christened Meisner but later changed the spelling of his name to Meissner. For most of his 40 ...

, 1842 * '' Heliophila brassicaefolia'' Eckl. & Zeyh. * '' Heliophila brassicifolia'' Eckl. & Zeyh. * '' Heliophila bulbostyla'' Barnes * '' Heliophila callosa'' (L.f.)

DC. Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...

, 1821 * '' Heliophila carnosa'' ( Thunb.) Steud., 1840 * '' Heliophila cedarbergensis'' Marais * '' Heliophila cinerea'' Marais, 1970 * '' Heliophila collina'' O.E. Schulz * '' Heliophila concatenata'' Sond., 1846 * '' Heliophila cornellsbergia'' B.J. Pienaar & Nicholas * '' Heliophila cornuta'' Sond., 1846 * '' Heliophila carnosa'' (Thunb.) Steud., 1840 * '' Heliophila coronopifolia'' L. * '' Heliophila crithmifolia''

Willd. Carl Ludwig Willdenow (22 August 1765 – 10 July 1812) was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants. Willdenow was also ...

, 1809 * '' Heliophila cuneata'' Marais * '' Heliophila decurva'' Schltr., 1913 * '' Heliophila deserticola'' Schltr., 1913 * '' Heliophila diffusa'' (Thunb.) DC. * '' Heliophila digitata'' L.f., 1782 * '' Heliophila dissecta'' Thunb. * '' Heliophila dregeana'' Sond., 1846 * '' Heliophila elata'' Sond. * '' Heliophila elongata'' DC. * '' Heliophila ephemera'' P.A.Bean * '' Heliophila esterhuyseniae'' Marais * '' Heliophila eximia'' Marais * '' Heliophila filicaulis'' Marais * '' Heliophila formosa'' Hilliard & B.L.Burtt * '' Heliophila gariepina'' Schltr. * '' Heliophila glauca'' Burch. ex DC. * '' Heliophila hurkana'' Al-Shehbaz & Mumm. * '' Heliophila juncea'' (P.J.Bergius) Druce * '' Heliophila katbergensis'' Marais * '' Heliophila laciniata'' Marais * '' Heliophila lactea'' Schltr. * '' Heliophila leptophylla'' Schltr. * '' Heliophila linearis'' DC. * '' Heliophila linoides'' Schltr. * '' Heliophila macowaniana'' Schltr. * '' Heliophila macra'' Schltr. * '' Heliophila macrosperma'' Burch. ex DC. * '' Heliophila maraisiana'' Al-Shehbaz & Mumm. * '' Heliophila meyeri'' Sond. * '' Heliophila minima'' (Stephens) Marais * '' Heliophila monosperma'' Al-Shehbaz & Mumm. * '' Heliophila namaquana'' Bolus * '' Heliophila namaquensis'' (Marais) Al-Shehbaz & Mumm. * '' Heliophila nubigena'' Schltr. * '' Heliophila obibensis'' Marais * '' Heliophila patens'' Oliv. * '' Heliophila pectinata'' Burch. ex DC. * '' Heliophila pendula'' Willd. * '' Heliophila pinnata'' L.f. * '' Heliophila polygaloides'' Schltr. * '' Heliophila promontorii'' Marais * '' Heliophila pubescens'' Burch. ex Sond. * '' Heliophila pusilla'' L.f. * '' Heliophila ramosissima'' O.E.Schulz * '' Heliophila refracta'' Sond. * '' Heliophila remotiflora'' O.E.Schulz * '' Heliophila rigidiuscula'' Sond. * '' Heliophila rimicola'' Marais * '' Heliophila scandens'' Harv. * '' Heliophila schulzii'' Marais * '' Heliophila scoparia'' Burch. ex DC. * '' Heliophila seselifolia'' Burch. ex DC. * '' Heliophila suavissima'' Burch. ex DC. * '' Heliophila suborbicularis'' Al-Shehbaz & Mumm. * '' Heliophila subulata'' Burch. ex DC. * '' Heliophila tabularis'' Dod * '' Heliophila thunbergii'' Steud. * '' Heliophila tricuspidata'' Schltr. * '' Heliophila trifurca'' Burch. ex DC. * '' Heliophila tulbaghensis'' Schinz * '' Heliophila variabilis'' Burch. ex DC.

References

{{Taxonbar, from=Q955407 Brassicaceae Brassicaceae genera