Gymnarrhena

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Gymnarrhena
Gymnarrhena micrantha.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Asterales
Family: Asteraceae
Subfamily: Gymnarrhenoideae
Tribe: Gymnarrheneae
Genus: Gymnarrhena
Desf.
Species:
G. micrantha
Binomial name
Gymnarrhena micrantha
Synonyms [1]
  • CryptadiaLindl. ex Endl. (1841)
  • FrankiaSteud. (1840)
  • Cryptadia euphratensisLindl. (1868)
  • Frankia schimperiSteud. (1840)
  • Gymnarrhena balansaeCoss. & Durieu ex Coss. & Kralik (1857)

Gymnarrhena is a deviant genus of plants in the family Asteraceae, with only one known species, Gymnarrhena micrantha. It is native to North Africa and the Middle East, as far east as Balochistan. Together with the very different Cavea tanguensis it constitutes the tribe Gymnarrheneae, and in the subfamily Gymnarrhenoideae. [2] [3]

Contents

Gymnarrhena is a small, flowering, winter annual with a rosette of simple, narrow leaves and flower heads cropped at its hart. It does not contain latex and does not carry spines. Gymnarrhena flowers in March and April. [4] One of the common names in Arabic is كَف الكَلْب meaning "dog's footprint", while in Hebrew it is called מוצנית קטנת-פרחים meaning "small chaff flower". [5] [6]

Description

Gymnarrhena micrantha is a dwarf annual herb of ½–2½ cm high, with all its leaves in rosette of up to 10 cm in diameter, and its flowers tucked away in the hart of this rosette, that is lacking latex, and does not have thorns. Two sources report twenty chromosomes (2n=20), but one other publication says eighteen (2n=18). [7] [8] [9]

Leaves

The leaves are simple and are arranged in a dense basal rosette. They are narrowly lanceolate to narrowly ovate in shape, more or less V-shaped in cross-section, lack leaf stalks and have a smooth surface. The tip is pointy or gradually narrowing. The leaf margins may carry some small, distanced teeth. [7]

Subterranean flower heads

The flower heads that develop underneath the leaves do not open and are self-pollinated. Each floret is fully enclosed in its involucral bracts, and the corolla shows very little development. The cypselas are relatively large and flattened, blackish in color, with ample hairs, and remain below the soil surface after the plant has died. Any pappus consists of somewhat scale-like bristles, hardly developed or is entirely absent. [7]

Aerial flower heads

The aerial flower heads are congested in the center of the leaf rosette, more or less arranged as a low cauliflower. Groups of florets are either functionally male or functionally female. The involucral bracts are overlapping in several series, papery, whitish and have a pointy tip. [7] The aerial flower heads have some semblance to a hedgehog and the hard, dry plants hurt the naked foot if stepped upon. [9]

Male florets

The functionally male florets occur in small groups and have very short individual stems, mostly in the centre of a larger cluster of female florets. The corollas are small, have (three or) four triangular lobes, greenish yellow and contain (three or) four stamens, carry yellowish or purplish anthers that are blunt on both ends and the filament is not extended beyond the anther. The fruit at the base of the male flower is much reduced and void, and pappus may consist of some irregular scales or be entirely abstent. Pollen is globe-shaped and has three sunken furrows (a type called tricolpate). These furrows have sharp ends and do not merge at the poles. The pollen has some unevenly distributed hollow spines, which are conical with a somewhat swollen base and a pointed tip, 1—2 μm high. [7]

Female florets and fruits

The functionally female florets are each stiffly enclosed by a large, cone-shaped green and white bract, and clustered with other one-flowered female flower heads, mostly surrounding groups of male florets. The inconspicuous corolla consists of whitish threads and surrounds the base of a whitish style with long arms which have rounded tips. The one-seeded indehiscent fruit (or cypsela) of the female florets are tiny, ovoid, with rows of stiff hairs on the ribs, and felty overall with long twin hairs, which have thin cell walls. The pappus on top consists of scales ending in a long drawn tip and with a row of hairs along the edges. [7]

Characters in common with Cavea

Gymnarrhena is most related to Cavea, but few morphological features would support this assignment, other than both having two types of flower heads and sharing a tendency towards dioecism. [3] Both also have basal leaf rosettes, stretched leaves, with few spaced teeth on the margin, and both lack spines and latex.

Differences with other Asteraceae

Gymnarrhena has aerial inflorescences that consist of many individual flower heads with disk florets which are either functionally male, with few florets each, or female with one floret only. This is a rare character combination, that is further known from the inflorescences of Gundelia . The latter however is a much larger, erect, thistle-like plant, which has latex and pentamerous florets. In Gymnarrhena the male florets (the only ones where a judgement can be made without enlargement) are (tri- or) tetramerous. The vast majority of Asteraceae have pentamerous florets, and several to many florets per flower head. Other asterids that have flower heads with only one floret are Corymbium , Hecastocleis shockleyi , Stifftia uniflora and Fulcaldea laurifolia , but these are pentamerous and hermaphrodite.

Taxonomy

specimens with and without aerial flowers Gymnarrhena micrantha003.jpg
specimens with and without aerial flowers

Gymnarrhena micrantha was first described by René Louiche Desfontaines in 1818. In 1857, a second species, G. balansae was distinguished by Ernest Cosson and Michel Charles Durieu de Maisonneuve, but it is doubtful this form from Algeria is sufficiently different. In 1868, the name Cryptadia euphratensis, with a description by John Lindley, was published, [10] but this name was later synonymised with G. micrantha.

The unusual morphology of Gymnarrhena has made it difficult for taxonomists to make a solid assignment. Bentham, Hoffmann, and Cronquist put this genus in the Inuleae tribe in its widest circumscription, while both Hoffmann and Cronquist note Gymnarrhena is similar to Geigeria . [11] [12] [13] In 1973, Peter Leins thought the pollen of Gymnarrhena too different from the Inuleae and proposed a position in the Cardueae (Cynareae) tribe. John J. Skvarla, Billie Lee Turner, and their colleagues in 1977 agreed that Gymnarrhena has some traits in common with the Cynareae but a pollen type that cannot be found in the Inuleae tribe. Kåre Bremer in his 1994 book Asteraceae: Cladistics & Classification included Gymnarrhena in the Cichorioideae, but without clarifying its position in this tribe. [7]

Modern classification

Gymnarrhena micrantha is now considered the sister group of Cavea tanguensis , who together constitute the tribe Gymnarrheneae and the subfamily Gymnarrhenoideae. [3] [7]

Phylogeny

Based on recent genetic analysis, it is now generally accepted that the Pertyoideae subfamily is sister to a clade that has as its basal member the Gymnarrhenoideae, and further consists of the Asteroideae, Corymbioideae and Cichorioideae. These three subfamilies share a deletion of nine base-pairs in the ndhF gene which is not present in Gymnarrhena micrantha. Current insights in the relationships of Gymnarrhena to the closest Asterid subfamilies are represented by the following tree. [3] [7]

subfamily Pertyoideae

subfamily Gymnarrhenoideae

Gymnarrhena micrantha

Cavea tanguensis

subfamily  Cichorioideae

Warionia saharae

all other Cichorioideae

subfamily Corymbioideae

subfamily Asteroideae

Evolutionary processes

The early production of few large seeds followed by production of many small seeds may have been the result of the variable and unpredictable growing season, fitting to a pioneer species. [7]

Etymology

The genus name Gymnarrhena may be a contraction of two Greek words, γυμνός (gymnos) meaning "naked" and ἄῤῥην (arrhēn), "male", while the species epithet micrantha is a contraction of the Greek words μικρός (mikrós), "small" and ἄνθος (ánthos), "flower".

Distribution and habitat

a herbarium specimen Gymnarrhena micrantha004.jpg
a herbarium specimen

Gymnarrhena is known from North Africa, such as Algeria and Egypt, the Middle-East, such as Sinai, Israel, Jordan, Siria, Kuwait, eastern Saudi Arabia, Qatar, the United Arab Emirates, Iraq, Iran and Pakistan (Balochistan). It grows on gravel plains, stony or rocky areas, in thin sandy deposits, alluvial plains and wadis. [7] [9] [14] [15] [16]

Ecology

Gymnarrhena is a dwarf herbaceous winter annual plant. It is said to be poisonous and animals appear to avoid it. It has aerial flowers in March, April and - when sufficient moisture is around - May. [14] Already after four leaves have grown, underground flowerheads develop that produce few larger cypselas, followed later on by many small cypselas from the aerial flower heads for as long as moisture is available. When the plant dies down in summer, the cypselas remain encased between the hardened bracts, presumably safe from harvester ants. After the first rain, which usually occurs the next winter, the bracts and pappus on the aerial flowerheads unfold, and the cypselas are dispersed by the wind, while many are gathered by ants. The cypselas in the underground flowerheads however germinate through the dead parts of the flowerhead, and remain protected against the ants. These seeds increase the chance that the plant continues its presence in a location that was favorable in the previous year. Aerial cypselas on average only weigh 5–6% of a subterranean fruit. After six days, seedlings of subterranean fruits are six times larger than those of aerial fruits and their survival rate is much higher. In very dry years, only subterranean fruits may develop and aerial florets may be entirely absent. [7]

Gymnarrhena is one of few species that grows where the sand has blown out from under tar tracks in Kuwait after the Gulf war, a strong confirmation of its ability to colonize disturbed habitat quickly. At Khirbet Faynan, in the southern Jordanian desert, Gymnarrhena grows on slag piles containing copper and lead and accumulates these heavy metals. [8]

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The family Asteraceae, with the original name Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae were first described in the year 1740. The number of species in Asteraceae is rivaled only by the Orchidaceae, and which is the larger family is unclear as the quantity of extant species in each family is unknown.

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

Gundelia or tumble thistle is a low to high (20–100 cm) thistle-like perennial herbaceous plant with latex, spiny compound inflorescences, reminiscent of teasles and eryngos, that contain cream, yellow, greenish, pink, purple or redish-purple disk florets. It is assigned to the family Asteraceae. Flowers can be found from February to May. The stems of this plant dry-out when the seeds are ripe and break free from the underground root, and are then blown away like a tumbleweed, thus spreading the seeds effectively over large areas with little standing vegetation. This plant is native to the eastern Mediterranean and the Middle-East. Opinions differ about the number of species in Gundelia. Sometimes the genus is regarded monotypic, Gundelia tournefortii being a species with a large variability, but other authors distinguish up to nine species, differing in floret color and pubescence. Young stems are cooked and eaten in the Middle-East and are said to taste like a combination of artichoke and asparagus.

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

Scolymus is a genus of annual, biennial or perennial, herbaceous plants that is assigned to the family Asteraceae, and can be found in Macaronesia, around the Mediterranean, and in the Middle East. All species are spiny, thistle-like in appearance, with flowerheads that consist of yellow ligulate florets, and canals that contain latex. It is sometimes called golden thistle or oyster thistle, and is known as سكوليمس (skwlyms) in Arabic and scolyme in French.

<i>Scolymus maculatus</i> Species of flowering plant

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<i>Catananche caerulea</i> Species of flowering plant

Catananche caerulea, or Cupid's dart, is a greyish green perennial herbaceous plant with a basal leaf rosette and conspicuous blue-purple or sometimes white flowerheads, belonging to the daisy family. It is a popular garden plant and is often used in dried flower arrangements.

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

Hecastocleis is a genus of low thorny shrubs with stiff branches, assigned to the daisy family. At the tip of each of the branches, inflorescences are subtended by oval, thorny, whitish to greenish bracts that enclose several flower heads which each contain only one pinkish bud, opening into a white corolla. It contains but one species, Hecastocleis shockleyi, the only representative of the tribe Hecastocleideae, and of the subfamily Hecastocleidoideae. Its vernacular name is prickleleaf. It is confined to the southwestern United States.

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

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

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

Catamixis is a genus assigned to the family Asteraceae, with only one known species, Catamixis baccharoides, a low to medium height, ¾—1¾ m, shrub. It is native to a very small area of western Nepal and northern India in the Himalayas. It has approximately spoon-shaped, leathery leaves with distanced rounded teeth alternately set along straight, shyly branching stems, and carries many flower heads of about 1 cm, with a few creamy white florets, sometimes with a hint of violet, in corymbs at the end of the branches. Flowers and fruits can be found between March and May. Its vernacular name in Hindi is विषपत्री (vishpatri) or विश्पत्र (vishpatra).

Cavea is a low perennial herbaceous plant that is assigned to the family Asteraceae. Cavea tanguensis is currently the only species assigned to this genus. It has a basal rosette of entire, slightly leathery leaves, and stems of 5–25 cm high, topped by bowl-shaped flower heads with many slender florets with long pappus and purplish corollas. The vernacular name in Chinese is 葶菊. It grows high in the mountains of China (Sichuan), Tibet, India (Sikkim), and Bhutan, and flowers in July and August.

<i>Corymbium</i> Genus of perennial plants from South Africa

Corymbium is a genus of flowering plants in the daisy family comprising nine species. It is the only genus in the subfamily Corymbioideae and the tribe Corymbieae. The species have leaves with parallel veins, strongly reminiscent of monocots, in a rosette and compounded inflorescences may be compact or loosely composed racemes, panicles or corymbs. Remarkable for species in the daisy family, each flower head contains just one, bisexual, mauve, pink or white disc floret within a sheath consisting of just two large involucral bracts. The species are all endemic to the Cape Floristic Region of South Africa, where they are known as plampers.

<span class="mw-page-title-main">Stifftioideae</span> Subfamily of flowering plants

The Stifftioideae are a subfamily of the family Asteraceae family of flowering plants. It comprises a single tribe, Stifftieae, of ten genera.

<i>Gorteria diffusa</i> Annual plant in the daisy family from South Africa

Gorteria diffusa is a highly variable, small annual herbaceous plant or rarely a shrublet that is assigned to the daisy family. Like in almost all Asteraceae, the individual flowers are 5-merous, small and clustered in typical heads, and are surrounded by an involucre, consisting of in this case several whorls of bracts, which are merged at their base. In G. diffusa, the centre of the head is taken by relatively few male and bisexual yellow to orange disc florets, and is surrounded by one complete whorl of 5–14 infertile cream to dark orange ray florets, sometimes with a few ray florets nearer to the centre. None, some or all of them may have darker spots at their base. The fruits remain attached to their common base when ripe, and it is the entire head that breaks free from the plant. One or few seeds germinate inside the flower head which can be found at the foot of plants during their first year. The species flowers between August and October. It is called beetle daisy in English and katoog in Afrikaans. It can be found in Namibia and South Africa.

<i>Hymenonema laconicum</i> Species of plant in the family Asteraceae

Hymenonema laconicum is a species of herbaceous perennial plant in the Asteraceae family. It is small to average height, with a rosette of greyish pinnately segmented leaves, and little branching solid stems carrying one to three heads of orange or yolk yellow ray-flowers, with a purple anther tube, and scaly pappus. The species is an endemic of the central and south-eastern Peloponnesos, and flowers in May and June.

<i>Scolymus grandiflorus</i> Species of flowering plant

Scolymus grandiflorus is a spiny annual or biennial plant in the family Asteraceae, native to the Mediterranean region. With up to 75 cm high stems, it is the smallest of the species of Scolymus. Its stems are lined with uninterrupted spiny wings. It also has the largest flowerheads in the genus, of approximately 5 cm wide. It has yellow, sometimes yolk-yellow ligulate florets. Its vernacular name in Maltese is xewk isfar kbir, meaning "large yellow fin", cardogna maggiore in Italian, scoddi on Sicily, and scolyme à grandes fleurs in French.

Catananche lutea, is a woolly annual plant, in the family Asteraceae, with most leaves in a basal rosette, and some smaller leaves on the stems at the base of the branches. Seated horizontal flowerheads develop early on under the rosette leaves. Later, not or sparingly branching erect stems grow to 8–40 cm high, carrying solitary flowerheads at their tips with a papery involucre whitish to beige, reaching beyond the yellow ligulate florets. Flowers are present between April and June. This plant is unique for the five different types of seed it develops, few larger seeds from the basal flowerheads, which remain in the soil, and smaller seeds from the flowerheads above ground that may be spread by the wind or remain in the flowerhead when it breaks from the dead plant. This phenomenon is known as amphicarpy. The seeds germinate immediately, but in one type, germination is postponed. It naturally occurs around the Mediterranean. Sources in English sometimes refer to this species as yellow succory.

<span class="mw-page-title-main">Gymnarrhenoideae</span> Subfamily of flowering plants

Gymnarrhenoideae is a subfamily within the family Asteraceae, with only one tribe, the Gymnarrheneae. Two very different species have been assigned to it, Gymnarrhena micrantha, a winter annual from the deserts of North-Africa and the Middle-East, and Cavea tanguensis, a perennial herb that grows on scree near streams and glaciers in the Eastern Himalayas. These species have very little in common, other than having two types of flower heads and sharing a tendency towards dioecism. Both also have basal leaf rosettes, stretched leaves, with few spaced teeth on the margin, and both lack spines and latex.

Famatinanthus is a genus in the family Asteraceae that was described in 2014 and has been assigned to its own tribe Famatinantheae and subfamily Famatinanthoideae. It contains only one known species, F. decussatus, a small shrub of ½—1¾ m high that is an endemic of the Andes of north-western Argentina, with small, entire, oppositely set leaves and flowerheads containing about ten cream-colored, ray and disk florets, with backward coiled lobes. It is locally known as sacansa. For more than 100 years, the species was known to science only from the type collection. It was described in 1885 and originally assigned to the genus Aphyllocladus.

<i>Felicia echinata</i> Shrublet in the daisy family from South Africa

Felicia echinata, commonly known as the dune daisy or prickly felicia, is a species of shrub native to South Africa belonging to the daisy family. It grows to 1 m (3.3 ft) high and bears blue-purple flower heads with yellow central discs. In the wild, it flowers April to October.

<i>Felicia</i> (plant) Genus of shrublets, perennials and annuals in the daisy family

Felicia is a genus of small shrubs, perennial or annual herbaceous plants, with 85 known species, that is assigned to the daisy family. Like in almost all Asteraceae, the individual flowers are 5-merous, small and clustered in typical heads, and which are surrounded by an involucre of, in this case 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 and are not individually subtended by a bract. Most species occur in the Cape Floristic Region, which is most 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 and 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.

References

  1. "Gymnarrhena micrantha Desf". Plants of the World Online. Retrieved 18 March 2024.
  2. Panero, Jose L.; Funk, Vicki A. (2009), "New tribes in Asteraceae", Phytologia, 91 (3): 568–570
  3. 1 2 3 4 Zhi-Xi Fu; Bo-Han Jiao; Bao Nie; Tiangang Gao (2016). "A comprehensive generic‐level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae". Journal of Systematics and Evolution. 54 (4): 416–437. doi: 10.1111/jse.12216 . S2CID   89115861 . Retrieved 2016-01-15.
  4. "Gymnarrhena micrantha | Compositae". Archived from the original on 2010-06-14. Retrieved 2012-05-09.
  5. "Gymnarrhena micrantha". Euromed Plantbase. Retrieved 2016-12-31.
  6. "Gymnarrhena micrantha". Wild Flowers of Israel. Retrieved 2016-12-29.
  7. 1 2 3 4 5 6 7 8 9 10 11 12 Funk, Vicki A.; Fragman-Sapir, Ori (2009). "22. Gymnarrheneae (Gymnarrhenoideae)" (PDF). In V.A. Funk; A. Susanna; T. Stuessy; R. Bayer (eds.). Systematics, Evolution, and Biogeography of Compositae. Vienna: International Association for Plant Taxonomy. pp. 327–332. Retrieved 2016-12-27.
  8. 1 2 Yitzak Guttermann (1989). "Gymnarrhena micrantha". In Halevy, Abraham H. (ed.). Handbook of Flowering. Vol. 6. CRC Press. pp. 356–358. ISBN   9780849339165.
  9. 1 2 3 "الكفنة [Gymnarrhena micrantha]". موقع السرحان [Sarhan site]. Retrieved 2016-12-31.
  10. Chesney, Francis Rawdon (1868). Narrative of the Euphrates Expedition carried on by Order of the British Government during the years 1835-1837. London: Longmans, Green & Co. cited on Vicki A. Funk. "Cryptadia euphratensis Lindley in Chesney, first listing of this species". compositae. Retrieved 2016-12-28.
  11. Bentham, George (1873). "Notes on the Classification, History, and Geographical Distribution of Compositæ". Botanical Journal of the Linnean Society. 13 (70–72): 335–577. doi:10.1111/j.1095-8339.1873.tb02575.x.
  12. Hoffmann, O. (1890–1894). "Compositae". In H.G.A. Engler; K.A.E. Prantl (eds.). Die natürlichen Pflanzenfamilien. Vol. 4. Leipzig, Germany: Wilhelm Engelmann Verlag. pp. 87–387.
  13. Cronquist, A. (1955). "Phylogeny and taxonomy of the Compositae". American Midland Naturalist . 53 (2): 478–511. doi:10.2307/2422084. JSTOR   2422084.
  14. 1 2 "Camel's Eye". Qatar e-Nature. Retrieved 2016-12-28.
  15. Petr Pokorny; Adela Pokorna. "Gymnarrhena micrantha". Common Plants of the Western Deserts of Egypt.
  16. Věra Svobodová (2010). "Gymnarrhena micrantha Desf". botany.cz. Retrieved 2016-12-31.