Nolana
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Nolana coelestis |
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Nolana is a genus fromthe nightshade family (Solanaceae). With 89 species it belongs to the genera nightshade ( Solanum , over 1000 species), Lycianthes (around 200 species), hammer bushes ( Cestrum , around 175 species), box thorn ( Lycium , around 83 species) and tobacco ( Nicotiana , around 75 species) some of the most biodiverse in the family. Their occurrence is limited to areas in Peru and Chile , one species is endemic to the Galápagos Islands , so it only occurs there. Mainly due to the five-fold ovary and the resulting aggregate fruits , which otherwise do not occur within the nightshade family, their taxonomic classification was controversial for a long time.
description
Vegetative characteristics
The Nolana species are low or prostrate annual or perennial herbaceous plants or woody and semi-woody shrubs . You have a beet root . Often the plants are more or less succulent . The stem grows upright, ascending, prostrate or lying on top, it is hairless to tomentose. Growth rings can be seen in the cross-section.
In phases of extreme drought, the plants almost completely stop growing. However, if they are evenly supplied with sufficient water, many of the rosette-forming species form an elongated central shoot that ends in another leaf rosette. In this way, the plants can spread like a mat over an area several meters in diameter.
The leaves are simple and without stipules , can be stalked or sessile. They form a rosette and / or are stem-like. They are alternate, but almost opposite to tufted. The leaf blades are entire, fleshy, dorsiventral , ovate to linear or spatulate, occasionally the leaf base runs down unevenly on the stem axes. The largest leaves have the kind Nolana rupicola with up to 15 cm long and 10 cm wide leaves. In some cases, the leaf blades are greatly reduced and only rounded and linear, then only 1 to 5 mm long and 1 mm in diameter. These reduced leaves are often densely covered with simple to star-shaped or branched trichomes , which sometimes form glandular tips. Occasionally, some of these peaks are pronounced salt glands.
Flowers and inflorescences
Some species of the genus bloom very young when the plant size is only a few centimeters. The most conspicuous flowers occur in the species complex around Nolana acuminata (corresponds to clade A, compare internal systematics ) with diameters of 3 to 5 cm. The flowers are typically solitary in the leaf axils; racemose inflorescences are rarely formed on ascending, modified branches , in which each flower arises from the axilla of a circular bract .
The five-fold, radial symmetry to zygomorphic flowers are bisexual. If the flowers are zygomorphic, both the flower envelope and the androeceum (the entirety of all stamens ) or just the androeceum can be asymmetrical. The five sepals are the same or unevenly shaped, fused together and regular to almost two-lipped. They are seldom reduced to an unlobed tube. The calyx remains on the fruit and encloses it. The crown is funnel-shaped, bell-shaped or tubular, more rarely also almost urn-shaped or saucer-shaped. The five fused petals are of the same shape and folded between the corolla lobes, regularly or almost double-lipped. They are blue, purple, lavender or white, the coronet is dark purple or white at the base and can have pale yellow, white or green stripes.
The five stamens are in front of the sepals. Three of the stamens have longer filaments than the other two stamens. The stamens are fused together at the base and are often hairy. The dust bags open via inwardly directed longitudinal slits. The pollen emerges from them in the form of individual, blue or white, triple-furrowed pollen grains.
The flower base is an upstanding, circular, notched or lobed disc from which nectar is secreted. The ovary consists of five intergrown carpels . The stylus is terminal in the subgenus Alona , in the subgenus Nolana it stands on the ovary bulge (gynobased). The scar is head-shaped and moist.
Fruits and seeds
The fruits of the Nolana are historically interpreted as derived berries . In the subgenus Alona , the carpels are usually combined into a five-chambered (but also three- to six-chambered) ovary. The placentation is basal-axial, the ovary develops into a dry fruit from five (or three to six) multi-seeded partial fruits. In the subgenus Nolana, however, the carpels are heavily lobed, with an ovule in each lobe . The ovary develops into a dry fruit, which consists of up to 30 mostly solitary partial fruits. There is a curved or screwed embryo in the seeds .
Other features
The base chromosome number is x = 12. In plant chemistry studies, hydroxyflavonols , quercetin and kaempferol were found in plants , substances that are also found in other nightshade plants. Furthermore, diterpenoids and sesquiterpenoids were found.
Distribution and locations
The species of the genus Nolana have their distribution areas between northern Peru and southern Chile , only Nolana galapagensis is endemic to the Galápagos Islands . Most species have a fairly small area , with Nolana adansonii , Nolana gracillima , Nolana lycioides and Nolana jaffuelii , only four species are native to a larger range between southern Peru and northern Chile. Two particularly species-rich centers can be distinguished: one in the southern Peruvian department of Arequipa , the other in northern Chile between Antofagasta and Atacama .
Most of the species are formative components of the so-called Loma Formations - extremely dry areas near the coast that reach up to an altitude of 1000 m. The formations extend between 5 and 30 ° south latitude over a length of 3500 km and are only occasionally interrupted by river valleys. In some places the areas are only 25 km wide. The vegetation of these formations is primarily dependent on the moisture that gets there through fog from the ocean. There is also a strong connection with the occurrence of the El Niño phenomenon . Due to the increased moisture that is present during these phases, the otherwise desert-like areas are transformed into densely vegetated landscapes. The greatest biodiversity of the genus has developed just a few kilometers from the Pacific coast at altitudes between 50 and 600 m. Only a few Nolana species also grow inland or at heights of up to 4000 m.
ecology
Site observations showed that the flowers are visited by a wide variety of pollinators, including beetles , butterflies , flies , ants, and wasps . The beetle genus Epicante feeds exclusively on Nolana crowns. Presumably, these beetles also act as pollinators. Various Thripidae were also observed in the flowers.
The propagation mechanisms of the seeds have not yet been fully investigated. Birds are believed to be involved in the spread, and fruit has also been found in rodent excreta.
Systematics
External system
Although Nolana has been excluded from the nightshade family (Solanaceae) by some authors due to its unusual flowers and fruits (e.g. Hunziker (2001)), relatively early molecular biological studies showed a close relationship to the goat's thorns ( Lycium ). In the systematics of the nightshade family , published in 2007 by Richard Olmstead and Lynn Bohs , summarizing various phylogenetic studies , the genus is classified in an unrated group called "Lyciina" alongside the buckthorn and the genus Sclerophylax . Together with the tribe Hyoscyameae and the genera Jaborosa and Latua , they form an equally rankless group called "Atropina", which is placed within the subfamily Solanoideae.
Internal system
Occasionally the genre is divided into the two sub- genres Nolana and Alona , especially older adaptations see independent genres in the groups. They differ mainly in the properties of the fruit. Phylogenetic studies of 63 of the 89 currently recognized species by Dillon et al. (2007) did not provide any clear support for this structure. A group of species within the Nolana , previously known in part as Sorema Lindl. were outsourced from the genus, could possibly receive the status of a sub-genus.
The phylogenetic studies by Dillon et al. showed that the genus itself is monophyletic and is in turn divided into three monophyletic groups, one of which consists of the species Nolana sessiliflora and is a basal sister taxon to the rest of the genus. The other two groups are described by Dillon et al. designated as clade I and II:
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A total of eight further monophyletic subclades labeled A to H could be identified within clades I and II, but their position within the clades could not be fully clarified. Nolana tarapacana could not be assigned to any clade within clade II . One examined individual of Nolana peruviana was unexpectedly placed in clade H, but others were expected to be placed in clade F.
The five subclades below clade I are made up of:
- Clade A
- Nolana acuminata , Nolana baccata , Nolana elegans , Nolana paradoxa , Nolana parviflora , Nolana pterocarpa , Nolana reichei , Nolana rupicola
- Clade B
- Nolana aticoana , Nolana chancoana , Nolana chapiensis , Nolana humifusa , Nolana inflata , Nolana laxa , Nolana lezamae , Nolana plicata , Nolana scaposa , Nolana spathulata , Nolana urubambae , Nolana weissiana
- Clade C
- Nolana carnosa , Nolana coelestis , Nolana filifolia , Nolana rostrata
- Clade E
- Nolana adansonii , Nolana arenicola , Nolana galapagensis
The three subclades below clade II are made up of:
- Clade F
- Nolana albescens , Nolana aplocaryoides , Nolana clivicola , Nolana diffusa , Nolana flaccida , Nolana lachimbensis , Nolana leptophylla , Nolana mollis , Nolana onoana , Nolana peruviana , Nolana ramosissima , Nolana salsoloides , Nolana sedifolia , Nolana sphaerophylla , Nolana villosa
- Clade G
- Nolana arequipensis , Nolana cerrateana , Nolana confinis , Nolana gayana , Nolana intonsa , Nolana johnstonii , Nolana lycioides , Nolana pallida , Nolana pilosa , Nolana thinophila , Nolana tomentella , Nolana volcanica ,
- Klade H
- Nolana crassulifolia , Nolana divaricata , Nolana incana , Nolana peruviana , Nolana werdermannii
The species that were not considered in the investigation were mostly endemic species known only from the type specimen , such as Nolana foliosa , Nolana insularis , Nolana pearcei , Nolana platyphylla , Nolana polymorpha and Nolana weberbaueri . Others have probably already died out due to the destruction of the site, such as Nolana minor .
In 2007, Michael Dillon and his colleagues described ten more species, some of which were also not part of the phylogenetic studies. The newly described species are Nolana aenigma , Nolana arequipensis , Nolana chancoana , Nolana chapiensis and Nolana lezamae from Peru, as well as Nolana dianae , Nolana reichei , Nolana onoana , Nolana lachimbensis and Nolana philippiana from Chile.
use
Some species of the genus have limited importance as ornamental plants. Nolana humifusa was introduced to Europe from Peru as early as 1760, seeds are occasionally available commercially, but no named varieties are known. The Chilean species Nolana paradoxa has been known from European gardens since around 1820 and is the best-known cultivated species, well-known varieties are for example 'Blue Bird' and 'Cliff Hanger Blue'.
Botanical history and etymology
The genus was founded in 1762 by Carl von Linné jr. first described. The name he chose for the genus is derived from the Latin nola , which means something like "little bell". A placement in an independent family Nolanaceae was proposed by Friedrich von Berchtold and Jan Svatopluk Presl in 1820 , but the systematic treatments by George Don (1838) and Michel Félix Dunal (1852) each subordinated the genus to the nightshade family (Solanaceae). George Bentham and Joseph Dalton Hooker placed the species in a tribe within the hoist plants (Convolvulaceae). The genus was also later placed in different positions, for example as a separate family Nolanaceae by Arthur John Cronquist (1981) and Armen Tachtadschjan (1980), or within the nightshade family as the subfamily Nolanoideae or as a tribe Nolaneae, for example by Rolf Martin Theodor Dahlgren (1980) , William D'Arcy (1979, 1991) and Robert Folger Thorne (1983). As early as 1992 Richard Olmstead and Jeffrey Palmer showed that Nolana is phylogenetically related to the goat thorns ( Lycium ); However, this view was rejected by some editors, including Armando Hunziker (2001). However, further molecular biological work confirmed this classification.
The internal system of the genre was also often understood very differently; the number of genera and species into which the now recognized scope of the genus was divided fluctuated greatly. Michael Dillon (2007) states that systematic treatments based only on examination of herbarium specimens recognize very few species; Researchers who also carried out investigations in the field usually recognized significantly more species. Dillon justifies this with the fact that many properties that enable species delimitation are not preserved in herbarium specimens. John Lindley (1844) recognized five genera ( Nolana , Alonia , Dolia , Sorema and Aplocarya ) with a total of 17 species. Michel Felix Dunal (1852), who recognized Nolana as a tribe, also divided them into five genera ( Nolana , Dolia , Alibrexia , Aplocarya and Bargemontia ) with 33 species. George Bentham and Joseph D. Hooker (1873) recognized four genera ( Nolana , Alona , Dolia and Bargemontia ) with a total of 27 species.
The first modern monographic treatment by Ivan Murray Johnston (1936) names two genera ( Nolana and Alona ) with 63 species. Aldo Mesa , who in 1981 initially classified only 18 species in a single genus and divided them into the subgenus Nolana and Alona , expanded his consideration of the genus and recognized 70 species in works from 1997 and 1998. Extensive studies and field work, in which new species were also described, currently distinguish 89 species.
swell
- ↑ a b c d e Michael O. Dillon: Nolanaceae . In: K. Kubitzki (Ed.) The Families and Genera of Vascular Plants . Volume ??. Springer Verlag, Berlin 200x. Pre-published manuscript from www.sacha.org .
- ↑ a b Michael O. Dillon, Gina Arancio and Federico Luebert: Five new species of Nolana (Solanaceae-Nolaneae) from Chile . In: Arnaldoa , Volume 14, Number 2, 2007. pp. 191-212.
- ↑ Sandra Knapp: Tobacco to tomatoes: a phylogenetic perspective on fruit diversity in the Solanaceae. In: Journal of Experimental Botany , Volume 53, Number 377, October 2002. pp. 2001-2022. doi : 10.1093 / jxb / erf068
- ↑ Michael O. Dillon: The Solanaceae Of The Lomas Formations of Costal Peru and Chile (PDF; 11.2 MB). In: V. Hollowell, T. Keating, W. Lewis, T. Croat (Eds.): Solanaceae: William G. D'Arcy Memorial , Monographs in Systematic Botany from the Missouri Botanical Garden, 2004. pp. 131-156.
- ↑ a b c Rosanna Freyre, Amy C. Douglas, Michael O. Dillon: Artificial Hybridizations in Five Species of Chilean Nolana (Solanaceae) (PDF; 1.7 MB). In: HortScience , Volume 40, Heft 3, 2005. pp. 532-536.
- ↑ a b c d e f g Michael O. Dillon et al .: Phylogeny of Nolana (Nolaneae, Solanoideae, Solanaceae) as inferred from granule-bound starch synthase I (GBSSI) sequences . In: Taxon , Volume 56, Number 4, November 2007. pp. 1000-1011.
- ^ A b Armando T. Hunziker: The Genera of Solanaceae. ARG Gantner Verlag KG, Ruggell, Liechtenstein 2001. ISBN 3-904144-77-4 .
- ↑ Richard G. Olmstead and Jeffrey D. Palmer: A chloroplast DNA phylogeny of the Solanaceae: Subfamilial relationships and character evolution (PDF; 2.0 MB). In: Annals of the Missouri Botanical Garden , Volume 89, 1992. pp. 346-360.
- ^ Richard G. Olmstead and Lynn Bohs: A Summary of Molecular Systematic Research in Solanaceae: 1982-2006 . In: DM Spooner et al. (Ed.): Solanaceae VI: Genomics Meets Biodiversity , ISHS Acta Horticulturae 745, June 2007. ISBN 978-90-6605-427-1 . Pp. 255-268.
- ^ J. Francis Macbride: Nolanaceae . In: Flora of Peru , Field Museum of History, Botany Series, Volume XIII, Part V, Number 2, 1960. pp. 829-854.
- ↑ Michael O. Dillon, Segundo Leiva Gonzáles and Victor Quipuscoa Silvestre: Five new species of Nolana (Solanaceae-Nolaneae) from Peru and notes on the classification of additional taxa . In: Arnaldoa , Volume 14, Number 2, 2007. pp. 171-190.
Web links
- Nolaneae information from Michael O. Dillon (English)
- Nolana - entry in the W3Tropicos database (English)
