Physalis

Cape gooseberry (Physalis peruviana ) Flower and shell of the immature fruit

The Physalis (Physalis ), also called Jews cherries, a genus of the family Solanaceae are (Solanaceae ). Most of the approximately 75 to 90 species are native to the Americas, while in Europe only the Chinese lantern plant (Physalis alkekengi ) occurs. Remarkable are especially the fruits that completely magnification of one to lantern -shaped calyx are enclosed. Some species of the genus, for example, often referred to only as Cape gooseberry Physalis, are cultivated as fruit and vegetables, others used for decorative purposes.

• 3.1 Botanical history and naming
• 3.2 External systematics
• 3.3 Internal systematics
• 3.4 phylogenetics

Description

Vegetative characteristics

Are jujubes year or (more rarely) perennial herbaceous plants, erect, slightly creeping grow prostrate bushy or in exceptional cases. Depending on the plants from 0.2 to 0.6 m ( often 0.1 to 0.7 m) can be large. Probably the hollow in all kinds Sprossachse sympodial branches. Individual species of the genus have a hair on the leaves, stem axis and partly also in the interior of the flower. This hair will often vary widely even within a species. The trichomes are straight or curved, short or long, the shapes can be simple, forked, branched or capitula -shaped with one or multicellular heads, in exceptional cases, the base of trichomes consist of several cells. For example, while P. Alkekengi is almost insensitive to frost and also survived harsh winters, there are other species that can not tolerate frost.

The perennial representatives usually form thick, woody beet roots, are more common in the genus, however, more or less horizontally to find growing brood buds forming roots that are deeply rooted in the earth and lead to clusters of rhizomes. The entire underground part of the plant can extend over an area with a diameter of several meters.

The alternate or often in pairs standing leaves are simply built, entire to coarsely toothed or lobed, they can elliptical, ovoid, elongated ovate, spatulate or rarely be linear. They are usually relatively thin, only in exceptional cases thick. The blade tip is blunted, scharfspitzig or pointed. The leaf base is asymmetrical, tapering at the stem and decurrent or heart-shaped. The lamina has a length of 5-11 cm (1.5 inches ) and a width of 4-6 cm (1 inch ). The petiole is 2-4 cm ( 0.7 to 6 cm ) long.

Inflorescences and flowers

The axillary flowers are single or in groups of two to seven flowers, usually they are hanging down and stand at 1.5-5 mm short or 11-35 mm (50 mm) long flower stems that occasionally arise from a very short peduncle. The bell-shaped and radiärsymmetrische calyx has five sepals or deeper divided into five parts, in which the separation is never more than two-thirds the length of the calyx. These parts are triangular, semi- ovate or long and linear shaped, 4-10 mm (2-14 mm ) long and usually invaginated at the base. The crown is radial symmetry, wheel-shaped to bell-like wheel-shaped, in exceptional cases ( Physalis Solanacea ) and urn -shaped. The diameter is 10-20 mm ( 5-35 mm). The coronary band is usually yellow, rarely white, in exceptional cases ( Physalis Solanacea ) purple to violet. The crown is set with five color or violet, violet - brown or greenish points, these are normal, confluent or consisting of several smaller points. The edge is provided with five short teeth, lobes or lobes or five long sections, which make up less than half the length of the crown. Inside the corolla tube usually is a ring of trichomes, which can have different shape and proportions. It is continuous or interrupted, so that the individual sectors are alternating with the stamens.

The five stamens are 1.5-3.5 mm ( 0.75 to 4.6 mm) long, and within a flower of the same length or only slightly different lengths. They are yellow or rarely blue. The stamens are fused with a broadened base at the bottom of the crown and fixed to the base or the back of the base of the dust bag. The counters of the anthers are only available in a small directed towards the flower center area free from each other, the connecting tissue between the counters is wide. The stamens are usually as long as or longer than the anthers, rarely shorter, they are occupied glabrous or with a few trichomes. The pollen grains are trizonocolpat (the three germ folds lie on pollen equator) and have a diameter of 25-29 microns, making them among the medium-sized pollen grains. The pollen grain wall ( exine ) is smooth, reticulate, filled with single free spines or rough.

The ovary is zweifächrig, the stylus is almost constantly medium and hollow, the scar is short saddle-shaped or shaped capitula - pressed, moist, fertile, the surface is covered with unicellular shallow to medium-sized papillae. To the ovary are annular nectaries.

Fruit and seeds

After fertilization of the flower petals, the first to be dropped. Subsequently, the five sepals enlarge with increasing fruit maturity, so that they join and form almost a lantern -shaped chalice to the developing berry. At maturity, these cover discolored yellow to bright orange. The enlarged sepals have a length of 6-15 mm ( 4-25 mm).

The berry itself is flattened globose to slightly, 4-7 or 10-16 mm (20 mm) in size. Depending on the ripe fruits are green to yellow or tangerine, in part, they are also covered in red or purple. They include a large number ( 100 to 180, in some species but only 5 to 16 ) of small, lens-shaped, light yellow - brown seeds. These have a size of 1.5-2.4 mm ( 1.2-2.8 mm) and are surrounded by parenchyma cells, the origin of the placenta and the pericarp. The surface of the seed is like a net, like a honeycomb or wrinkled - warty. The embryo is almost rotated or rotated, the cotyledons shorter than the rest of the embryo the endosperm is abundant.

Dissemination and locations

Most species of the genus originate from the area between Central America and the southern United States, only the origin of the Chinese lantern plant (Physalis alkekengi ) is made in China or possibly also suspected in Europe. Most species grow in Mexico, of which two thirds are endemic. Only twelve species are also found in South America. Many species have spread beyond the original distribution areas also, so Physalis example, Alkekengi also found in the northeastern United States and Physalis philadelphica was introduced among others in Turkey.

They grow between sea level and positions 2300-2700 m in forests, as ruderal, or disturbed sites with secondary vegetation.

System

Botanical history and naming

The use of the name Physalis is already occupied by works of Dioscorides. The term derives from the Greek Physa ( "bubble "). The name Jews cherry is derived from the shape of the surrounding lanterns ( the calyx ), which is similar in shape and color according to the different dress codes prescribed for Jews hats, which in turn can be derived from the Phrygian cap. These hats were common since the early Middle Ages.

A first known graphical representation of occurring in Europe Lampionblume (Physalis alkekengi ) comes from the julianae Aniciae Codex ( Vienna Dioscorides ), which is dated at about 512. In later medieval herbals are various pictorial representations of this kind to be found. A first picture of a coming from America type is probably the drawing of Physalis philadelphica from Francisco Hernandez 'Nova plantarum animalum et Mineralium mexicanorum historia of 1651st Before the introduction of the binary nomenclature by Carolus Linnaeus different names were as Solanum vesicarium vulgatius repens, fructu et vesica rubra (Robert Morison ), Alkekengi barbadense nanum, alliariae folio (Johann Jacob Dillen ) or Alkekengi virginianum fructo luteo (Louis Feuillée ) used.

Linnaeus described in 1753 in his work Species Plantarum the genus Physalis with a total of nine species, of which five perennial and four annual species. Soon after, a number of other species have been described, so that for example in 1817 in the work Systema Vegetabilium of Johann Jacob Roman and Joseph August Schultes total of 37 species have been described. Often the species descriptions were, however, placed carelessly, so that Christian Gottfried Daniel Nees found by Esenbeck 1831 that the genus Physalis a " remarkable example of synonyms Pilings " since you " re the species already designated designated time and time again, without to give an account where now want the old ways, who had escaped the eyes are hidden. " He reduced the known species of the genus to only 17 established the three of him, but not named sections in 1837 by Don George named. In Physalodendron were the woody species in Eurostorhiza persevering, rhizomatic types and classified the annual species in Epeteiorhiza. A fourth section introduced by him Anomalae contained two species with tubular calyces. His conjecture that these types probably constitute another genus, should prove to be correct later - today these species are assigned to the genus Deprea.

Other arrangements of the genus with different classifications were made in 1852 by Michel Félix Dunal, which recognized only two of Don's sections, and in 1896 by Per Axel Rydberg, who did not use any of the sections described above and divided the genus into three separate sections. Two of these sections were monotypic, the third ( Euphysalis, today a section Physalis ) was again divided into seven groups of species.

In the 20th century, initially employed Margaret Y. Menzel (1951 ), comprising the genus, led among others by crossing experiments with 28 species and provided for these karyological data. The previously established by Rydberg species groups were collected by them to the sections. A summary of all treatments carried out up to that of the genus was published in 1989 by Radovan Hendrych, last major taxonomic considerations were made in 1994 and 1999 by Mahinda Martínez. Up to the outsourcing of the genus Quincula this classification was also confirmed by phylogenetic analyzes.

External system

While an enlarged fruit cup can be observed within the entire family of Solanaceae, is a bloating, as can be observed in the jujubes, very rare. Thus, it is usually very easy to distinguish jujubes from other members of the nightshade family. Through morphological similarities, especially in terms of the enlarged calyx, a variety of species have been described as physaloid. In many of these physaloiden genres, such as Archiphysalis, Athenaea, Deprea, Exodeconus, Jaltomata, Larnax, Nicandra, Physalisatrum or Saracha no close relationship to Physalis could be determined by cpDNA analysis.

Discrepancies there are associated primarily with the classification under the subtribe Physalinae, especially Margaranthus solanaceous (or physalis Solanacea ) and Quincula lobata (or Physalis lobata ) are traded by different authors either as a separate species, each with a species or genus of jujubes.

The external systematics of the genus Physalis is as follows (quoted from Olmstead et al. )

• Family: Solanaceae [ ... ]
• Tribus: Physaleae [ ... ]
• Subtribe: Physalinae Genus: Brachistus
• Genus: Leucophysalis
• Genus: Margaranthus ( genus status to be confirmed)
• Genus: Oryctes
• Genus: Quincula ( genus status to be confirmed)
• Genus: Physalis
• Genus: Tzeltalia (. Away by Olmstead et al in the genus Physalis, but mapped to morphological reasons of the subtribe )
• Genus: Witheringia

Another proposal for the assignment of different genres to subtribe Physalinae presented Armando Hunziker 2000, he took on only the genera Physalis, Quincula, Leucophysalis and Chamaesaracha.

Internal system

The genus of jujubes includes an estimated 75 to 90 species. As with many genera of the family Solanaceae is a generally recognized system has not yet been established and is complemented by new research results again and again.

In the last revision of the systematics of the genus Physalis it was divided in 1999 by Martinez in four subgenera with twelve sections. Among other things, which was previously considered independently managed genera Quincula and Margaranthus were subordinated to the jujubes.

Accordingly, the internal systematics of this genus subdivided as follows: ( expanded gradually, with changes marked *)

• Physalis (Physalis )

• Physalis subgenus

• Lampionblume (Physalis alkekengi L.)

• Subgenus Physalodendron

• Physalis arborescens L.
• Physalis melanocystis Bitter

• Subgenus Quincula

• Subgenus Rydbergis

• Section Angulatae

• Physalis acutifolia ( Miers ) Sandwith
• Physalis ampla waterf.
• Physalis angulata L.
• Physalis carnosa Standley
• Physalis crassifolia Benth.
• Physalis lagascae Roem. & Schult.
• Physalis microcarpa Urb. & Eckman
• Tomatillo (Physalis philadelphica Lam. )
• Physalis Solanacea ( Schltdl. ) axelius ( syn. Margaranthus solanaceous Schltdl. )
• Physalis sulphurea ( Fernald ) waterf.

• Section Campanulae

• Physalis campanula Standlschmaus. & Steyerm.
• Physalis glutinosa Schltdl.

• Section Coztomatae

• Physalis aggregata waterf.
• Physalis angustior waterf.
• Physalis chenopodifolia Lam.
• Physalis coztomatl Dunal
• Physalis greenmanii waterf.
• Physalis hintonii waterf.
• Physalis Lassa Standley & Steyerm. *
• Physalis lignescens waterf. *
• Physalis longiloba Vargas, M. Martínez & Dávila *
• Physalis longipedicellata waterf.
• Physalis mcvaughii waterf. *
• Physalis orizabae Dunal *
• Physalis pennellii waterf.
• Physalis philippiensis Fernald
• Physalis pringlei Greenm.
• Physalis sancti - Josephi Dunal
• Physalis subrepens waterf.
• Physalis waterfallii Vargas, M. Martínez & Dávila *

• Section Epeteiorhiza

• Physalis angustiphysa waterf.
• Physalis cordata Mill
• Physalis grisea ( Waterf. ) M. Martínez
• Physalis ignota Britton
• Physalis latiphysa waterf.
• Physalis leptophylla B.L. Rob. & Greenm.
• Physalis minuta Griggs
• Physalis missouriensis Mack. & Bush
• Physalis neomexicana Rydb.
• Physalis nicandroides Schltdl.
• Physalis patula Mill
• Physalis porrecta waterf.
• Physalis pruinosa L.
• Physalis pubescens L.
• Physalis tamayoi Vargas, M. Martínez & Dávila *

• Section lanceolate

• Physalis caudella Standlschmaus.
• Physalis fendleri A. Gray
• Physalis gracilis Miers
• Physalis hastatula waterf.
• Physalis hederifolia A. Gray
• Physalis heterophylla Nees
• Physalis ingrata Standley
• Physalis lanceolata Michx.
• Physalis longifolia Nutt.
• Physalis muelleri waterf.
• Cape gooseberry (Physalis peruviana L.)
• Physalis pumila Nutt.
• Physalis queretaroensis M. Martinez & L. Hernandez *
• Physalis sordida Fernald
• Physalis virginiana Mill
• Physalis volubilis waterf.

• Section Rydbergae

• Physalis minimaculata waterf.
• Physalis rydbergii Fernald

• Section Tehuacanae

• Physalis tehuacanensis waterf.

• Section Viscosa

• Physalis angustifolia Nutt.
• Physalis cinerascens A. P. Hitchcock
• Physalis mollis Nutt.
• Physalis vestita waterf.
• Physalis viscosa L.
• Physalis walteri Nutt.

After the revision of Martínez ( 1999) Physalis amphitricha, Physalis calidaria, P. and P. parvianthera microphysa were excluded from the genus. The first two species were later assigned to the genus Tzeltalia, for the other two types is an assignment to other genera pending. Species with an uncertain assignment within the Physalis Physalis are jalicensis waterf. and Physalis Lassa Standley & Steyerm. (both suspected in section Coztomatae ) and Physalis lignescens (possibly synonymous with P. pubescens). In the Flora de Jalisco (2003 ), which ' is based on the revision Martínez, Physalis Lassa was placed in the section Coztomatae and Physalis lignescens, also recognized as a distinct species in section Coztomatae. The type Physalis carpenteri was until 2012 the only species of the section Carpenterianae, but since no close relationship was found for the remaining species of the genus, it was moved by Maggie Whitson in a separate genus Calliphysalis.

Phylogenetics

2005 were carried out by Whitson phylogenetic studies to confirm this division. Only the subgenus Rydbergis has - proved by these investigations as monophyletic, the classification of the genus Physalis in Margaranthus was confirmed by inclusion in this subgenus - after excluding the section Carpenterianae. Most species of this subgenus are characterized by herbaceous growth, separately growing flowers, ungelappte, yellow petals and a greatly enlarged fruit cup. Species that do not fall into this subgenus are characterized by different morphological features, such as multiple flowers per shoot nodes, lobed purple or white petals or an atypical fruit cup magnification. The genus Physalis in previously approved scope has proved to be paraphyletic, a spin-off the kind Physalis lobata in a monotypic genus Quincula was supported by the results.

Section Carpenterianae (2012 described as a species Calliphysalis )

Physalis subgenus

Physalis microphysa

Quincula lobata

Subgenus Physalodendron

Chamaesaracha

Subgenus Rydbergis (without section Carpenterianae )

Cladogram simplified after

The eponymous ancestral species for the genus P. Alkekengi proved to be far away from most other species of the genus as kladistisch. To resolve the resulting taxonomic problems within the genus would either all species except P. Alkekengi a new genus associated with, or class, would have to be extended within the subtribe Physalinae so that all genera is currently located there a common genus Physalis be assigned.

Importance

Among the jujubes, there are a few species that are of economic importance. These include, for example, used as a fruit cape gooseberry (Physalis peruviana ) or the well-known, especially in Mexico and the southern United States and cultivated as a vegetable tomatillo (Physalis philadelphica ). While the tomatillo is almost unknown in German supermarkets, the Cape gooseberry is not least used because of their exceptional fruit shell as a decoration. It is grown primarily in Africa, South America, India and Java and exported all over the world. Other producing countries are Australia, Kenya, New Zealand, the USA and South of France. The main harvest season is from December to July. In Germany it is usually sold under the generic name Physalis. More rarely, the pineapple cherry or strawberry tomato is (Physalis pruinosa Physalis grisea and ) cultivated as a fruit. The fruit of most other species have an unpleasant aftertaste.

A use as medicine against urinary tract is known both from European herbals as well as American cultures. The scarlet berries originating from China or Europe Lampionblume (Physalis alkekengi ) are edible, the plant is because of its orange calyxes as cut and dried flower popular.