flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in the subfamily
Betoideae
The Betoideae are a small subfamily of the flowering plant amaranth family, Amaranthaceae ''sensu lato'' (or in Chenopodiaceae ''sensu stricto''). Commonly known members include beet, sugar beet, chard, and mangelwurzel, which all are cultivars o ...
of the family
Amaranthaceae
Amaranthaceae is a family of flowering plants commonly known as the amaranth family, in reference to its type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making it ...
. These are mostly procumbent herbs occurring in the Western
Mediterranean region
In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...
and
Macaronesia
Macaronesia (Portuguese: ''Macaronésia,'' Spanish: ''Macaronesia'') is a collection of four volcanic archipelagos in the North Atlantic, off the coasts of Africa and Europe. Each archipelago is made up of a number of List of islands in the At ...
, with some isolated occurrences in
North Africa
North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
and at the
Horn of Africa
The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
. They are interesting as
crop wild relative
A crop wild relative (CWR) is a wild plant closely related to a domesticated plant. It may be a wild ancestor of the domesticated (cultivated) plant or another closely related taxon.
Overview
The wild relatives of crop plants constitute an i ...
s of
sugar beet
A sugar beet is a plant whose root contains a high concentration of sucrose and which is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together wi ...
.
Description
''Patellifolia'' are
annual
Annual may refer to:
*Annual publication, periodical publications appearing regularly once per year
** Yearbook
** Literary annual
*Annual plant
*Annual report
*Annual giving
*Annual, Morocco, a settlement in northeastern Morocco
*Annuals (band), ...
or perennial herbs, growing erect or often procumbent. The alternate
leaves
A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
have a petiole, their leaf blade is heart-shaped or hastate.
The spike-like
inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s consist of glomerules of one to three flowers sitting in the axils of leaf-like
bract
In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
s. The free
flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are hermaphrodite. The
perianth
The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...
consists of five herbaceous, slightly keeled
tepal
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
lobes which are connate at base. There are five
stamen
The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filame ...
s opposite to the tepals, their filaments are fused at base forming a disc. The ovary is semi-inferior with 2 stigmas.
In fruit, the ovary is partly immersed in the enlarged base of the perianth. The tepal lobes are not modified and appressed to the fruit. The solitary fruits are nearly globular capsules, they fall down when ripe and open with a circumscissile lid. The vertical seed contains an annular embryo and copious perisperm (feeding tissue).
Distribution and habitat
''Patellifolia'' is mainly distributed in the Western
Mediterranean region
In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...
(
Spain
, image_flag = Bandera de España.svg
, image_coat = Escudo de España (mazonado).svg
, national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond")
, national_anthem = (English: "Royal March")
, i ...
,
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
),
Macaronesia
Macaronesia (Portuguese: ''Macaronésia,'' Spanish: ''Macaronesia'') is a collection of four volcanic archipelagos in the North Atlantic, off the coasts of Africa and Europe. Each archipelago is made up of a number of List of islands in the At ...
Islands. There are also some isolated occurrences in North Africa (
Hoggar Mountains
The Hoggar Mountains ( ar, جبال هقار, Berber: ''idurar n Ahaggar'') are a highland region in the central Sahara in southern Algeria, along the Tropic of Cancer. The mountains cover an area of approximately 550,000 km.
Geography
This ...
and
Tassili n'Ajjer
Tassili n'Ajjer ( Berber: ''Tassili n Ajjer'', ar, طاسيلي ناجر; "Plateau of rivers") is a national park in the Sahara desert, located on a vast plateau in southeastern Algeria. Having one of the most important groupings of prehistoric ...
,
Libya
Libya (; ar, ليبيا, Lībiyā), officially the State of Libya ( ar, دولة ليبيا, Dawlat Lībiyā), is a country in the Maghreb region in North Africa. It is bordered by the Mediterranean Sea to the north, Egypt to Egypt–Libya bo ...
), and in the
Horn of Africa
The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
region (
Socotra
Socotra or Soqotra (; ar, سُقُطْرَىٰ ; so, Suqadara) is an island of the Republic of Yemen in the Indian Ocean, under the ''de facto'' control of the UAE-backed Southern Transitional Council, a secessionist participant in Yemen’s ...
,
Somalia
Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constituti ...
).
The plants grow in more or less coastal habitats from sea level up to 200 m, in Macaronesia and the Western Mediterranean region, and also in Socotra. But in the North African mountains they were found at higher altitudes up to 2000 m, and also in Somalia at 1150 m.
type species
In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''Patellifolia webbiana''. The same authors had described this taxon already in 1976 as ''Patellaria'' J.T. Williams, A.J.Scott & Ford-Lloyd, but that was an illegitimate name, (because the
lichen
A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.Patellaria'' Hoffmann was published earlier in 1789).
These plants were first grouped together in 1927 by
Vladimir Andreevich Tranzschel
Vladimir Andreevich Tranzschel (russian: Владимир Андреевич траншель) (–January 21, 1942) was a Russian botanist, mycologist and plant pathologist, especially an expert on rust fungi.
He graduated from Saint Petersburg ...
as an informal unranked taxon ''Patellares'' in the genus ''Beta''. In 1934,
Oskar Eberhard Ulbrich
Oskar Eberhard Ulbrich (17 September 1879 – 4 November 1952) was a German botanist and mycologist.
Ulbrich was born in Berlin. He studied natural sciences at the University of Berlin, where his instructors included Adolf Engler (1844–1930) and ...
placed them in a new section ''Beta'' sect. ''Procumbentes''. Until 2006, ''Patellifolia'' was often not accepted as a separate genus. But recent molecular genetic studies by Kadereit et al. (2006) and Romeiras et al. (2016) revealed a deep genetic differentiation between ''Beta'' and ''Patellifolia'' and confirmed the status as own genus.
''Patellifolia'' comprises one to three species:
* '' Patellifolia patellaris'' (Moq.) A.J. Scott & al. (Syn. ''Beta patellaris'' Moq.), in dry coastal sites or on rocks, widely distributed in Macaronesia (Canary Islands, Madeira, Cape Verde) and in the western Mediterranean (Spain, Balearic islands, Sicily, Morocco), North Africa, to the Horn of Africa.
* '' Patellifolia procumbens'' (Chr. Sm.) A.J. Scott & al. (Syn. ''Beta procumbens'' Chr. Sm.), in dry coastal sites in Macaronesia (Canary Islands, Madeira, Cape Verde).
* '' Patellifolia webbiana'' (Moq.) A.J. Scott & al. (Syn. ''Beta webbiana'' Moq.), endemic on
Gran Canaria
Gran Canaria (, ; ), also Grand Canary Island, is the third-largest and second-most-populous island of the Canary Islands, an archipelago off the Atlantic coast of Northwest Africa which is part of Spain. the island had a population of that co ...
, in ruderal nitrophilous sites, Critically Endangered.
It was suggested that ''Patellifolia procumbens'' and ''Patellifolia webbiana'' might not be different species. The identification of the three species is difficult, and some distinguishing characters were found to be unreliable. Therefore, Thulin et al. (2010) proposed to treat all members of ''Patellifolia'' as a single variable species, '' Patellifoia procumbens''. This proposal was followed in the Euro+Med Plant Base.
But ''Patellifolia patellaris'' could not be hybridised with the other two species, supporting the rank of an own species. The island species evolved recently, and because of their isolation they developed only small genetic differentiation. As a result, they show only weak genetic barriers to hybridization.
Evolution
''Patellifolia'' is a rather old genus; the divergence from its sister genus ''Beta'' probably occurred early in the Late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
. Its subtaxa are relatively young, diversifying in the late
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Quaternary
The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS). It follows the Neogene Period and spans from 2.58 million years ...
. The present distribution area suggests that it was previously a widespread taxon, but later the area fragmented and extensive extinctions occurred.
Crop wild relatives
''Patellifolia'' is related to the economically important cultivated beets (''
Beta vulgaris
''Beta vulgaris'' (beet) is a species of flowering plant in the subfamily Betoideae of the family Amaranthaceae. Economically, it is the most important crop of the large order Caryophyllales. It has several cultivar groups: the sugar beet, of gre ...
'' subsp. ''vulgaris'', with
sugar beet
A sugar beet is a plant whose root contains a high concentration of sucrose and which is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together wi ...
,
beetroot
The beetroot is the taproot portion of a beet plant, usually known in North America as beets while the vegetable is referred to as beetroot in British English, and also known as the table beet, garden beet, red beet, dinner beet or golden beet ...
,
chard
Chard or Swiss chard (; ''Beta vulgaris'' subsp. ''vulgaris'', Cicla Group and Flavescens Group) is a green leafy vegetable. In the cultivars of the Flavescens Group, the leaf stalks are large and often prepared separately from the leaf blade; ...
and
fodder beet
Mangelwurzel or mangold wurzel (from :de:Futterrübe, German ''Mangel/Mangold'', "chard" and ''Wurzel'', "root"), also called mangold,Wright, Clifford A. (2001) ''Mediterranean Vegetables: a cook's ABC of vegetables and their preparation in Spain ...
). Therefore, ''Patellifolia'' is of great interest as
Crop wild relative
A crop wild relative (CWR) is a wild plant closely related to a domesticated plant. It may be a wild ancestor of the domesticated (cultivated) plant or another closely related taxon.
Overview
The wild relatives of crop plants constitute an i ...
and potential gene donor. They seem to be resistant to some of the most serious diseases of sugar beets, to
leaf spot
A leaf spot is a limited, discoloured, diseased area of a leaf that is caused by fungal, bacterial or viral plant diseases, or by injuries from nematodes, insects, environmental factors, toxicity or herbicides. These discoloured spots or lesions ...
disease caused by ''
Cercospora beticola
''Cercospora beticola'' is a fungus, fungal plant pathogen which typically infects plants of the genus ''Beta (plant), Beta'', within the family of Chenopodiaceae. It is the cause of Cercospora leaf spot disease in sugar beets, spinach and swiss ...
'',
curly top
Curly Top is a 1935 American musical romantic comedy film starring Shirley Temple, John Boles and Rochelle Hudson.
Plot
A bachelor wants to adopt an orphan, but she refuses to leave behind her older sister, so he adopts them both. The man even ...
powdery mildew
Powdery mildew is a fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant diseases to identify, as ...
(''
Erysiphe polygoni
''Erysiphe betae'' is a plant pathogen. It is a form of powdery mildew that can affect crops of sugar beet, when it can cause up to a 30% yield loss.Heffer, V., M. L. Powelson, K. B. Johnson, and N. Shishkoff. "Identification of Powdery Mildew Fu ...
''). All three species proved to be highly resistant to the Beet cyst eelworm (''Heterodera schachtii''). The three species differ in their resistance level, whereas ''Patellifolia patellaris'' is not completely immune, but eelworms never develop to maturity in the other two species.
References
Bolibar, S.C. ''Patellifolia'' in Flora Iberica p.482-484., accessed 20 August 2016.Santos-Guerra, A.; Frese, L. (2011). ''Patellifolia webbiana''. The IUCN Red List of Threatened Species 2011: e.T172260A6859581. . Accessed 19 August 2016.Kadereit, G., Hohmann, S. & Kadereit, J.W. (2006). "A synopsis of Chenopodiaceae subfam. Betoideae and notes on the taxonomy of ''Beta''". - Willdenowia 36, p. 9-19.Romeiras, M.M., Vieira, A., Silva, D.N., Moura, M., Santos-Guerra, A., Batista, D., Duarte, M.C., & Paulo, O.S. (2016). "Evolutionary and Biogeographic Insights on the Macaronesian ''Beta-Patellifolia'' Species (Amaranthaceae) from a Time-Scaled Molecular Phylogeny." PLoS One. 2016; 11(3): e0152456. Scott, A.J., Ford-Lloyd, B.V. & Williams, J.T. (1977). "''Patellifolia'', nomen novum (Chenopodiaceae)". Taxon 26(2–3): 284. Thulin, M.; Rydberg, A.; Thiede, H. (2010). "Identity of ''Tetragonia pentandra'' and taxonomy and distribution of ''Patellifolia'' (Chenopodiaceae)". Willdenowia 40(1): 5-11. Ulbrich, O.E. (1934). "Chenopodiaceae", in: Engler, A. & Prantl, K.A.E. (eds.): "Die natürlichen Pflanzenfamilien", ed.2 16c, Engelmann, Leipzig: 455.Uotila, P. (2011) ''Patellifolia.'' In: "Chenopodiaceae (pro parte majore)." In: Euro+Med Plantbase - the information resource for Euro-Mediterranean plant diversity.