Patellifolia
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''Patellifolia'' is a genus of
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 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, maki ...
. These are mostly procumbent herbs occurring in the Western Mediterranean region and
Macaronesia Macaronesia (; ) is a collection of four volcanic archipelagos in the Atlantic Ocean#North Atlantic, North Atlantic Ocean, off the coast of North Africa and Europe. Each archipelago is made up of a number of list of islands in the Atlantic Oc ...
, with some isolated occurrences in
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
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 of 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 ...
s of
sugar beet A sugar beet is a plant whose root contains a high concentration of sucrose and that is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together with ...
.


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), a ...
or perennial herbs, growing erect or often procumbent. The alternate
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
have a
petiole Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, and ...
, their leaf blade is heart-shaped or hastate. The spike-like
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 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, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
s. The free
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s are hermaphrodite. The
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...
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 (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
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 (
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
,
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
),
Macaronesia Macaronesia (; ) is a collection of four volcanic archipelagos in the Atlantic Ocean#North Atlantic, North Atlantic Ocean, off the coast of North Africa and Europe. Each archipelago is made up of a number of list of islands in the Atlantic Oc ...
and
Cape Verde Cape Verde or Cabo Verde, officially the Republic of Cabo Verde, is an island country and archipelagic state of West Africa in the central Atlantic Ocean, consisting of ten volcanic islands with a combined land area of about . These islands ...
Islands. There are also some isolated occurrences in North Africa (
Hoggar Mountains The Hoggar Mountains (; 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 km2. The Hoggar Mountains are home to ...
and
Tassili n'Ajjer Tassili n'Ajjer (Berber: ''Tassili n Ajjer'', ; "Plateau of rivers") is a mountain range in the Sahara desert, located in south-eastern Algeria. It holds one of the most important groupings of prehistoric cave art in the world, and covers an ar ...
,
Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...
), 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, locally known as Saqatri, is a Yemeni island in the Indian Ocean. Situated between the Guardafui Channel and the Arabian Sea, it lies near major shipping routes. Socotra is the largest of the six islands in the Socotra archipelago as ...
,
Somalia Somalia, officially the Federal Republic of Somalia, is the easternmost country in continental Africa. The country is located in the Horn of Africa and is bordered by Ethiopia to the west, Djibouti to the northwest, Kenya to the southwest, th ...
). 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.


Systematics

The genus ''Patellifolia'' was published in 1977 by A J Scott, Brian V. Ford-Lloyd and J. Trevor Williams, with the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''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 hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
genus '' Patellaria'' Hoffmann was published earlier in 1789). These plants were first grouped together in 1927 by Vladimir Andreevich Tranzschel 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, a Spain, Spanish archipelago off the Atlantic coast of Northwest Africa. the island had a population of that constitut ...
, 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 (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
. Its subtaxa are relatively young, diversifying in the late
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 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), as well as the current and most recent of the twelve periods of the ...
. 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 that is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together with ...
,
beetroot The beetroot (British English) or beet (North American English) is the taproot portion of a ''Beta vulgaris'' subsp. ''vulgaris'' plant in the Conditiva Group. The plant is a root vegetable also known as the table beet, garden beet, dinner bee ...
,
chard Chard (; '' Beta vulgaris'' subsp. ''vulgaris'', Cicla Group and Flavescens Group) is a green leafy vegetable. In the cultivars of the Flavescens Group, or Swiss chard, the leaf stalks are large and often prepared separately from the leaf b ...
and fodder beet). 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 viral disease that affects many crops. This disease causes plants to become smaller in size, have shriveled petals and leaves, and are twisted and pulled out of shape. They are often caused by curtoviruses (genus ''Curtovirus''), m ...
virus, rhizomania, and
powdery mildew Powdery mildew is a fungus, fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of Ascomycota, ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant disea ...
(''
Erysiphe polygoni ''Erysiphe betae'' is a fungal plant pathogen. It is a form of powdery mildew that can affect crops of sugar beet A sugar beet is a plant whose root contains a high concentration of sucrose and that is grown commercially for sugar productio ...
''). 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.


External links

* *
Illustration of ''Patellifolia patellaris'' at Flora Iberica
{{Taxonbar, from=Q6066269 Amaranthaceae Amaranthaceae genera