Davallia
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'' Davallia'' (deersfoot fern, hare's foot fern, shinobu fern, rabbit foot fern, ball fern) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of about 40
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
. In the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 (PPG I), it is the only genus in the family Davalliaceae, which is placed in the suborder
Polypodiineae Polypodiineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods I in earlier systems, and to the very broadly defined family Polypodiaceae in the classification of Christenhusz & Chase (2014). It probably ...
, order
Polypodiales The Order (biology), order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including Tropics, tropical, semitropical and Temperate clima ...
. Alternatively, the family may be placed in a very broadly defined family
Polypodiaceae Polypodiaceae is a Family (biology), family of ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family includes around 65 genus, genera and an estimated 1,650 species and is placed in the order Polypodiales, suborder ...
''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'' as the subfamily Davallioideae. The family is
sister A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
to the largest family of ferns, Polypodiaceae, and shares some morphological
characters Character or Characters may refer to: Arts, entertainment, and media Literature * ''Character'' (novel), a 1936 Dutch novel by Ferdinand Bordewijk * ''Characters'' (Theophrastus), a classical Greek set of character sketches attributed to Theoph ...
with it.Karl U. Kramer. 1990. "Davalliaceae". pages 74-80. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and Genera of Vascular Plants'' volume I. Springer-Verlag: Berlin;Heidelberg, Germany. Species are
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
ferns, with
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s arising from long aerial
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s which grow on and over thick
bark Bark may refer to: Common meanings * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Arts and entertainment * ''Bark'' (Jefferson Airplane album), ...
on
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s or on rock crevices.


Description

Usually
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
or epipetric.
Rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s dictyostelic,
dorsiventral A dorsiventral (Lat. ''dorsum'', "the back", ''venter'', "the belly") organ is one that has two surfaces differing from each other in appearance and structure, as an ordinary leaf. This term has also been used as a synonym for dorsoventral organs, ...
, densely scaly.
Stipes Crucifixion is a method of capital punishment in which the condemned is tied or nailed to a large wooden cross, beam or stake and left to hang until eventual death. It was used as a punishment by the Achaemenid Empire, Persians, Carthaginians, ...
articulate at base. Phyllopodia short.
veins Veins () are blood vessels in the circulatory system of humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are those of the pulmonary and fetal c ...
free.
Sporangium A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
stalk 3-rowed. Annulus vertical.
Spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s monolete.


Taxonomy

'' Gymnogrammitis'' and ''
Leucostegia ''Leucostegia'' is a genus of ferns in the family Hypodematiaceae in the Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classifi ...
'' were once included in Davalliaceae, but these are now known to belong elsewhere. ''Gymnogrammitis'' is in a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
with '' Selliguea'' and others in the family Polypodiaceae.Harald Schneider, Alan R. Smith, Raymond Cranfill, Christopher H. Haufler, Tom A. Ranker, and Terri J. Hildebrand. 2002. "''Gymnogrammitis dareiformis'' is a polygrammoid fern (Polypodiacee) - Resolving an apparent conflict between morphological and molecular data". ''Plant Systematics and Evolution'' 234(1-4):121-136. ''Leucostegia'' is in the family
Hypodematiaceae Hypodematiaceae is a family of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Hypod ...
, which consists of ''
Hypodematium ''Hypodematium'' is a genus of ferns and is the type genus of the family Hypodematiaceae Hypodematiaceae is a family of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is pl ...
'' and ''Leucostegia'',Hong-Mei Liu, Xian-Chun Zhang, Wei Wang, Yin-Long Qiu, and Zhi-Duan Chen. 2007. "Molecular Phylogeny of the Fern Family Dryopteridaceae inferred from Chloroplast rbcL and atpB Genes". ''International Journal of Plant Sciences'' 168(9):1311-1323. and possibly ''
Didymochlaena ''Didymochlaena'' is a genus of ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is the only genus in the family ''Didymochlaenaceae''. Alternatively, the family may be placed in a very broadly defined family Polypodi ...
'' as well. In 2008, a
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study Study or studies may refer to: General * Education **Higher education * Clinical trial * Experiment * Field of study * Observational study * Scientific study * Research * Study skills, abilities and approaches applied to learning Other * Study ...
of Davalliaceae showed that none of the polyspecific genera recognized at that time were
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
.Chie Tsutsumi, Xian-Chun Zhang, and Masahiro Kato. 2008. "Molecular Phylogeny of Davalliaceae and Implications for Generic Classification". ''Systematic Botany'' 33(1):44-48. In that same year, a revision of the family divided it into five genera. One of these, ''Araiostegiella'', was newly described. The genus ''Davallia'' was divided into two
sections Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
, named ''Davallia'' and ''Trogostolon''. Based on
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies, the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 (PPG I) accepts only one genus in the family, ''Davallia'', sinking the other genera into synonymy. The study on which the PPG I
circumscription Circumscription may refer to: * Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) * Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthr ...
is based divides the genus into seven sections.


Phylogeny

The following cladogram for the suborder
Polypodiineae Polypodiineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods I in earlier systems, and to the very broadly defined family Polypodiaceae in the classification of Christenhusz & Chase (2014). It probably ...
(eupolypods I), based on the consensus cladogram in the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 (PPG I), shows a likely
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationship between Davalliaceae and the other families of the clade. Other species include: * †'' Davallia aspera'' Botscharnikova 1960 * '' Davallia brevipes'' Copeland 1906 * †'' Davallia bullatiformis'' Chlonova 1960 * †'' Davallia cavernosa'' Botscharnikova 1960 * '' Davallia dimorpha'' (Copeland) Holttum 1937 * †'' Davallia haidingeri'' von Ettingshausen 1858 * '' Davallia kinabaluensis'' (Copeland) Chen 2017 * '' Davallia leptocarpa'' Mettenius ex Kuhn 1869 * †'' Davallia mantellii'' (Brongniart 1828) de Saporta 1891 * †'' Davallia montana'' Knowlton 1899 * '' Davallia napoensis'' Wang & Xing 2011 * '' Davallia pusilloides'' (Copeland) Parris 2018 * '' Davallia rouffaeriensis'' Nooteboom 1994 * '' Davallia seramensis'' Kato 1989 * '' Davallia serrata'' Willdenow 1810 * '' Davallia sessilifolioides'' Kato 1989 * †'' Davallia solidites'' Graham 1963 * '' Davallia speciosa'' Mettenius ex Kuhn 1869 * '' Davallia subvestita'' (Christensen) Parris 2018 * '' Davallia wagneriana'' Copeland 1905


Selected species

Many of the species of ''Davallia'' are closely related and hard to
distinguish In law, to distinguish a case means a court decides the holding or legal reasoning of a precedent case that will not apply due to materially different facts between the two cases. Two formal constraints constrain the later court: the expresse ...
from each other. In 1990, a treatment of Davalliaceae estimated the number of species at 110. A 2008
paper Paper is a thin sheet material produced by mechanically or chemically processing cellulose fibres derived from wood, Textile, rags, poaceae, grasses, Feces#Other uses, herbivore dung, or other vegetable sources in water. Once the water is dra ...
listed all of the species, recognizing only 63.Masahiro Kato and Chie Tsutsumi. 2008. "Generic Classification of Davalliaceae". ''Acta Phytotaxonomica et Geobotanica'' 59(1):1-14. A new species, '' Davallia napoensis'' was described in 2011.Fa-Guo Wang, Hong-Feng Chen, and Fu-Wu Xing. 2011. "''Davallia napoensis'', a New Species of Davalliaceae from Guangxi, China". ''Novon'' 21(3):380-384. The
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 (PPG I) suggests there are about 65 species. * '' Davallia bullata'' -
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, and tropical
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
. * '' Davallia canariensis'' -
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 ...
,
Iberian Peninsula The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
and
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 ...
. * '' Davallia denticulata'' - Africa, India, China, Malesia, Indonesia, Polynesia, Australia * '' Davallia divaricata'' ( syn.: ''Davallia polyantha'') - Tropical
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
. * '' Davallia embolostegia'' * '' Davallia fejeensis'' Hook ( syn.: ''Davallia fijiensis'') -
Fiji Islands Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...
and
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. * '' Davallia mariesii'' or "Squirrel's-foot fern" – tropical Asia and Malaysia * '' Davallia pectinata'' * '' Davallia repens'' * '' Davallia solida'' - Sri Lanka, tropical and subtropical Asia (east Himalaya to Southeast Asia and China), eastern Australia, and Pacific Islands ** '' Davallia solida var. pyxidata'' -
New South Wales New South Wales (commonly abbreviated as NSW) is a States and territories of Australia, state on the Eastern states of Australia, east coast of :Australia. It borders Queensland to the north, Victoria (state), Victoria to the south, and South ...
** ''Davallia solida'' var. ''fejeensis'' (Hook.) Noot. - endemic to
Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...
** ''Davallia solida'' var. ''solida'' * '' Davallia tasmanii'' - Davallia fern, native to the
Three Kings Islands 3 (three) is a number, numeral (linguistics), numeral and numerical digit, digit. It is the natural number following 2 and preceding 4, and is the smallest odd prime number and the only prime preceding a square number. It has religious and cul ...
. * '' Davallia trichomanoides'' ( syn.: ''Davallia dissecta'') - Malaysia.


Distribution and habitat

Davalliaceae is
native Native may refer to: People * '' Jus sanguinis'', nationality by blood * '' Jus soli'', nationality by location of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Nat ...
to
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
regions of the
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
,
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, and
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
.Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider, and Paul G. Wolf. 2008. "Davalliaceae". pages 443-444. In: "Fern Classification". pages 417-467. In: Tom A. Ranker and Christopher H. Haufler (editors). ''Biology and Evolution of Ferns and Lycophytes''. Cambridge University Press. Plants are usually
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
, sometimes
lithophytic Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are also ...
or
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
.


''Davallia'' as house plants

Many species of ''Davallia'' are in cultivation,
Anthony Huxley Anthony Julian Huxley (2 December 1920 – 26 December 1992) was a British botanist and writer. An elected council member of the Royal Horticultural Society, he became its vice president in 1991. He edited '' Amateur Gardening'' from 1967 to 19 ...
, Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. (set).
with '' Davallia tyermanii'', '' Davallia fejeensis'', and '' Davallia solida'' being perhaps the most well-known.George W. Staples and Derral R. Herbst. 2005. "A Tropical Garden Flora" Bishop Museum Press: Honolulu, HI, USA. A key to the cultivated species of ''Davallia'' is available.Barbara Joe Hoshizaki. 1981. "The fern genus ''Davallia'' in cultivation". ''Baileya'' 21(1):1-42. ''D. polypodiaceae'', ''D. canariensis'' and ''D. trichomanoides'' are also grown as
ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
s.Perry, Leonard, ''Davillia'', http://pss.uvm.edu/pss123/ferndav.html ''D. fejeensis'' is the most common ''Davallia'' species in commerce, and ''D. canariensis'' is widely grown as a
house plant A houseplant, also known as a pot plant, potted plant, or indoor plant, is an ornamental plant cultivated indoors. for aesthetic or practical purposes. These plants are commonly found in homes, offices, and various indoor spaces, where they contr ...
. The plants have furry rhizomes which cover the surface of the potting mixture as well as root down into it. The fronds are triangular in shape and about 1½ feet long by 1 foot wide. They divide into three to four
pinnae Pinna may refer to: Biology * Pinna (anatomy), or auricle, the outer part of the ear * ''Pinna'' (bivalve), a genus of molluscs * Pinna (botany), a primary segment of a compound leaf People Surname * Christophe Pinna (born 1968), French marti ...
which subdivide into many pinnules. ''Davallia'' are often used in hanging baskets because the rhizomes split into sections and the surface is covered quickly. Unlike other ferns, ''Davallia'' tolerate low levels of humidity.Reader's Digest. Success with House Plants. The Reader's Digest Association, Inc. 1979:173


References

*Lorenzi, H. & Souza, M. S. (2001). ''Plantas Ornamentais no Brasil: arbustivas, herbáceas e trepadeiras.'
Plantarum
*Key, K. & Baines, J. (1974). ''El ABC de las Plantas de Interior.'' Blume *Hay, R., McQuown G., & Beckett, K. (1976). ''Diccionario ilustrado en color de plantas de interior.'' Gustavo Gili *Hellyer, A. (1976). ''The Collingridge Encyclopedia of Gardening.'' Hamlyn *Bornhorst, Heidi. ''Davallia fern has many names, various uses.'' Honolulu Advertiser, November 11, 2001

(accessed October 8, 2015) {{Taxonbar, from=Q2390278 Davalliaceae Epiphytes Fern genera Taxa named by Carl Linnaeus