Dryopteridaceae
Encyclopedia
Dryopteridaceae, is a family
of leptosporangiate fern
s in the order
Polypodiales
. They are known colloquially
as the wood ferns. They comprise about 1700 species
and have a cosmopolitan distribution
. They may be terrestrial
, epipetric
, hemiepiphytic
, or epiphytic. Many are cultivated
as ornamental plant
s. The largest genera
are Elaphoglossum (600), Polystichum (260), Dryopteris (225), and Ctenitis (150). These four genera contain about 70% of the species. Dryopteridaceae diverged
from the other families in eupolypods I
about 100 Mya (million years ago).
, with non-clathrate scales
at apices. Frond
s usually monomorphic, less often dimorphic
, sometimes scaly or glandular
, less commonly hairy. Petioles
with numerous round, vascular bundles
arranged in a ring, or rarely as few as 3; the adaxial bundles largest. Veins pinnate
or forking
, free to variously anastomosing
; the areole
s with or without included veinlets; Sori usually round, acrostichoid (covering the entire abaxial surface of the lamina) in a few lineages
; usually indusiate, sometimes exindusiate. Indusia, when present, round-reniform or peltate. Sporangia with 3-rowed, short to long stalks; spore
s reniform, monolete, perine winged.
sensu lato, Onocleaceae
and most of Tectariaceae
. Molecular phylogenetic studies
found Kramer's version
of Dryopteridaceae to be polyphyletic and it was split up
by Smith and others in 2006. The inclusion of Didymochlaena, Hypodematium
, and Leucostegia in Dryopteridaceae is doubtful. If these three are excluded, then the family is strongly supported
as monophyletic in cladistic analyses. Some authors have already treated
these genera outside of Dryopteridaceae.
Nothoperanema is now included in Dryopteris
. In 2007, a phylogenetic study of DNA sequence
s showed that Pleocnemia should be transferred from Tectariaceae to Dryopteridaceae. In 2010, in a paper on bolbitidoid ferns, Arthrobotrya was resurrected from Teratophyllum. Later that year, Mickelia was described as a new genus.
Some species have been removed from the genus Oenotrichia because they do not belong there or even in the family Dennstaedtiaceae
where Oenotrichia sensu stricto is placed. These species probably belong in Dryopteridaceae, but have not yet been given a generic name.
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of leptosporangiate fern
Leptosporangiate fern
Leptosporangiate ferns are the largest group of living ferns. They are often considered to be the class Pteridopsida or Polypodiopsida, although other classifications assign them a different rank...
s in the order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
Polypodiales
Polypodiales
The 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 tropical, semitropical and temperate areas...
. They are known colloquially
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
as the wood ferns. They comprise about 1700 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
and have a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...
. They may be terrestrial
Terrestrial plant
A terrestrial plant is one that grows on land. Other types of plants are aquatic , epiphytic , lithophytes and aerial ....
, epipetric
Epipetric
An epipetric plant, is one that is found growing on rocks. Many ferns fall into this category of plant habitat, including Asplenium . Epipetric plants are called Lithophytes....
, hemiepiphytic
Hemiepiphyte
A hemiepiphyte is a plant which begins its life as an epiphyte but which later grows roots down into the ground. The seeds of hemiepiphytes germinate in the canopy and initially live epiphytically...
, or epiphytic. Many are cultivated
Gardening
Gardening is the practice of growing and cultivating plants. Ornamental plants are normally grown for their flowers, foliage, or overall appearance; useful plants are grown for consumption , for their dyes, or for medicinal or cosmetic use...
as ornamental plant
Ornamental plant
Ornamental plants are plants that are grown for decorative purposes in gardens and landscape design projects, as house plants, for cut flowers and specimen display...
s. The largest genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
are Elaphoglossum (600), Polystichum (260), Dryopteris (225), and Ctenitis (150). These four genera contain about 70% of the species. Dryopteridaceae diverged
Evolutionary radiation
An evolutionary radiation is an increase in taxonomic diversity or morphological disparity, due to adaptive change or the opening of ecospace. Radiations may affect one clade or many, and be rapid or gradual; where they are rapid, and driven by a single lineage's adaptation to their environment,...
from the other families in eupolypods I
Eupolypods I
Eupolypods I is a clade of ferns in the order Polypodiales. It probably diverged from Eupolypods II during the mid-Cretaceous. The divergence is supported by both molecular data and an often over-looked morphological characteristic which lies in the vasculature of the petiole. Most species that...
about 100 Mya (million years ago).
Genera
In the list below, the first column consists of the 36 genera recognized by Alan R. Smith and coauthors in 2006. The second column contains extinct genera and genera that have been added since the 2006 classification.
|
Mickelia Mickelia is a genus of ferns in the family Dryopteridaceae. It is one of the six genera of bolbitidoid ferns and is sister to the very large genus Elaphoglossum. It consists of 10 species. All are native to the neotropics.- Species :... Wessiea Wessiea is an extinct monotypic genus of fern in the Dryopteridaceae family with the sole species Wessiea yakimaensis. Wessia is known from Langhian age Miocene fossils found in Central Washington.... |
Description
Rhizomes often stout; creeping, ascending or erect, sometimes scandent or climbingVine
A vine in the narrowest sense is the grapevine , but more generally it can refer to any plant with a growth habit of trailing or scandent, that is to say climbing, stems or runners...
, with non-clathrate scales
Trichome
Trichomes are fine outgrowths or appendages on plants and certain protists. These are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae.- Algal trichomes :...
at apices. Frond
Frond
The term frond refers to 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...
s usually monomorphic, less often dimorphic
Frond dimorphism
Frond dimorphism refers to a difference in ferns between the fertile and sterile fronds. Since ferns, unlike flowering plants, bear spores on the leaf blade itself, this may affect the form of the frond itself...
, sometimes scaly or glandular
Gland (botany)
In plants, a gland is defined functionally as a plant structure which secretes one or more products. This may be located on or near the plant surface and secrete externally, or be internal to the plant and secrete into a canal or reservoir...
, less commonly hairy. Petioles
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...
with numerous round, vascular bundles
Vascular tissue
Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue:...
arranged in a ring, or rarely as few as 3; the adaxial bundles largest. Veins pinnate
Pinnate
Pinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
or forking
Divaricate
Divaricate is a term used in identifying plants describing the pattern of branching. Divaricate branching is roughly horizontal, usually only diverging about 15 degrees upward or downward....
, free to variously anastomosing
Anastomosis
An anastomosis is the reconnection of two streams that previously branched out, such as blood vessels or leaf veins. The term is used in medicine, biology, mycology and geology....
; the areole
Areole
Areoles are an important diagnostic feature of cacti, and identify them as a family distinct from other succulent plants. The areoles on cacti are clearly visible; they generally appear as small light- to dark-colored bumps, out of which grow clusters of spines...
s with or without included veinlets; Sori usually round, acrostichoid (covering the entire abaxial surface of the lamina) in a few lineages
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
; usually indusiate, sometimes exindusiate. Indusia, when present, round-reniform or peltate. Sporangia with 3-rowed, short to long stalks; spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
s reniform, monolete, perine winged.
History
In 1990, Karl U. Kramer and coauthors defined Dryopteridaceae broadly to include the present family as well as WoodsiaceaeWoodsiaceae
Woodsiaceae or Cliff Fern is a family of fern within the Aspleniales order....
sensu lato, Onocleaceae
Onocleaceae
Onocleaceae is a small family of terrestrial ferns. There are four genera of onocleoids: Matteuccia, Onoclea, Onocleopsis, and Pentarhizidium, consisting of five species largely in north temperate climes....
and most of Tectariaceae
Tectariaceae
Tectariaceae is a family of Leptosporangiate ferns in the order Polypodiales. It comprises 10 genera. Tectaria is much larger than the other genera....
. Molecular phylogenetic studies
Research
Research can be defined as the scientific search for knowledge, or as any systematic investigation, to establish novel facts, solve new or existing problems, prove new ideas, or develop new theories, usually using a scientific method...
found Kramer's version
Circumscription (taxonomy)
In taxonomy, circumscription is the definition of the limits of a taxonomic group of organisms. One goal of taxonomy is to achieve a stable circumscription for every taxonomic group. Achieving stability can be simple or difficult....
of Dryopteridaceae to be polyphyletic and it was split up
Segregate (taxonomy)
In taxonomy, a segregate, or a segregate taxon is created when a taxon is split off, from another taxon. This other taxon will be better known, usually bigger, and will continue to exist, even after the segregate taxon has been split off...
by Smith and others in 2006. The inclusion of Didymochlaena, Hypodematium
Hypodematium
-Species:*Hypodematium crenatum Kuhn*Hypodematium daochengense K. H. Shing*Hypodematium fordii Ching*Hypodematium glabrius Holtt.*Hypodematium glabrum Ching ex Shing*Hypodematium glandulosum Ching ex Shing...
, and Leucostegia in Dryopteridaceae is doubtful. If these three are excluded, then the family is strongly supported
Resampling (statistics)
In statistics, resampling is any of a variety of methods for doing one of the following:# Estimating the precision of sample statistics by using subsets of available data or drawing randomly with replacement from a set of data points # Exchanging labels on data points when performing significance...
as monophyletic in cladistic analyses. Some authors have already treated
Treatise
A treatise is a formal and systematic written discourse on some subject, generally longer and treating it in greater depth than an essay, and more concerned with investigating or exposing the principles of the subject.-Noteworthy treatises:...
these genera outside of Dryopteridaceae.
Nothoperanema is now included in Dryopteris
Dryopteris
Dryopteris , commonly called wood ferns, male ferns, and buckler ferns, is a genus of about 250 species of ferns with distribution in the temperate Northern Hemisphere, with the highest species diversity in eastern Asia. Many of the species have stout, slowly creeping rootstocks that form a crown,...
. In 2007, a phylogenetic study of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
s showed that Pleocnemia should be transferred from Tectariaceae to Dryopteridaceae. In 2010, in a paper on bolbitidoid ferns, Arthrobotrya was resurrected from Teratophyllum. Later that year, Mickelia was described as a new genus.
Some species have been removed from the genus Oenotrichia because they do not belong there or even in the family Dennstaedtiaceae
Dennstaedtiaceae
Dennstaedtiaceae is one of fifteen families in the order Polypodiales, the most derived families within monilophytes . It includes the world's most abundant fern, Pteridium aquilinum . Members of the order generally have large, highly divided leaves and have either small, round intramarginal sori...
where Oenotrichia sensu stricto is placed. These species probably belong in Dryopteridaceae, but have not yet been given a generic name.