Thymelaeaceae
Encyclopedia
Thymelaeaceae is a cosmopolitan family
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 flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...

s composed of 50 genera (listed below) and 898 species. It was established in 1789 by Antoine Laurent de Jussieu
Antoine Laurent de Jussieu
Antoine Laurent de Jussieu was a French botanist, notable as the first to propose a natural classification of flowering plants; much of his system remains in use today.-Life:...

.

Thymelaeaceae is in the order Malvales
Malvales
Malvales are an order of flowering plants. As circumscribed by APG II-system, it includes about 6000 species within nine families. The order is placed in the eurosids II, which are part of the eudicots....

. Except for its sister relationship with Tepuianthaceae, little is known for sure about its relationships with the other families 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...

.

The family is more diverse in the southern hemisphere
Southern Hemisphere
The Southern Hemisphere is the part of Earth that lies south of the equator. The word hemisphere literally means 'half ball' or "half sphere"...

 than in the northern
Northern Hemisphere
The Northern Hemisphere is the half of a planet that is north of its equator—the word hemisphere literally means “half sphere”. It is also that half of the celestial sphere north of the celestial equator...

, with its major concentrations of species in Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 and Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

. The genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 are overwhelmingly African

The species include mostly tree
Tree
A tree is a perennial woody plant. It is most often defined as a woody plant that has many secondary branches supported clear of the ground on a single main stem or trunk with clear apical dominance. A minimum height specification at maturity is cited by some authors, varying from 3 m to...

s and shrub
Shrub
A shrub or bush is distinguished from a tree by its multiple stems and shorter height, usually under 5–6 m tall. A large number of plants may become either shrubs or trees, depending on the growing conditions they experience...

s, with a few vine
Vine
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...

s and herbaceous plant
Herbaceous plant
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...

s.

Several genera are of economic importance. Gonystylus (Ramin), is valued for its hard, white wood. The bark of Edgeworthia and Wikstroemia is used as a component of paper.

Daphne
Daphne (plant)
Daphne is a genus of between 50 and 95 species of deciduous and evergreen shrubs in the family Thymelaeaceae, native to Asia, Europe, and north Africa...

, is grown for its sweetly scented flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

s. Species of Wikstroemia, Daphne, Phaleria
Phaleria
Phaleria is genus of flowering plant....

, Dais, Pimelea and other genera are grown as ornamentals
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...

.

Many of the species are poison
Poison
In the context of biology, poisons are substances that can cause disturbances to organisms, usually by chemical reaction or other activity on the molecular scale, when a sufficient quantity is absorbed by an organism....

ous if eaten.

A good collection of pictures of plants in this family has appeared in a scientific paper.

Herber (2003) recognized 45 genera, excluding Tepuianthus from the family, sinking Atemnosiphon and Englerodaphne into Gnidia, Eriosolena into Daphne, and Thecanthes into Pimelea.

The largest genera and the approximate number of species in each are: Gnidia (160), Pimelea (110), Daphne (95), Wikstroemia (70), Daphnopsis (65), Struthiola (35), Lachnaea (30), Thymelaea (30), Phaleria (30), and Gonystylus (25).

Genera

  • Aetoxylon
  • Amyxa
  • Aquilaria
    Aquilaria
    Aquilaria is a genus of fifteen species of trees in the Thymelaeaceae, native to southeast Asia. They occur particularly in the rain forests of Indonesia, Thailand, Cambodia, Laos, Vietnam, Malaysia, Northern India, the Philippines, Borneo and New Guinea. The trees grow to 6-20 m tall...

  • Arnhemia
  • Atemnosiphon
  • Craterosiphon
  • Dais
  • Daphne
    Daphne (plant)
    Daphne is a genus of between 50 and 95 species of deciduous and evergreen shrubs in the family Thymelaeaceae, native to Asia, Europe, and north Africa...

  • Daphnopsis
  • Deltaria
  • Diarthron
  • Dicranolepis
  • Dirca
    Dirca
    Dirca is a genus of three or four species of flowering plants in the family Thymelaeaceae, native to North America. The genus is named after Dirce in Greek mythology. The general common name for this deciduous shrub is leatherwood; others include moosewood, ropebark and Powhatan-derived wicopy...

  • Drapetes
  • Edgeworthia
    Edgeworthia
    Edgeworthia is a genus of plants in the family Thymelaeaceae. When the genus was first described, it was published twice in the same year , in two separate publications: Plantarum vascularium genera: secundum ordines naturales digesta eorumque differentiae et affinitates tabulis diagnostacis...

  • Englerodaphne
  • Enkleia
  • Eriosolena
  • Funifera
  • Gnidia
    Gnidia
    Gnidia is a genus of the plant family Thymelaeaceae. It contains the following species :* Gnidia socotrana, Gilg...

  • Gonystylus
  • Goodallia
  • Gyrinops
  • Jedda
    Jedda (plant)
    Jedda is a monotypic plant genus in the Thymelaeaceae family, its only species being J. multicaulis. The species is an upright, evergreen bush indigenous to tropical Australia, discovered in the early 1980s on the Cape York Peninsula in northern Queensland, in an area of grassy forest land....

  • Kelleria
  • Lachnaea
  • Lagetta
  • Lasiadenia
  • Lethedon
  • Linodendron
  • Linostoma
  • Lophostoma
  • Octolepis
  • Ovidia
  • Passerina
    Passerina (plant)
    Passerina is a genus in the plant family Thymelaeaceae.-List of species:*Passerina aragonensis*Passerina argentata*Passerina broteriana*Passerina burchellii*Passerina choulettei*Passerina clementii*Passerina comosa...

  • Peddiea
    Peddiea
    Peddiea is a genus of plant in family Thymelaeaceae. It contains the following species :* Peddiea kivuensis, Robyns...

  • Phaleria
    Phaleria
    Phaleria is genus of flowering plant....

  • Pimelea
    Pimelea
    Pimelea is a genus of plants belonging to the family Thymelaeaceae. There are about 80 species in the genus, native to Australia and New Zealand. Many of the species are poisonous to cattle.Selected species...

  • Rhamnoneuron
  • Schoenobiblus
  • Solmsia
    Solmsia
    Solmsia is a genus, containing two species, of flowering plants belonging to the family Thymelaeaceae.It was named to honor Hermann zu Solms-Laubach by Henri Ernest Baillon....

  • Stellera
  • Stephanodaphne
  • Struthiola
  • Synandrodaphne
  • Synaptolepis
  • Tepuianthus
  • Thecanthes
  • Thymelaea
    Thymelaea
    Thymelaea is a genus of about 30 species of evergreen shrubs and herbs in the flowering plant family Thymelaeaceae, native to the Canary Islands, the Mediterranean region, north to central Europe, and east to central Asia.Species-References:**...

  • Wikstroemia
    Wikstroemia
    Wikstroemia is a genus of 55-70 species of flowering shrubs and small trees in the mezereon family, Thymelaeaceae.-Medicinal uses:Wikstroemia indica is one of the 50 fundamental herbs used in traditional Chinese medicine.-Species:...



  • In the past, different authors have defined Thymelaeaceae in different ways. For example, John Hutchinson
    John Hutchinson (botanist)
    John Hutchinson, OBE, FRS was a renowned English botanist, taxonomist and author.-Life and career:...

     excluded Gonystylus and its close relatives, as well as Aquilaria and its close relatives from the family, forming 2 segregate families: Gonystylaceae and Aquilariaceae. But today, the only controversy that still remains over the circumscription of the family is the question of whether Tepuianthus should be included, or segregated as a separate, monogeneric family. Stevens includes Tepuianthus, but Kubitzki rejects this hypothesis and treats Tepuianthaceae as a separate family.

    Description

    This is not intended as a full botanical description, but only as a few notes on some of the conspicuous or unusual traits of the family when Tepuianthus is excluded. The bark is usually shiny and fibrous. Attempts to break the stem often result in a strip of bark peeling down the side. The number of stamen
    Stamen
    The stamen is the pollen producing reproductive organ of a flower...

    s is usually once or twice the number of calyx lobes. If twice, then they often occur in 2 well separated series. Exceptions include Gonystylus, which may have up to 100 stamens, and Pimelea, which has only 1 or 2. The floral tube appears to be a calyx or corolla, but is actually a hollow receptacle. This feature is probably unique to Thymelaeaceae. The sepal
    Sepal
    A sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...

    s are mounted on the rim of the floral tube. Stamens may be mounted on the rim or inside. What appear to be petal
    Petal
    Petals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...

    s are actually stipular
    Stipule
    In botany, stipule is a term coined by Linnaeus which refers to outgrowths borne on either side of the base of a leafstalk...

     appendages of the sepals. The fruit is a 1-seeded berry
    Berry
    The botanical definition of a berry is a fleshy fruit produced from a single ovary. Grapes are an example. The berry is the most common type of fleshy fruit in which the entire ovary wall ripens into an edible pericarp. They may have one or more carpels with a thin covering and fleshy interiors....

     or an achene
    Achene
    An achene is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate and indehiscent...

    . The 1-seeded berries have often been mistaken for drupe
    Drupe
    In botany, a drupe is a fruit in which an outer fleshy part surrounds a shell of hardened endocarp with a seed inside. These fruits develop from a single carpel, and mostly from flowers with superior ovaries...

    s whenever the seed coat was mistaken for an endocarp.

    When using a key
    Identification key
    In biology, an identification key is a printed or computer-aided device that aids the identification of biological entities, such as plants, animals, fossils, microorganisms, and pollen grains...

     to the families of flowering plants, Thymelaeaceae is often difficult or impossible to recognize because of equivocal interpretation of the flower parts. Sepals, petals, and staminode
    Staminode
    In botany, a staminode is an often rudimentary, sterile or abortive stamen. This means that it does not produce pollen. Staminodes are frequently inconspicuous and stamen-like, usually occurring at the inner whorl of the flower, but are also sometimes long enough to protrude from the...

    s are hard to distinguish, and many keys are ambiguous about whether staminodes should be counted as stamens. Moreover, in Wikstroemia, individual plants often produce anomalous flowers. In these, the nonfunctional organs are much deformed and bear little resemblance to the parts that they represent.

    Defining the genera

    The circumscription
    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 genera in Thymelaeaceae has always been especially difficult, and is to some degree, artificial. For example, the difficulty of distinguishing Daphne from Wikstroemia has been commented upon by Rautenbach and Herber. Several small genera are probably embedded in Daphne or Wikstroemia, or if Daphne and Wikstroemia are intermingled, these small genera might be embedded in both simultaneously. Stellera, for example, is nested within Wikstroemia, at least (see the phylogenetic
    Phylogenetics
    In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

     tree below).

    A recent comparison of DNA sequences has established the monophyly
    Monophyly
    In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

     of Thymelaea and the polyphyly
    Polyphyly
    A polyphyletic group is one whose members' last common ancestor is not a member of the group.For example, the group consisting of warm-blooded animals is polyphyletic, because it contains both mammals and birds, but the most recent common ancestor of mammals and birds was cold-blooded...

     of Diarthron, but there was not sufficient sampling in Wikstroemia and Daphne to exclude the possibility that Thymelaea, Diarthron, and others might be embedded in them.

    The large genus Gnidia is polyphyletic and its species fall into 4 separate clades, each of which contains other genera of the family (see the phylogenetic tree below). The type species
    Type species
    In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

     for Gnidia is Gnidia pinifolia. If Gnidia is divided into 4 or more separate genera, the segregate genus which contains G. pinifolia will retain the name Gnidia. Zachary S. Rogers published a revision of the Gnidia of Madagascar in 2009 in Annals of the Missouri Botanical Garden
    Annals of the Missouri Botanical Garden
    Annals of the Missouri Botanical Garden is a long-established major peer-reviewed journal of botany, established in 1914 by the Missouri Botanical Garden, under the directorship of botanist and phycologist, George Thomas Moore, and still published quarterly by the Missouri Botanical Garden Press. ...

    .

    Some of the older treatments of Thymelaeaceae recognize Lasiosiphon as a separate genus from Gnidia. This distinction was later shown to be artificial. However, Van der Bank et al. (2002) suggested that Lasiosiphon might be resurrected if redefined. The type species for Lasiosiphon is Gnidia glauca, formerly known as Lasiosiphon glaucus.

    Classification

    Unlike most recent authors, who recognize 4 subfamilies, B.E. Herber has divided Thymelaeaceae into 2 subfamilies. He has retained the subfamily Gonostyloideae, but renamed it Octolepidoideae. The other 3 traditional subfamilies (Synandrodaphnoideae, Aquilarioideae, and Thymelaeoideae) were combined into a Thymelaeoideae s.l.(sensu lato), and reduced to tribal
    Tribe (biology)
    In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...

     rank, as Synandrodaphneae, Aquilarieae, and Daphneae, respectively. No tribes were designated in subfamily Octolepidoideae, but it was provisionally divided into 2 informal groups, the Octolepis group and the Gonystylus group. Likewise, no subtribes
    Tribe (biology)
    In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...

     were designated in the tribe Daphneae, but it was informally divided into 4 groups: the Linostoma group, the Daphne group, the Phaleria group, and the Gnidia group. The 45 genera recognized by Herber are grouped as follows. Three genera in Daphneae were placed incertae sedis
    Incertae sedis
    , is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

     (not assigned to any particular group or in a separate group by themselves).

    Octolepidoidae

    Octolepis group: Arnhemia, Deltairia, Lethedon, Octolepis, Solmsia
    Gonystylus group: Aetoxylon, Amyxa, Gonystylus

    Thymelaeoideae

    Synandrodaphneae: Synandrodaphne
    Aquilarieae: Aquilaria, Gyrinops
    Daphneae

    Linostoma group: Craterosiphon, Dicranolepis, Enkleia, Jedda, Linostoma, Lophostoma, Synaptolepis
    Phaleria group: Phaleria, Peddiea
    Daphne group: Daphne, Daphnopsis, Diarthron, Dirca, Edgeworthia, Funifera, Goodallia, Lagetta, Ovidia, Rhamnoneuron, Schoenobiblus, Stellera, Thymelaea, Wikstroemia
    Gnidia group: Dais, Drapetes, Gnidia, Kelleria, Lachnaea, Passerina, Pimelea, Struthiola
    Incertae sedis: Linodendron, Stephanodaphne, Lasiadenia

    Phylogeny

    The first molecular phylogeny for Thymelaeaceae was published in 2002. It was based on 2 regions of chloroplast
    Chloroplast
    Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

     DNA
    DNA
    Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

    . These were the rbcL gene
    Gene
    A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...

     and the intergenic spacer between the transfer RNA
    Transfer RNA
    Transfer RNA is an adaptor molecule composed of RNA, typically 73 to 93 nucleotides in length, that is used in biology to bridge the three-letter genetic code in messenger RNA with the twenty-letter code of amino acids in proteins. The role of tRNA as an adaptor is best understood by...

     genes trnL and trnF. Forty one species in the family were sampled. In 2008, Marline Rautenbach performed a phylogenetic study in which 143 species in the family were sampled. The sampling in this study was concentrated in the Gnidia group, but the sampling in the rest of the family was as extensive as in the previous study, or more so. In addition to rbcL and trnL-F data, sequences of the ITS (internal transcribed spacer
    Internal transcribed spacer
    ITS refers to a piece of non-functional RNA situated between structural ribosomal RNAs on a common precursor transcript. Read from 5' to 3', this polycistronic rRNA precursor transcript contains the 5' external transcribed sequence , 18S rRNA, ITS1, 5.8S rRNA, ITS2, 28S rRNA and finally the 3'ETS...

    ) region of nrDNA (nuclear ribosomal DNA) were used. All of the clades that were strongly supported in the previous study were recovered with even stronger statistical support.

    The tree below is an excerpt from the Rautenbach (2002) phyogeny. The species of Gnidia were chosen from among the most common or well known species in a way that shows which clades contain species of Gnidia.

    Open questions

    Rautenbach used different names from Herber for some of the groups and placed some of the groups at different taxonomic rank, but her phylogeny supports Herber's classification with the few exceptions noted below. The only strongly supported difference (99% (bootstrap percentage)
    Bootstrapping (statistics)
    In statistics, bootstrapping is a computer-based method for assigning measures of accuracy to sample estimates . This technique allows estimation of the sample distribution of almost any statistic using only very simple methods...

    from Herber's classification was that Dais was found to be sister to Phaleria. The phylogeny casts significant doubt upon the monophyly of the subfamily Octolepidoideae, and upon the monophyly of the informal Octolepis and Gonostylus groups, but this result had only weak statistical support. Only a sampling of more species and more DNA from each will determine whether these groups are monophyletic or not. Stephanodaphne and Peddiea might need to be transferred to the Gnidia group, but support was not strong (60% BP) for a clade consisting of the Gnidia group with Stephanodaphne and Peddiea. Again, more extensive sampling will be required to resolve this question. Two of the 3 genera placed incertae sedis by Herber (Linodendron and Lasiadenia) have not yet been sampled and their relationships to other genera remain obscure.

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