The Thymelaeaceae are a cosmopolitan

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

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 t ...

s composed of 50 genera (listed below) and 898 species.Zachary S. Rogers (2009 onwards)
A World Checklist of Thymelaeaceae (version 1)
Missouri Botanical Garden Website, St. Louis. It was established in 1789 by Antoine Laurent de Jussieu.Antoine Laurent de Jussieu ''Genera Plantarum'', page 76. Herrisant & Barrois, Paris. The Thymelaeaceae are mostly

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, including only woody plants with secondary growth, plants that are ...

s and

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from tree ...

s, with a few

vine A vine (Latin ''vīnea'' "grapevine", "vineyard", from ''vīnum'' "wine") is any plant with a growth habit of trailing or scandent (that is, climbing) stems, lianas or runners. The word ''vine'' can also refer to such stems or runners themsel ...

s and

herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition ...

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.Ernst Schmidt, Mervyn Lotter and Warren McCleland The number of

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 fila ...

s is usually once or twice the number of

calyx Calyx or calyce (plural "calyces"), from the Latin ''calix'' which itself comes from the Ancient Greek ''κάλυξ'' (''kálux'') meaning "husk" or "pod", may refer to: Biology * Calyx (anatomy), collective name for several cup-like structures ...

lobes. If twice, then they often occur in two 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

corolla Corolla may refer to: *Corolla (botany), the petals of a flower, considered as a unit *Toyota Corolla, an automobile model name *Corolla (headgear) A ''corolla'' is an ancient headdress in the form of a small circlet or crown.receptacle. This feature is probably unique to Thymelaeaceae. The

sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...

s are mounted on the rim of the floral tube. Stamens may be mounted on the rim or inside. What appear to be petals are actually stipular appendages of the sepals. The fruit is a 1-seeded

berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, rasp ...

or an

achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not o ...

. The 1-seeded berries have often been mistaken for drupes whenever the seed coat was mistaken for an endocarp. When using a key to the families of flowering plants, Thymelaeaceae are often difficult or impossible to recognize because of equivocal interpretation of the flower parts. Sepals, petals, and

staminode In botany, a staminode is an often rudimentary, sterile or abortive stamen, which means that it does not produce pollen.Jackson, Benjamin, Daydon; ''A Glossary of Botanic Terms with their Derivation and Accent''; Published by Gerald Duckworth & C ...

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.Warren L. Wagner, Derral R. Herbst, and Sy H. Sohmer. ''Manual of the Flowering Plants of Hawaii'', Revised Edition, 1999. Bishop Museum Press: Hololulu In these, the nonfunctional organs are much deformed and bear little resemblance to the parts that they represent. A good collection of pictures of plants in this family has appeared in a scientific paper.

Taxonomy

The family is named for the genus '' Thymelaea'', the name of which is a combination of the Greek name for the herb

thyme Thyme () is the herb (dried aerial parts) of some members of the genus ''Thymus'' of aromatic perennial evergreen herbs in the mint family Lamiaceae. Thymes are relatives of the oregano genus '' Origanum'', with both plants being mostly indigen ...

θύμος (thúmos) and that for the

olive The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ' ...

ἐλαία (elaía) - in reference to its thyme-like foliage (i.e. minuscule leaves) and olive-like fruit.

Classification

The Thymelaeaceae are in the order Malvales.Peter F. Stevens (2001 onwards) Angiosperm Phylogeny Website In: Missouri Botanical Garden Website Except for a sister relationship with Tepuianthaceae, little is known for sure about their relationships with the other families in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...

.Clemens Bayer, Michael F. Fay, Anette Y. de Bruijn, Vincent Savolainen, Cynthia M. Morton, Klaus Kubitzki, William S. Alverson, and Mark W. Chase (1999). "Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atpB and rbcL DNA sequences". ''Botanical Journal of the Linnean Society'' 129(4): 267-381 Unlike most recent authors, who recognize four subfamilies, B.E. Herber has divided Thymelaeaceae into two subfamilies. He has retained the subfamily Gonostyloideae, but renamed it Octolepidoideae. The other three traditional subfamilies (Synandrodaphnoideae, Aquilarioideae, and Thymelaeoideae) were combined into a Thymelaeoideae s.l.( sensu lato), and reduced to tribal rank, as Synandrodaphneae, Aquilarieae, and Daphneae, respectively. No tribes were designated in subfamily Octolepidoideae, but it was provisionally divided into two informal groups, the Octolepis group and the Gonystylus group. Likewise, no

subtribes Subtribe is a taxonomic category ranking which is below the rank of tribe and above genus. The standard suffix for a subtribe is -ina (in animals) or -inae (in plants Plants are predominantly photosynthetic eukaryotes of the kingdom Plant ...

were designated in the tribe Daphneae, but it was informally divided into four 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 (not assigned to any particular group or in a separate group by themselves).

Octolepidoideae

:''Octolepis'' group: '' Arnhemia'', ''

Deltaria ''Deltaria brachyblastophora'' is a species of shrubs in the Thymelaeaceae family. It is endemic to New Caledonia and the only species of the genus ''Deltaria''. It is related to '' Arnhemia'', ''Gonystylus'', ''Lethedon'' and ''Solmsia ''Solm ...

'', ''

Lethedon ''Lethedon'' is a genus of shrubs in the Thymelaeaceae family from Australia and New Caledonia. It is related to '' Arnhemia'', '' Deltaria'', ''Gonystylus'' and ''Solmsia ''Solmsia'' is a genus containing one or two species of flowering pla ...

'', '' Octolepis'', ''

Solmsia ''Solmsia'' is a genus containing one or two species of flowering plants belonging to the family Thymelaeaceae. It is endemic to New Caledonia. The genus was named to honor Hermann zu Solms-Laubach by Henri Ernest Baillon. It is related to '' ...

'' :''Gonystylus'' group: '' Aetoxylon'', '' Amyxa'', ''

Gonystylus ''Gonystylus'' is a southeast Asian genus of about 30 species of hardwood trees also known as ramin, melawis ( Malay) and ramin telur (Sarawak). Description Ramin is native to Malaysia, Singapore, Indonesia, Brunei, the Philippines, and Papua ...

Thymelaeoideae

:Synandrodaphneae: '' Synandrodaphne'' :Aquilarieae: '' Aquilaria'', '' Gyrinops'' :Daphneae ::''Linostoma'' group: '' Craterosiphon'', '' Dicranolepis'', '' Enkleia'', '' Jedda'', '' Linostoma'', ''

Lophostoma ''Lophostoma'' is a genus of Central and South American bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true ...

'', '' Synaptolepis'' ::''Phaleria'' group: '' Peddiea'', '' Phaleria'' ::''Daphne'' group: '' Daphne'', ''

Daphnopsis ''Daphnopsis'' is a plant genus in the family Thymelaeaceae The Thymelaeaceae are a cosmopolitan family of flowering plants composed of 50 genera (listed below) and 898 species.Zachary S. Rogers (2009 onwards)A World Checklist of Thymelaeace ...

'', '' Diarthron'', ''

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; other ...

'', ''

Edgeworthia ''Edgeworthia'' (paper bush) is a genus of plants in the family Thymelaeaceae. When the genus was first described, it was published twice in the same year (1841), in two separate publications: '; and ''Denkschriften der Regensburgischen Botanisch ...

'', '' Funifera'', '' Goodallia'', '' Lagetta'', '' Ovidia'', '' Rhamnoneuron'', '' Schoenobiblus'', ''

Stellera ''Stellera'' is a genus of flowering plant in the family Thymelaeaceae, with a single species ''Stellera chamaejasme'' found in mountainous regions of Central Asia, China, Siberia and South Asia. ''S. chamaejasme'' is a herbaceous perennial ...

'', '' Thymelaea'', ''

Wikstroemia ''Wikstroemia'' is a genus of 55-70 species of flowering shrubs and small trees in the mezereon family, Thymelaeaceae. Hawaiian species are known by the common name ‘ākia. Medicinal uses '' Wikstroemia indica'' () is one of the 50 fundamental ...

'' ::''Gnidia'' group: ''

Dais A dais or daïs ( or , American English also but sometimes considered nonstandard)dais
in the Random House Dictionary< ...

'', '' Drapetes'', '' Gnidia'', '' Kelleria'', '' Lachnaea'', '' Passerina'', ''

Pimelea ''Pimelea'', commonly known as rice flowers, is a genus of plants belonging to the family Thymelaeaceae. There are about 150 species, including 110 in Australia and thirty six in New Zealand. Description Plants in the genus ''Pimelea'' are h ...

'', '' 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 A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it i ...

DNA. These were the rbcL

gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...

and the intergenic spacer between the transfer RNA 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) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...

) 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) phylogeny. 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''.

Defining the genera

Ruby Gnidia (Gnidia rubescens) close-up (11421758284)

The circumscription 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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...

tree below). A recent comparison of DNA sequences has established the

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...

of ''Thymelaea'' and the polyphyly of ''Diarthron'',David Galicia-Herbada (2006). "Origin and diversification of Thymelaea(Thymelaeaceae): inferences from a phylogenetic study based on ITS (rDNA) sequences". ''Plant Systematics and Evolution'' 257(3-4):159-187. 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 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 specim ...

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''. 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)Michelle van der Bank, Michael F. Fay, and Mark W. Chase (2002). "Molecular Phylogenetics of Thymelaeaceae with particular reference to African and Australian genera". ''Taxon'' 51(2):329-339. suggested that ''Lasiosiphon'' might be resurrected if redefined. The type species for ''Lasiosiphon'' is ''Gnidia glauca'', formerly known as ''Lasiosiphon glaucus''.

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) 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.

Genera

Herber (2003)B.E. Herber. "Thymelaeaceae" In: ''The Families and Genera of Vascular Plants vol.V'' (Klaus Kubitzki and Clemens Bayer, volume editors). Springer-Verlag: Berlin, Heidelberg (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). , 50+ genera are accepted by the World Checklist of Thymelaceae: In the past, different authors have defined Thymelaeaceae in different ways. For example, John Hutchinson excluded ''Gonystylus'' and its close relatives, as well as ''Aquilaria'' and its close relatives from the family, forming 2 segregate families: Gonystylaceae and Aquilariaceae.Hutchinson,John. ''The Families of Flowering Plants, Third Edition (1973)''. Oxford University Press: London. 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 In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

family.Horn,J.W., (2004). "The morphology and relationships of the Sphaerosepalaceae(Malvales)". ''Botanical Journal of the Linnean Society'' 144(1):1-40 Stevens includes ''Tepuianthus'', but Kubitzki treats Tepuianthaceae as a separate family.Klaus Kubitzki. "Tepuianthaceae" In: ''The Families and Genera of Vascular Plants, vol.V'' Klaus Kubitzki and Clemens Bayer. (volume editors). Springer-Verlag: Berlin,Heidelberg. (2003).

Distribution

The family is more diverse in the Southern Hemisphere than in the

Northern Northern may refer to the following: Geography * North, a point in direction * Northern Europe, the northern part or region of Europe * Northern Highland, a region of Wisconsin, United States * Northern Province, Sri Lanka * Northern Range, a r ...

, with major concentrations of species in

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

and

Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...

.Vernon H. Heywood, Richard K. Brummitt, Alastair Culham, and Ole Seberg. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada (2007) The genera are overwhelmingly African.Marline Rautenbach. Figure 1.2, page 7 In: "Gnidia is not monophyletic: taxonomic implications for Gnidia and its relatives in Thymelaeoideae" Digispace at the University of Johannesburg. 8 Jul 2008. (see External links below).

Ethnobotany and economic use

Several genera are of economic importance. ''

'' (Ramin) is valued for its comparatively soft, easily worked yellowish wood, but trade in all species in the genus are controlled by CITES. Many genera have inner bark yielding strong

fibre Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...

suitable for the making of cordage and

paper Paper is a thin sheet material produced by mechanically or chemically processing cellulose fibres derived from wood, rags, grasses or other vegetable sources in water, draining the water through fine mesh leaving the fibre evenly distribu ...

- a fact actually acknowledged in the naming of one of the genera, '' Funifera'' being the Latin for "bearer (provider) of rope". The barks of '' Daphne'', ''

'', '' Rhamnoneuron'', '' Thymelaea'', ''

'', and ''

'' are used in paper-making, while '' Lagetta'' species are known as lacebark for their lacelike inner bark, the attractive appearance of which has led to their being used to make clothing and other utilitarian objects.

Toxicity and medicinal uses

Many of the species (e.g. ''

Wikstroemia indica ''Wikstroemia indica'', also known as tie bush, Indian stringbush, bootlace bush, or small-leaf salago () is a small shrub with glossy leaves, small greenish-yellow flowers and toxic red fruits. It grows in forests and on rocky, shrubby slopes i ...

'' and '' Stellera chamaejasme'') have actual or potential uses in medicine and are

poison Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...

ous if eaten, acting as violent purges (e.g. '' Daphne mezereum''), this toxicity often being related to the plants' containing phorbol esters which, as the name suggests, are also common in the

spurge ''Euphorbia'' is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to t ...

family Euphorbiaceae.

Use as ornamental plants

'' Daphne'' is grown (despite the high toxicity of its attractive fruits) for its sweetly scented flowers. Species of ''

'', '' Daphne'', '' Phaleria'', ''

Dais A dais or daïs ( or , American English also but sometimes considered nonstandard)dais
in the Random House Dictionary< ...

'', ''

'' and other genera are grown as Ornamental plant, ornamentals.George W. Staples and Derral R. Herbst. 2005. "A Tropical Garden Flora". Bishop Museum Press: Honolulu. .Marline Rautenbach. "Gnidia is not monophyletic: taxonomic implications for Gnidia and its relatives in Thymelaeoideae". Digispace at the University of Johannesburg 8 Jul 2008. (see External links below)

Gallery

File:Strettle Road Reserve pimelea.jpg, Inflorescence of the

n ''Pimelea spectabilis''. File:Daphne striata 100604.jpg, ''Daphne striata'', native to the Alps and the Dolomites. File:Struthiola myrsinites Gonnabos IMG 1983s.jpg, The South African ''Struthiola myrsinites''. File:Passerina (Thymelaeaceae) details of flowering sprig EOS 009.jpg, Flowers of an unidentified ''Passerina'' species. File:Hortus Botanicus Leiden - Phaleria capitata Jack (Sumatra).JPG, ''Phaleria capitata'' of Sumatra exhibiting cauliflory. File:Stellera chamaejasme 2.jpg, '' Stellera chamaejasme'' of Central Asia, Central and East Asia. File:Daphnopsis racemosa Griseb. (12751985583).jpg, The Brazilian ''Daphnopsis racemosa'' in flower. File:Daphnopsis racemosa Griseb. (17144500699).jpg, Fruits of ''Daphnopsis racemosa''. File:Dirca palustris - Edwards.jpg, Flowers of ''Dirca palustris'' of the US File:Akabana-mitsumata.JPG, ''Edgeworthia chrysantha'', native to China. File:Wikstroemia phillyreifolia (Hawai'i false ohelo) flowers (6592426883).jpg, Flowers of the Hawaiian ''Wikstroemia phillyreifolia''.

References

External links

* Zachary S. Rogers (2009 onwards)
A Worldwide Checklist of Thymelaeaceae (version 1).

Peter F. Stevens (2001 onwards) In
Missouri Botanical Garden

Rautenbach(2008)
in
UJDigiSpace @ The University of Johannesburg

distribution
in: Gnidia is not monophyletic: taxonomic implications for Gnidia and its relatives in Thymelaeoideae
Thymelaeaceae of Mongolia in FloraGREIF
{{Taxonbar, from=Q156109 Thymelaeaceae, Malvales families