Thymelaeaceae

{{Short description|Family of flowering plants}} {{more footnotes needed|date=November 2017}} {{Automatic taxobox |image = Thymelaea hirsuta kz2.jpg |image_caption = ''Thymelaea hirsuta'' |taxon = Thymelaeaceae |authority = Juss. |subdivision_ranks = Genera |subdivision = See text }}

The '''Thymelaeaceae''' {{IPAc-en|ˌ|θ|ɪ|m|ᵻ|l|iː|ˈ|eɪ|s|iː|,_|-|s|i|%|aI}} are a cosmopolitan family of flowering plants composed of 50 genera (listed below) and 898 species.<ref name="Rogers2009">Zachary S. Rogers (2009 onwards). [http://www.tropicos.org/Project/Thymelaeaceae A World Checklist of Thymelaeaceae (version 1)]. Missouri Botanical Garden Website, St. Louis.</ref> It was established in 1789 by Antoine Laurent de Jussieu.<ref name="jussieu1789">Antoine Laurent de Jussieu ''Genera Plantarum'', page 76. Herrisant & Barrois, Paris.</ref> The Thymelaeaceae are mostly trees and shrubs, with a few vines and herbaceous plants, the latter including some annual species.

==Description== Several conspicuous or unusual traits are characteristic of the family (when ''Tepuianthus'' is excluded). The bark is usually shiny and fibrous, with strips of bark peeling down the side of broken stems.<ref name=Kruger>Ernst Schmidt, Mervyn Lotter and Warren McCleland {{google books|RpVJJH3kV0oC|Trees and Shrubs of Mpumalanga and Kruger National Park (2002)|page=448}}</ref> The number of stamens is usually once or twice the number of calyx lobes; when twice, 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.

Thymelaeaceae are often difficult to identify because of equivocal interpretation of the flower parts. Sepals, petals, and staminodes 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.

==Taxonomy== The family is named from the genus ''Thymelaea'', the name of which is a combination of the Greek name for the herb thyme θύμος (thúmos) and that for the olive ἐλαία (elaía)—in reference to its thyme-like foliage (i.e. minuscule leaves) and olive-like fruit.<ref>''The Royal Horticultural Society Dictionary of Gardening'' ed. Chittenden, Fred J., 2nd edition, by Synge, Patrick M. Volume III : Je-Pt. Pub. Oxford at the Clarendon Press 1965. Reprinted 1984. {{ISBN|0-19-869106-8}}</ref>

===Classification=== The Thymelaeaceae are in the order Malvales.<ref name="Steven2001">Peter F. Stevens (2001 onwards) Angiosperm Phylogeny Website In: Missouri Botanical Garden Website</ref> Except for a sister relationship with Tepuianthaceae, little is known for sure about their relationships with the other families in the order.<ref name="Bayer1999">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</ref>

Unlike most recent authors, who accept four subfamilies, B. E. Herber has divided Thymelaeaceae into two subfamilies.{{cn|date=August 2025}} 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 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 accepted 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'', ''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: ''Peddiea'', ''Phaleria'' ::'''''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.<ref name=vanderbank2002/> It was based on two regions of chloroplast DNA. These were the rbcL gene 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) 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''.

{{clade | style=font-size:75%;line-height:100% |1={{clade |1=''Octolepis'' }} |2={{clade |1=''Gonystylus'' |2=''Lethedon'' |3=''Deltaria'' |4=''Solmsia'' |5=''Arnhemia'' }} |label3=Thymelaeoideae |3={{clade |label1=Synandrodaphneae |1=''Synandrodaphne'' |2={{clade |label1=Aquilarieae |1={{clade |1=''Aquilaria'' |2=''Gyrinops'' }} |label2=Daphneae |2={{clade |label1=Linostoma group |1={{clade |1=''Dicranolepis'' |2={{clade |1=''Enkleia'' |2={{clade |1=''Synaptolepis'' |2=''Craterosiphon'' }} }} }} |2={{clade |label1=Daphne group |1={{clade |1=''Edgeworthia'' |2={{clade |1={{clade |1=''Wikstroemia gemmata'' |2={{clade |1=''Wikstroemia canescens'' |2=''Stellera'' }} }} |2={{clade |1=''Diarthron'' |2={{clade |1=''Daphne'' |2=''Thymelaea'' }} }} }} }} |2={{clade |label1=Phaleria group |1={{clade |1=''Phaleria'' |2=''Dais'' }} |label2=Gnidia group |2={{clade |1={{clade |1=''Gnidia glauca'' |2={{clade |1=''Gnidia kraussiana'' |2=''Gnidia calocephala'' }} }} |2=''Stephanodaphne'' |3={{clade |1=''Dirca'' |2=''Ovidia'' }} |4=''Peddiea'' |5={{clade |1=''Gnidia humilis'' |2={{clade |1=''Gnidia squarrosa'' |2={{clade |1=''Gnidia pilosa'' |2=''Pimelea'' }} }} }} |6={{clade |1={{clade |1=''Gnidia racemosa'' |2={{clade |1=''Gnidia pinifolia'' |2=''Struthiola'' }} }} |2={{clade |1=''Lachnaea'' |2=''Passerina'' |3={{clade |1=''Drapetes'' |2=''Gnidia anomala'' }} }} }} }} }} }} }} }} }} }}

===Defining the genera=== [[File:Ruby Gnidia (Gnidia rubescens) close-up (11421758284).jpg|thumb|upright|left|The striking flowers of ''Gnidia rubescens'']] thumb|upright|left|''Gnidia pinifolia'' in bloom [[File:Gnidia glauca (6666724377).jpg|thumb|upright|right|''Gnidia glauca'' (known formerly as ''Lasiosiphon glaucus'')]] [[File:Nacchinar (in Tamil) (3127220503).jpg|thumb|upright|right|Detail of flowers of ''Gnidia glauca'' (known formerly as ''Lasiosiphon glaucus'')]] The circumscription of genera in Thymelaeaceae has always been difficult, and is to some degree artificial. For example, the difficulty of distinguishing ''Daphne'' from ''Wikstroemia'' has been commented upon by Rautenbach and Herber.<ref name="rautenbach2008"/><ref name="herber2003"/> 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 tree below).

A recent comparison of DNA sequences has established the monophyly of ''Thymelaea'' and the polyphyly of ''Diarthron'',<ref name="galicia-herbada2006">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.</ref> 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 four separate clades, each of which contains other genera of the family (see the phylogenetic tree below). The type species for ''Gnidia'' is ''Gnidia pinifolia''. If ''Gnidia'' is divided into four 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 treat ''Lasiosiphon'' as a separate genus from ''Gnidia''. This distinction was later shown to be artificial. However, Van der Bank et al. (2002)<ref name="vanderbank2002">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.</ref> suggested that ''Lasiosiphon'' might be resurrected if redefined. The type species for ''Lasiosiphon'' is ''Gnidia glauca'', formerly known as ''Lasiosiphon glaucus''.

===Open questions=== {{original research|subsection|date=November 2025}} 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 three genera placed incertae sedis by Herber (''Linodendron'' and ''Lasiadenia'') have not yet been sampled and their relationships to other genera remain obscure.

===Genera=== [[File:Gonystylus bancanus.jpg|thumb|upright|right|''Gonystylus bancanus'' native to Brunei, Indonesia, and Malaysia: botanical line drawing of detailed anatomy]] Herber (2003)<ref name="herber2003">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)</ref> accepts 45 genera, excluding ''Tepuianthus'' from the family, sinking ''Atemnosiphon'' and ''Englerodaphne'' into ''Gnidia'', ''Eriosolena'' into ''Daphne'', and ''Thecanthes'' into ''Pimelea''.<ref name="herber2003"/> 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).<ref name="herber2003"/>

{{As of|2024|September}}, ''Plants of the World Online'' accepts 52 genera.<ref name= powo>{{cite web |title=''Thymelaeaceae'' Juss. |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:50000375-1 |website=Plants of the World Online |publisher=Royal Botanic Gardens, Kew |access-date=16 September 2024}}</ref> {| border="0" |- | style="vertical-align:top;"| * ''Aetoxylon'' {{small|(Airy Shaw) Airy Shaw}} * ''Amyxa'' {{small|Tiegh.}} * ''Aquilaria'' {{small|Lam.}} * ''Arnhemia'' {{small|Airy Shaw}} * ''Atemnosiphon'' {{small|Leandri}} * ''Craterosiphon'' {{small|Engl. & Gilg.}} * ''Dais'' {{small|D.Royen ex L.}} * ''Daphne'' {{small|Tourn. ex L.}} * ''Daphnimorpha'' {{small|Nakai}} * ''Daphnopsis'' {{small|Mart.}} * ''Deltaria'' {{small|Steenis}} * ''Diarthron'' {{small|Turcz.}} * ''Dicranolepis'' {{small|Planch.}} * ''Dirca'' {{small|L.}} * ''Drapetes'' {{small|Banks ex Lam.}} * ''Edgeworthia'' {{small|Meisn.}} * ''Englerodaphne'' {{small|Gilg}} | style="vertical-align:top;"| * ''Enkleia'' {{small|Griff.}} * ''Eriosolena'' {{small|Blume}} * ''Funifera'' {{small|Andrews ex C.A.Mey.}} * ''Gnidia'' {{small|L.}} * ''Gonystylus'' {{small|Teijsm. & Binn.}} * ''Goodallia'' {{small|Benth.}} * ''Gyrinops'' {{small|Gaertn.}} * ''Jedda'' {{small|J.R.Clarkson}} * ''Kelleria'' {{small|Endl.}} * ''Lachnaea'' {{small|L.}} * ''Lagetta'' {{small|Juss.}} * ''Lasiadenia'' {{small|Benth.}} * ''Lethedon'' {{small|Biehler}} * ''Linodendron'' {{small|Griseb.}} * ''Linostoma'' {{small|Wall. ex Endl.}} * ''Lophostoma'' {{small|Meisn.}} * ''Octolepis'' {{small|Oliv.}} | style="vertical-align:top;"| * ''Ovidia'' {{small|Meisn.}} * ''Passerina'' {{small|L.}} * ''Peddiea'' {{small|Harv.}} * ''Phaleria'' {{small|Jack}} * ''Pimelea'' {{small|Banks ex Gaertn.}} * ''Restella'' {{small|Pobed.}} * ''Rhamnoneuron'' {{small|Gilg}} * ''Schoenobiblus'' {{small|Mart.}} * ''Solmsia'' {{small|Baill.}} * ''Stellera'' {{small|L.}} * ''Stephanodaphne'' {{small|Baill.}} * ''Struthiola'' {{small|L.}} * ''Synandrodaphne'' {{small|Gilg}} * ''Synaptolepis'' {{small|Oliv.}} * ''Tepuianthus'' {{small|Maguire & Steyerm.}} * ''Thymelaea'' {{small|Mill.}} * ''Wikstroemia'' {{small|Endl.}} |}

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 two segregate families, Gonystylaceae and Aquilariaceae.<ref name="hutchinson1973">Hutchinson, John. ''The Families of Flowering Plants, Third Edition (1973)''. Oxford University Press: London.</ref> 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.<ref name="horn2004">Horn, J.W., (2004). "The morphology and relationships of the Sphaerosepalaceae(Malvales)". ''Botanical Journal of the Linnean Society'' '''144'''(1):1-40</ref> Stevens includes ''Tepuianthus'', but Kubitzki treats Tepuianthaceae as a separate family.<ref name="kubitzky2003">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).</ref>

==Distribution== The family is more diverse in the Southern Hemisphere than in the Northern, with major concentrations of species in Africa and Australia.<ref name=heywood2007>Vernon H. Heywood, Richard K. Brummitt, Alastair Culham, and Ole Seberg. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada (2007)</ref> The genera are overwhelmingly African.<ref name="rautenbach page7">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).</ref>

==Ethnobotany and economic use== thumb|upright|''Lagetta lagetto'' the "lacebark": botanical illustration showing plant with samples of cordage and fabric made from its fibre [[File:Jamaican-lacebark-souvenirs2.jpg|thumb|upright|right|Intricate Jamaican souvenirs woven from "lacebark" fibre]] [[File:Flickr - BioDivLibrary - n39 w1150 (1).jpg|thumb|upright|right|The Brazilian ''Funifera utilis'', its genus named from the suitability of its fibre for rope-making (under the obsolete name ''Lagetta funifera'')]] Several genera are of economic importance. ''Gonystylus'' (ramin) is valued for its comparatively soft, easily worked yellowish wood, but trade in all species in the genus is controlled by CITES. Many genera have inner bark yielding strong fibre suitable for the making of cordage and paper, a fact acknowledged in the naming of one of the genera, ''Funifera'' meaning the Latin for "bearer (provider) of rope". The bark of ''Aquilaria'', ''Daphne'', ''Edgeworthia'', ''Gnidia'', ''Linostoma'', ''Rhamnoneuron'', ''Thymelaea'', ''Stellera'', and ''Wikstroemia'' are used in paper-making,<ref>Ojascastro, J., Phạm, V. P., Trần H. N., and Hart, R. (2024). Hand Papermaking Traditions of Việt Nam. Society of Ethnobiology. https://ethnobiology.org/publications/hand-papermaking-traditions-viet-nam</ref> while ''Lagetta'' was once harvested as a source of natural lace for making doilies and trimmings for luxury garments.<ref>Brennan, E., Harris, L. A., & Nesbitt, M. (2013). Object lesson Jamaican lace-bark: its history and uncertain future. Textile History, 44(2), 235-253. https://citeseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=39946822df30eeb0fcd93ce7d5a26b7a944cf6e7</ref>

==Toxicity and medicinal uses== [[File:Daphne mezereum 003.JPG|thumb|The attractive, but poisonous, fruit of ''Daphne mezereum'']] Many of the species (e.g. ''Wikstroemia indica'' and ''Stellera chamaejasme'') have actual or potential uses in medicine and are poisonous if eaten, acting as violent purges (e.g. ''Daphne mezereum''). This toxicity is often related to the plants' containing phorbol esters which, as the name suggests, are also common in the spurge family Euphorbiaceae.<ref>{{cite journal | pmid = 17661218| year = 2007| last1 = Goel| first1 = G| title = Phorbol esters: Structure, biological activity, and toxicity in animals| journal = International Journal of Toxicology| volume = 26| issue = 4| pages = 279–88| last2 = Makkar| first2 = H. P.| last3 = Francis| first3 = G| last4 = Becker| first4 = K| doi = 10.1080/10915810701464641| citeseerx = 10.1.1.320.6537| s2cid = 11550625}}</ref>

==Use as ornamental plants== [[File:Daphne bholua (3307537340).jpg|thumb|upright|right|The sweetly-scented and highly ornamental flowers of ''Daphne bholua'', a Nepalese species also used in traditional paper-making]] ''Daphne'' is grown (despite the high toxicity of its attractive fruit) for its sweetly scented flowers. Species of ''Wikstroemia'', ''Daphne'', ''Phaleria'', ''Dais'', ''Pimelea'' and other genera are grown as ornamental plants.<ref>George W. Staples and Derral R. Herbst. 2005. "A Tropical Garden Flora". Bishop Museum Press: Honolulu. {{ISBN|978-1-58178-039-0}}.</ref><ref name="rautenbach2008">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)</ref>

==Gallery== <gallery> File:Strettle Road Reserve pimelea.jpg|Inflorescence of the Australian ''Pimelea spectabilis'' File:Daphne striata 100604.jpg|''Daphne striata'', native to the Alps and the Dolomites File:Struthiola ciliata 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 and East Asia File:Daphnopsis racemosa Envira flor Oct 2007 031.jpg|Flower of ''Daphnopsis racemosa'' File:Daphnopsis racemosa Griseb. (17144500699).jpg|Fruit of ''Daphnopsis racemosa'' File:Dirca palustris - Edwards.jpg|Flowers of ''Dirca palustris'' from the United States 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'' </gallery>

==References== {{Reflist}}

== External links == {{Commons category|Thymelaeaceae}} * Zachary S. Rogers (2009 onwards). [http://legacy.tropicos.org/Project/Thymelaeaceae A Worldwide Checklist of Thymelaeaceae (version 1).] * [https://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny] Peter F. Stevens (2001 onwards) In: Missouri Botanical Garden * [https://ujcontent.uj.ac.za/esploro/outputs/doctoral/Gnidia-L-Thymelaeaceae-is-not-monophyletic/9913629807691 Rautenbach (2008)] in: [https://ujcontent.uj.ac.za/esploro/ UJDigiSpace @ The University of Johannesburg] * [https://web.archive.org/web/20110831201241/http://ujdigispace.uj.ac.za:8080/dspace/bitstream/10210/784/27/Bl.%207%20_Fig.%201.2%20-%20Distribution%20Thymelaeoideae.pdf Distribution] in: Gnidia is not monophyletic: taxonomic implications for Gnidia and its relatives in Thymelaeoideae *[https://floragreif.uni-greifswald.de/taxon/?flora_search=Taxon&fam_id=119 Thymelaeaceae of Mongolia in FloraGREIF]

{{Angiosperm families}} {{Taxonbar|from=Q156109}} {{Authority control}}

Category:Thymelaeaceae Category:Malvales families