Acanthaceae

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Cladus: Lamiids
Ordo: Lamiales

Familia: Acanthaceae
Subfamiliae: Acanthoideae – Avicennioideae – Nelsonioideae – Thunbergioideae

Genera: Acanthopale – Acanthopsis – Acanthus – Achyrocalyx – Afrofittonia – Ambongia – Ancistranthus – Andrographis – Angkalanthus – Anisacanthus – Anisosepalum – Anisostachya – Anisotes – Anomacanthus – Aphanosperma – Aphelandra – Ascotheca – Asystasia – Avicennia – Aymoreana – Ballochia – Barleria – Barleriola – Blepharis – Borneacanthus – Boutonia – Brachystephanus – Bravaisia – Brillantaisia – Brunoniella – Calacanthus – Calycacanthus – Camarotea – Carlowrightia – Celerina – Cephalacanthus – Cephalophis – Chalarothyrsus – Chamaeranthemum – Champluviera – Chileranthemum – Chlamydacanthus – Chlamydocardia – Chorisochora – Chroesthes – Clinacanthus – Clistax – Codonacanthus – Conocalyx – Cosmianthemum – Crabbea – Crossandra – Crossandrella – Cyclacanthus – Cynarospermum – Cyphacanthus – Dasytropis – Dianthera – Diceratotheca – Dichazothece – Dicladanthera – Dicliptera – Dinteracanthus – Dischistocalyx – Dolichostachys – Duosperma – Dyschoriste – Ecbolium – Echinacanthus – Elytraria – Encephalosphaera – Eranthemum – Eremomastax – Filetia – Fittonia – Forcipella – Glossochilus – Graphandra – Graptophyllum – Gymnophragma – Gymnostachyum – Gypsacanthus – Haplanthodes – Haplanthus – Harpochilus – Hemigraphis – Henrya – Herpetacanthus – Heteradelphia – Holographis – Hoverdenia – Hulemacanthus – Hygrophila – Hypoestes – Ichtyostoma – Isoglossa – Isotheca – Jadunia – Justicia – Kalbreyeriella – Kenyacanthus – Kosmosiphon – Kudoacanthus – Lankesteria – Lasiocladus – Leandriella – Lepidagathis – Leptosiphonium – Liberatia – Linariantha – Louteridium – Mackaya – Marcania – Mcdadea – Megalochlamys – Megaskepasma – Melittacanthus – Mellera – Mendoncia – Metarungia – Mexacanthus – Mimulopsis – Mirandea – Monechma – Monothecium – Morsacanthus – Nelsonia – Neriacanthus – Neuracanthus – Nicoteba – Odontonema – Oplonia – Orophochilus – Pachystachys – Pararuellia – Pericalypta – Petalidium – Phaulopsis – Phialacanthus – Phlogacanthus – Physacanthus – Podorungia – Poikilacanthus – Polylychnis – Populina – Pranceacanthus – Pseuderanthemum – Pseudocalyx – Pseudodicliptera – Psilanthele – Ptyssiglottis – Pulchranthus – Rhaphidospora – Rhinacanthus – Rhombochlamys – Ritonia – Rostellularia – Ruellia – Ruelliopsis – Rungia – Ruspolia – Ruttya – Saintpauliopsis – Salpinctium – Salpixantha – Samuelssonia – Sanchezia – Sapphoa – Satanocrater – Schaueria – Schaueriopsis – Sclerochiton – Sebastiano-schaueria – Spathacanthus – Sphacanthus – Sphinctacanthus – Staurogyne – Stenandrium – Stenostephanus – Stenothyrsus – Streblacanthus – Streptosiphon – Strobilanthes – Strobilanthopsis – Suessenguthia – Symplectochilus – Tabascina – Tessmanniacanthus – Tetramerium – Thunbergia – Thyrsacanthus – Thysanostigma – Trichanthera – Trichaulax – Trichocalyx – Trichosanchezia – Vavara – Vindasia – Whitfieldia – Wuacanthus – Xantheranthemum – Xerothamnella – Xylacanthus – Yeatesia – Zygoruellia
Paleogenera: †Kapgateophyllum
Genera of uncertain synonymy: Cuenotia – Pseudacanthopale – Pseudosiphonium – Psiloesthes – Stachyacanthus
Name

Acanthaceae Juss., Gen. Pl. 102 (1789), nom. cons. emend. Manzitto‐Tripp et al. (2021)

Type genus: Acanthus L. Sp. Pl. 2: 639. (1753)

Synonyms

Heterotypic
Avicenniaceae Miq., Pl. Preiss. (J.G.C. Lehmann) 1: 353 (1845)
Justiciaceae Raf., Fl. Tellur. 4: 60 (1838)
Thunbergiaceae Lilja, Skånes Fl., ed. 2: 979 (1870)
Mendonciaceae Bremek., Proc. Kon. Ned. Akad. Wetensch., ser. C, 56: 540 (1954)
Meyeniaceae Sreem., Phytologia 37: 412 (1977)
Nelsoniaceae Sreem., Phytologia 37: 412 (1977)

References
Primary references

Jussieu, A.L. de 1789. Genera plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. 498 pp. Paris: Herissant et Theophile Barrois. BHL Reference page. 102–103

Additional references

Darbyshire, I., Kiel, C.A., Daniel, T.F., McDade, L.A. & Luke, W.Q. 2019. Two new genera of Acanthaceae from tropical Africa. Kew Bulletin 74(3): 39. DOI: 10.1007/s12225-019-9828-z Reference page.
Manzitto‐Tripp, E.A., Darbyshire, I., Daniel, T.F., Kiel, C.A. & McDade, L.A. 2021. Revised classification of Acanthaceae and worldwide dichotomous keys. Taxon. DOI: 10.1002/tax.12600 Open access Reference page.
McDade, L.A., Daniel, T.F. & Kiel, C.A. 2008. Toward a comprehensive understanding of phylogenetic relationships among lineages of Acanthaceae s.l. (Lamiales). American Journal of Botany 95(9): 1136–1152. DOI: 10.3732/ajb.0800096 Reference page.
McDade, L., Hammel, B. & Kiel, C. 2018. New Species, New Combinations and New Synonymies Towards a Treatment of Acanthaceae for the Manual de Plantas de Costa Rica. Aliso: A Journal of Systematic and Evolutionary Botany 36(1): 27-45. DOI: 10.5642/aliso.20183601.04 PDF Reference page.
Olmstead, R.G. (ed.) 2016. A Synoptical Classification of the Lamiales. Version 2.6.2 (in prog.). 20 pp. PDF (Internet Archive) Reference page.
Tripp, E.A. 2007. Evolutionary Relationships Within the Species-Rich Genus Ruellia (Acanthaceae). Systematic Botany 32(3): 628–649. DOI: 10.1600/036364407782250625 Paywall JSTOR Hybrid open access journal Reference page.

Links

Govaerts, R. et al. 2022. Acanthaceae in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2022 Jan. 21. Reference page.
Hassler, M. 2022. Acanthaceae. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2022. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: 2022 Jan. 21. Reference page.
International Plant Names Index. 2022. Acanthaceae. Published online. Accessed: Jan. 24 2022.
Profice, S.R., Kameyama, C., Côrtes, A.L.A., Braz, D.M., Indriunas, A., Vilar, T., Pessoa, C., Ezcurra, C., & Wasshausen, D. 2014. Acanthaceae, in Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Online.
Tropicos.org 2022. Acanthaceae. Missouri Botanical Garden. Published online. Accessed: 1 Jan. 2022.

Vernacular names
العربية: أقنثية
azərbaycanca: Akantkimilər
català: Acantàcies
čeština: Paznehtníkovité
dansk: Akantus-familien
Deutsch: Akanthusgewächse
English: Acanthus family
Esperanto: Akantacoj
español: Acantáceas
فارسی: پاخرسیان
suomi: Akanttikasvit
עברית: קוציציים
hrvatski: Primogovke
hornjoserbsce: Akantusowe rostliny
magyar: Medvekörömfélék
հայերեն: Ականթազգիներ
italiano: Acantacee
日本語: キツネノマゴ科
перем коми: Акант котыр
한국어: 쥐꼬리망초과
kurdî: Famîleya pelkerengan
коми: Акант котыр
lietuvių: Akantiniai
മലയാളം: അക്കാന്തേസീ
नेपाली: असुरो
Nederlands: Acanthusachtigen
norsk: Akantusfamilien
polski: Akantowate
română: Acantacee
русский: Акантовые
svenska: Akantusväxter
தமிழ்: முண்மூலிகைக் குடும்பம்
తెలుగు: అకాంథేసి
ไทย: วงศ์เหงือกปลาหมอ
Türkçe: Ayıpençesigiller
українська: Акантові
Tiếng Việt: Họ Ô rô
中文: 爵床科

Acanthaceae is a family (the acanthus family) of dicotyledonous flowering plants containing almost 250 genera and about 2500 species. Most are tropical herbs, shrubs, or twining vines; some are epiphytes. Only a few species are distributed in temperate regions. The four main centres of distribution are Indonesia and Malaysia, Africa, Brazil, and Central America. Representatives of the family can be found in nearly every habitat, including dense or open forests, scrublands, wet fields and valleys, sea coast and marine areas, swamps, and mangrove forests.

Description

Plants in this family have simple, opposite, decussated leaves with entire (or sometimes toothed, lobed, or spiny) margins, and without stipules. The leaves may contain cystoliths, calcium carbonate concretions, seen as streaks on the surface.

The flowers are perfect, zygomorphic to nearly actinomorphic, and arranged in an inflorescence that is either a spike, raceme, or cyme. Typically, a colorful bract subtends each flower; in some species, the bract is large and showy. The calyx usually has four or five lobes; the corolla tubular, two-lipped or five-lobed; stamens number either two or four, arranged in pairs and inserted on the corolla, and the ovary is superior and bicarpellated, with axile placentation.

The fruit is a two-celled capsule, dehiscing somewhat explosively. In most species, the seeds are attached to a small, hooked stalk (a modified funiculus called a jaculator or a retinaculum) that ejects them from the capsule. This trait is shared by all members of the clade Acanthoideae. A 1995 study of seed expulsion in Acanthaceae used high speed video pictures to show that retinacula propel seeds away from the parent plant when the fruits dehisce, thereby helping the plant gain maximum seed dispersal range.[3]

A species well known to temperate gardeners is bear's breeches (Acanthus mollis), a herbaceous perennial plant with big leaves and flower spikes up to 2 m tall. Tropical genera familiar to gardeners include Thunbergia and Justicia.

Avicennia, a genus of mangrove trees, usually placed in Verbenaceae or in its own family, Avicenniaceae, is included in Acanthaceae by the Angiosperm Phylogeny Group on the basis of molecular phylogenetic studies that show it to be associated with this family.
Medicinal uses

Traditionally the most important part use in Acanthaceae is the leaves and they are used externally for wounds. Some research has indicated that Acanthaceae possess antifungal, cytotoxic, anti-inflammatory, anti-pyretic, antioxidant, insecticidal, hepatoprotective, immunomodulatory, anti-platelet aggregation and anti-viral potential.[4]

For instance, Acanthus ilicifolius, whose chemical composition has been heavily researched, is widely used in ethnopharmaceutical applications, including in Indian and Chinese traditional medicine.[5] Various parts of Acanthus ilicifolius have been used to treat asthma, diabetes, leprosy, hepatitis, snake bites, and rheumatoid arthritis.[6] The leaves of Acanthus ebracteatus, noted for their antioxidant properties, are used for making Thai herbal tea in Thailand and Indonesia.[7]
Phytochemistry

Phytochemical reports on family Acanthaceae are glycosides, flavonoids, benzonoids, phenolic compounds, naphthoquinone and triterpenoids.[4]
Overview of systematics

Since the first comprehensive classification of Acanthaceae in 1847 by Nees,[8] there have been a few major revisions presented since for the whole family.

Lindau, in 1895, divided the family into the subfamilies Mendoncioideae, Thunbergioideae, Nelsonioideae, and Acanthoideae.[9] Critically, Mendoncioideae, Thunbergioideae, and Nelsonioideae do not possess retinaculate fruits—and it is this distinction, between classifying Acanthaceae into a family that includes those clades with non-retinaculate fruits and one that excludes them, that still persists to the modern day.

Bremekamp, in 1965, presented a classification of Acanthaceae that differed from that of Lindau, for his Acanthaceae excluded genera that lack retinaculate fruits.[10] He placed Nelsonioideae within Scrophulariaceae, classified Thunbergiaceae and Mendonciaceae as distinct families and divided his Acanthaceae into two groups (Acanthoideae and Ruelloideae) based on the presence or absence of cystoliths, articulate stems, monothecate anthers, and colpate pollen.

In Scotland and Vollesen's 2000 study,[11] they accepted 221 genera and detailed five major groups within Acanthaceae s.s. (that is, those possessing retinaculate fruits), which is equivalent to Acanthoideae Link sensu Lindau 1895. Out of those 221 genera, they placed 201 of them into seven infrafamilial taxa of Acanthaceae, leaving only 20 unplaced.

In the current understanding of Acanthaceae, Acanthaceae s.s. includes only those clades with retinaculate fruits (that is, Acantheae, Barlerieae, Andrographideae, Whitfieldeae, Ruellieae, and Justiceae), while Acanthaceae s.l. includes those clades as well as Thunbergioideae, Nelsonioideae, and Avicennia.[12]
Dating the Acanthaceae lineage

Much research, using both molecular data and fossils, has been conducted in recent years regarding the dating and distribution of the Acanthaceae and Lamiales lineage, although there still remains some ambiguity.

In a 2004 study on the molecular phylogenetic dating of asterid flowering plants, researchers estimated 106 million years (MY) for the stem lineage of Lamiales, 67 MY for the stem lineage of Acanthaceae, and 54 MY for the crown node of Acanthaceae (that is, the age of extant lineages with the family).[13] These estimates are older than those based on fossils that can confidently be assigned to Lamiales, which are middle Eocene in age, ca. 48-37 MY.[14] Palynomorphs that definitively show the existence of Acanthaceae are known from the upper Miocene, with the oldest ca. 22 MY.[15]
Selected genera
Chinese violet (Asystasia gangetica)
Barleria sp.
Leaf of the nerve plant (Fittonia verschaffeltii)
Polka dot plant (Hypoestes phyllostachya)
Justicia aurea
Louteridium panemensis
Peristrophe speciosa
Pseuderanthemum maculatum
Popping pod (Ruellia tuberosa )
Rostellularia sp.
Thunbergia laurifolia

As of 17 January 2017, the Germplasm Resources Information Network accepts 217 genera,[16] 206 Accepted Genera are listed by Plants of the World Online (the Royal Botanic Gardens, Kew).[17]

Acanthopale C.B.Clarke
Acanthopsis Harv.
Acanthostelma Bidgood & Brummitt
Acanthura Lindau
Acanthus L. (bear's breech)
Achyrocalyx Benoist
Aechmanthera
Adhatoda Mill. (sometimes included in Justicia)
Afrofittonia Lindau
Ambongia Benoist
Ancistranthus Lindau (desert honeysuckle)
Ancistrostylis T.Yamaz.
Andrographis Wall. ex Nees (false waterwillow)
Angkalanthus Balf.f.
Anisacanthus Nees
Anisosepalum E.Hossain
Anisostachya Nees (sometimes included in Justicia)
Anisotes Nees
Anomacanthus R.D.Good
Apassalus Kobuski
Aphanosperma T.F.Daniel
Aphelandra R.Br.
Aphelandrella Mildbr.
Ascotheca Heine
Asystasia Blume
Asystasiella Lindau
Avicennia L.
Ballochia Balf.f.
Barleria L. (bush violets)
Barleriola Oerst
Benoicanthus Heine & A.Raynal
Blechum P.Browne
Blepharis Juss.
Borneacanthus Bremek.
Boutonia DC.
Brachystephanus Nees
Bravaisia DC.
Brillantaisia P.Beauv.
Calacanthus T.Anderson ex Benth. & Hook.f.
Calophanoides (C.B.Clarke) Ridl. (sometimes included in Justicia)
Calycacanthus K.Schum.
Camarotea Scott-Elliot
Carlowrightia A.Gray (wrightwort)
Celerina Benoist
Cephalacanthus Lindau
Chaetacanthus (synonym of Dyschoriste) Nees
Chalarothyrsus Lindau
Chamaeranthemum Nees
Championella Bremek.
Chileranthemum Oerst.
Chlamydacanthus Lindau (sometimes included in Theileamea)
Chlamydocardia Lindau
Chlamydostachya Mildbr.
Chroesthes Benoist
Clinacanthus Nees
Clistax Mart.
Codonacanthus Nees
Conocalyx Benoist
Corymbostachys Lindau
Cosmianthemum Bremek.
Crabbea Harv.
Crossandra Salisb.
Crossandrella C.B.Clarke
Cyclacanthus S.Moore
Cylindrosolenium Lindau
Cyphacanthus Leonard
Dactylostegium Nees (sometimes included in Dicliptera)
Danguya Benoist
Dasytropis Urb.
Dichazothece Lindau
Dicladanthera F.Muell.
Dicliptera Juss. (Foldwing)
Didyplosandra Wight ex Bremek.
Dipteracanthus Nees (sometimes included in Ruellia)
Dischistocalyx T.Anderson ex Benth. & Hook.f.
Dolichostachys Benoist
Drejera Nees
Drejerella Lindau (sometimes included in Justicia)
Duosperma Dayton
Dyschoriste Nees (snakeherb)
Ecbolium Kurz
Echinacanthus Nees
Elytraria Michx. (scalystem)
Encephalosphaera Lindau
Epiclastopelma Lindau
Eranthemum L.
Eremomastax Lindau
Eusiphon Benoist (often included in Ruellia)
Filetia Miq.
Fittonia Coem.
Forcipella Baill.
Forsythiopsis Baker (sometimes included in Oplonia)
Geissomeria Lindl.
Glossochilus Nees
Golaea Chiov.
Graphandra J.B.Imlay
Graptophyllum Nees
Gymnophragma Lindau
Gymnostachyum Nees
Gynocraterium Bremek.
Gypsacanthus E.J.Lott et al.
Haplanthodes Kuntze
Harpochilus Nees
Hemiadelphis Nees
Hemigraphis Nees (sometimes included in Strobilanthes)
Henrya Nees
Herpetacanthus Nees
Heteradelphia Lindau
Holographis Nees
Hoverdenia Nees
Hulemacanthus S.Moore
Hygrophila R.Br. (swampweed)
Hypoestes (including Periestes)
Ionacanthus Benoist
Isoglossa Oerst.
Isotheca Turrill
Jadunia Lindau
Juruasia Lindau
Justicia L. (justicia, water-willow, shrimp plant)
Kalbreyeriella Lindau
Kosmosiphon Lindau
Kudoacanthus Hosok.
Lankesteria Lindl.
Lasiocladus Bojer ex Nees
Leandriella Benoist
Lepidagathis Willd.
Leptostachya Nees
Liberatia Rizzini (sometimes included in Lophostachys)
Linariantha B.L.Burtt & R.M.Sm.
Lophostachys Pohl
Louteridium S.Watson
Lychniothyrsus Lindau
Marcania J.B.Imlay
Megalochlamys Lindau
Megalostoma Leonard
Megaskepasma Lindau
Melittacanthus S.Moore
Mellera S.Moore
Mendoncia Vand.
Metarungia Baden
Mexacanthus T.F.Daniel
Meyenia Nees
Mimulopsis Schweinf.
Mirandea Rzed.
Monechma Hochst. (sometimes included in Justicia)
Monothecium Hochst.
Morsacanthus Rizzini
Nelsonia R.Br.
Neohallia Hemsl.
Neriacanthus Benth.
Neuracanthus Nees
Odontonema Nees (toothedthread)
Ophiorrhiziphyllon Kurz
Oplonia Raf.
Oreacanthus Benth.
Orophochilus Lindau
Pachystachys Nees
Pachystrobilus: P. involucratus (Blume) Bremek.[18]
Pelecostemon Leonard
Pentstemonacanthus Nees
Perenideboles Ram. (Goyena)
Pericalypta Benoist
Peristrophe Nees
Petalidium Nees
Phaulopsis Willd.
Phialacanthus Benth.
Phidiasia Urb.
Phlogacanthus Nees
Physacanthus Benth.
Podorungia Baill.
Poikilacanthus Lindau
Polylychnis Bremek. (often included in Ruellia)
Populina Baill.
Pranceacanthus Wassh.
Pseuderanthemum Radlk.
Pseudocalyx Radlk.
Pseudodicliptera Benoist
Pseudoruellia Benoist
Psilanthele Lindau
Ptyssiglottis T.Anderson
Pulchranthus V.M.Baum et al.
Pupilla Rizzini (sometimes included in Justicia)
Razisea Oerst.
Rhinacanthus Nees
Rhombochlamys Lindau
Ritonia Benoist
Rostellularia Rchb. (sometimes included in Justicia)
Ruellia L. (Wild Petunia)
Ruelliopsis C.B.Clarke
Rungia Nees
Ruspolia Lindau
Ruttya Harv.
Saintpauliopsis Staner (sometimes included in Staurogyne)
Salpinctium T.J.Edwards
Salpixantha Hook.
Samuelssonia Urb. & Ekman
Sanchezia Ruiz & Pav.
Santapaua N.P.Balakr. & Subr. (sometimes included in Hygrophila)
Sapphoa Urb.
Satanocrater Schweinf.
Sautiera Decne.
Schaueria Nees
Schwabea Endl. & Fenzl
Sclerochiton Harv.
Sebastiano-schaueria Nees
Sericospora Nees
Siphonoglossa Oerst. (tubetongue)
Spathacanthus Baill.
Sphacanthus Benoist
Sphinctacanthus Benth.
Spirostigma Nees
Standleyacanthus Leonard
Staurogyne Wall.
Steirosanchezia Lindau
Stenandriopsis S.Moore
Stenandrium Nees (shaggytuft)
Stenostephanus Nees
Streblacanthus Kuntze
Streptosiphon Mildbr.
Strobilanthes Blume
Strobilanthopsis S.Moore
Styasasia S.Moore
Suessenguthia Merxm.
Synchoriste Baill.
Taeniandra Bremek.
Tarphochlamys Bremek.
Teliostachya Nees
Tessmanniacanthus Mildbr.
Tetramerium Nees
Theileamea Baill.
Thunbergia Retz.
Thysanostigma J.B.Imlay
Tremacanthus S.Moore
Triaenanthus Nees
Trichanthera Kunth
Trichaulax Vollesen
Trichocalyx Balf.f.
Trichosanchezia Mildbr.
Ulleria Bremek. (often included in Ruellia)
Vavara Benoist
Vindasia Benoist
Warpuria Stapf
Whitfieldia Hook.
Xantheranthemum Lindau
Xerothamnella C.T.White
Xylacanthus Aver. & K.S.Nguyen
Yeatesia Small (Bractspike)
Zygoruellia Baill.[19]

Excluded genera

Thomandersia Baill. → Thomandersiaceae[20][21]

References

"Family: Acanthaceae Juss., nom. cons". Germplasm Resources Information Network. United States Department of Agriculture. 2003-01-17. Retrieved 2011-07-29.
Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
Witztum, A; Schulgasser, K (1995). "The mechanics of seed expulsion in Acanthaceae". Journal of Theoretical Biology. 176 (4): 531–542. Bibcode:1995JThBi.176..531W. doi:10.1006/jtbi.1995.0219.
Awan, A.J., Aslam, M.S. (2014). "FAMILY ACANTHACEAE AND GENUS APHELANDRA: ETHNOPHARMACOLOGICAL AND PHYTOCHEMICAL REVIEW". International Journal of Pharmacy and Pharmaceutical Sciences. 6 (10): 44–55.
Wostmann, R; Liebezeit, G (2008). "Chemical composition of the mangrove holly Acanthus ilicifolius (Acanthaceae)—review and additional data". Senckenbergiana Maritima. 38: 31–37. doi:10.1007/bf03043866. S2CID 38592501.
Bandaranayake, W. M. (1998). "Traditional and medicinal uses of mangroves". Mangroves and Salt Marshes. 2 (3): 133–148. doi:10.1023/a:1009988607044. S2CID 129317332.
Chan, E. W. C.; Eng, S. Y.; Tan, Y. P.; Wong, Z. C.; Lye, P. Y.; Tan, L. N. (2012). "Antioxidant and Sensory Properties of Thai Herbal Teas with Emphasis on Thunbergia laurifolia Lindl". Chiang Mai J. Sci. 39 (4): 599–609.
Nees, C. G. (1847). de Candolle, A. P. (ed.). "Acanthaceae". Prodromus Systematis Naturalis Regni Vegetabilis. 11: 46–519.
Lindau, G. (1895). Engler, A.; Prantl, K. (eds.). "Acanthaceae". Die Natuirlichen Pflanzenfamilien. 4 (3b): 274–353.
Bremekamp, C. E. B. (1965). "Delimitation and subdivision of the Acanthaceae". Bull. Bot. Surv. India. 7: 21–30.
Scotland, R. W.; Vollesen, K. (2000). "Classification of Acanthaceae". Kew Bulletin. 55 (3): 513–589. doi:10.2307/4118776. JSTOR 4118776.
Tripp, E. A.; Daniel, T. F.; Fatimah, S.; McDade, L. A. (2013). "Phylogenetic Relationships within Ruellieae (Acanthaceae) and a Revised Classification". International Journal of Plant Sciences. 174 (1): 97–137. doi:10.1086/668248. S2CID 84423889.
Bremer, K.; Friis, E. M.; Bremer, B. (2004). "Molecular phylogenetic dating of asterid flowering plants shows Early Cretaceous diversification". Systematic Biology. 53 (3): 496–505. doi:10.1080/10635150490445913. PMID 15503676.
Pigg, K. B.; Wehr, W. C. (2002). "Tertiary flowers, fruits, and seeds of Washington state and adjacent areas—Part III" (PDF). Washington Geology. 30: 3–16.
Medus, J. (1975). "Palynologie de sediments tertiaires de Sénégal mé ridional". Pollen et Spores. 17: 545–608.
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