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Classification System: APG IV

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

Familia: Rubiaceae
Subfamilia: Rubioideae
Tribus: Spermacoceae
Genera: AgathisanthemumAmphiasmaAmphistemonAnthospermopsisArcytophyllumAstiellaBorreriaBouvardiaCarajasiaCarterellaConostomiumCordylostigmaCruseaDebiaDenscantiaDentellaDiacrodonDibrachionostylusDimetiaDiodiaDolichocarpaDolichometraEdrastimaEmmeorhizaErnodeaExallageGaliantheGomphocalyxHedyotisHedythyrsusHemidiodiaHexasepalumHoustoniaHydrophylaxInvolucrellaKaduaKohautiaLathraeocarpaLelyaLeptopetalumLeptoscelaLucyaManettiaManostachyaMartensianthusMerumeaMexotisMicrasepalumMitracarpusMitrasacmopsisNeanotisNesohedyotisNodocarpaeaOldenlandia – Oldenlandiopsis – Opercularia – Parainvolucrella – ParanotisPentanopsisPentodonPhialiphora – Phyllocrater – Phylohydrax – Planaltina – Polyura – Pseudonesohedyotis – Psyllocarpus – Richardia – Sacosperma – Schwendenera – Scleromitrion – Spermacoce – Staelia – Stenaria – Stenotis – Stephanococcus – SynaptanthaTessieraThamnoldenlandiaTobagoaTortuella

Name

Spermacoceae Bercht. & J.Presl, Prir. Rostlin 256. (1820)

Type genus: Spermacoce L., Sp. Pl. 1: 102. (1753)

Synonyms

Homotypic
Spermacoceaceae Bercht. & J.Presl, Přir. Rostlin: 256. 1820.
Spermacoceae Cham. & Schltdl. ex DC., Prodr. 4: 538. 1830.
Spermacocineae J.Presl, Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 835. 1846.
Spermacocoideae Chevall., Fl. Gén. Env. Paris 2: 605. 1828.

Heterotypic
Hedyotidaceae Dumort., Comment. Bot. 57. 1822.
Hedyotideae Cham. & Schltdl. ex DC., Prodr. 4: 342, 401. 1830.
Hedyotidinae DC., Prodr. 4: 342, 419. 1830.
Hedyotidineae J.Presl, Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 812. 1846.
Hedyotidoideae Burnett, Outlines Bot. 906, 909. 1835.
Type genus: Hedyotis L., nom. cons.

Note: There is still uncertainty regarding the generic circumscription within this tribe - see Discussion Page for examples.
References

Berchtold, F.W.v. & Presl, J.S. 1820. O Prirozenosti Rostlin 256.
Bremer, B. & Manen, J.F. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Plant Systematics and Evolution 225(1-4): 43-72. DOI: 10.1007/BF00985458 PDF Reference page.
Fader, A.A.C., Salas, R.M., Dessein, S. & Cabral, E.L., 2016. Synopsis of Hexasepalum (Rubiaceae), the priority name for Diodella and a new species from Brazil. Systematic Botany 41(2): 408–422. DOI: 10.1600/036364416X691803 PDF from ResearchGate Reference page.
Gibbons, K.L. 2020. Hedyotis, Oldenlandia and related genera (Rubiaceae: Spermacoceae) in Australia: New genera and new combinations in an Asian‐Australian‐Pacific lineage. Taxon. The Journal of the International Association for Plant Taxonomy 69(3): 515-542. DOI: 10.1002/tax.12236 Paywall Reference page.
Groeninckx, I., Dessein, S., Ochoterena, H., Persson, C., Motley, T.J., Kårehed, J., Bremer, B., Huysmans, S. & Smets, E. 2009. Phylogeny of the Herbaceous Tribe Spermacoceae (Rubiaceae) Based on Plastid DNA Data1. Annals of the Missouri Botanical Garden 96(1): 109-133. DOI: 10.3417/2006201 PDF Reference page.
Groeninckx, I., Ochoterena, H., Smets, E. & Dessein, S. 2010. Molecular phylogenetic and morphological study of Kohautia (Spermacoceae, Rubiaceae), with the recognition of the new genus Cordylostigma. Taxon 59(5): 1457-1471. DOI: 10.1002/tax.595011 Paywall ResearchGate Reference page.
Guo, X., Wang, R.J., Simmons, M.P., But, P.P.H. & Yu, J. 2013. Phylogeny of the Asian Hedyotis–Oldenlandia complex (Spermacoceae, Rubiaceae): Evidence for high levels of polyphyly and the parallel evolution of diplophragmous capsules. Molecular Phylogenetics and Evolution 67(1): 110-122. DOI: 10.1016/j.ympev.2013.01.006 PDF Reference page.
Janssens, S.B., Groeninckx, I., Petra, J., Verstraete, B., Smets, E.F. & Dessein, S. 2016. Dispersing towards Madagascar: Biogeography and evolution of the Madagascan endemics of the Spermacoceae tribe (Rubiaceae). Molecular Phylogenetics and Evolution 95: 58-66. DOI: 10.1016/j.ympev.2015.10.024 Reference page.
Neupane, S., Dessein, S., Wikström, N., Lewis, P.O., Long, C., Bremer, B. & Motley, T.J. 2015. The Hedyotis-Oldenlandia complex (Rubiaceae: Spermacoceae) in Asia and the Pacific: Phylogeny revisited with new generic delimitations. Taxon 64(2): 299-322. DOI: 10.12705/642.8 PDF. Reference page.
Rydin, C., Razafimandimbison, S.G., Khodabandeh, A. & Bremer, B. 2009. Evolutionary relationships in the Spermacoceae alliance (Rubiaceae) using information from six molecular loci: insights into systematic affinities of Neohymenopogon and Mouretia. Taxon 58(3): 793-810. JSTOR PDF Reference page.
Xu, Y.D., Yuan, M.D. & Wang, R.J. 2021. Morphology and molecules support the new monotypic genus Parainvolucrella (Rubiaceae) from Asia. PhytoKeys 180: 53-64. DOI: 10.3897/phytokeys.180.67624 Open access Reference page.

Spermacoceae is a tribe of flowering plants in the family Rubiaceae and contains about 1346 species in 57 genera. Its representatives are found in the tropics and subtropics.[1]
Genera

Currently accepted names[1][2][3][4][5][6][7]

Agathisanthemum Klotzsch (5 sp.)
Amphiasma Bremek. (7 sp.)
Amphistemon Groeninckx (2 sp.)
Anthospermopsis (K.Schum.) J.H.Kirkbr. (1 sp.)
Arcytophyllum Willd. (18 sp.)
Astiella Jovet (1 sp.)
Bouvardia Salisb. (51 sp.)
Carajasia R.M.Salas, E.L.Cabral & Dessein (1 sp.)
Carterella Terrell (1 sp.)
Conostomium (Stapf) Cufod. (5 sp.)
Cordylostigma Groeninckx (9 sp.)
Crusea Schltdl. & Cham. (15 sp.)
Debia Neupane & N.Wikstr. (4 sp.)
Dentella J.R.Frost. & G.Frost. (8 sp.)
Dibrachionostylus Bremek. (1 sp.)
Dimetia (Wight & Arn.) Meisn. (6 sp.)
Diodella Small (12 sp.)
Diodia L. (23 sp.)
Edrastima Raf. (5 sp.)
Emmeorhiza Pohl ex Endl. (1 sp.)
Ernodea Sw. (8 sp.)
Exallage Bremek. (18 sp.)
Galianthe Griseb. (48 sp.)
Gomphocalyx Baker (1 sp.)
Hedyotis L. (182 sp.)
Hedythyrsus Bremek. (3 sp.)
Houstonia L. (24 sp.)
Hydrophylax L.f. (1 sp.)
Involucrella (Benth. & Hook.f.) Neupane (2 sp.)
Kadua Cham. & Schltdl. (30 sp.)
Kohautia Cham. & Schltdl. (28 sp.)
Lathraeocarpa Bremek. (2 sp.)
Lelya Bremek. (1 sp.)
Leptopetalum Hook. & Arn. (8 sp.)
Manettia Mutis ex L. (125 sp.)
Manostachya Bremek. (3 sp.)
Mitracarpus Zucc. (65 sp.)
Mitrasacmopsis Jovet (1 sp.)
Neanotis W.H.Lewis (34 sp.)
Nesohedyotis (Hook.f.) Bremek. (1 sp.)
Oldenlandia L. (217 sp.)
Oldenlandiopsis Terrell & W.H.Lewis (1 sp.)
Pentanopsis Rendle (2 sp.)
Pentodon Hochst. (2 sp.)
Phylohydrax Puff (2 sp.)
Planaltina R.M.Salas & E.L.Cabral (3 sp.)
Psyllocarpus Mart. & Zucc. (9 sp.)
Richardia L. (16 sp.)
Sacosperma G.Taylor (2 sp.)
Schwendenera K.Schum. (1 sp.)
Spermacoce L. (286 sp.)
Staelia Cham. & Schltdl. (21 sp.)
Stenaria (Raf.) Terrell (6 sp.)
Stephanococcus Bremek. (1 sp.)
Synaptantha Hook.f. (2 sp.)
Tessiera DC. (2 sp.)
Thamnoldenlandia Groeninckx (1 sp.)

Synonyms

Adenothola Lem. = Manettia
Aeginetia Cav. = Bouvardia
Allaeophania Thwaites = Hedyotis
Anistelma Raf. = Hedyotis
Anotis DC. = Arcytophyllum
Arbulocarpus Tennant = Spermacoce
Bigelovia Spreng. = Spermacoce
Bigelowia DC. = Spermacoce
Borreria G.Mey. = Spermacoce
Chaenocarpus Neck. ex A.Juss. = Spermacoce
Chamisme Nieuwl. = Houstonia
Chenocarpus Neck. = Spermacoce
Conotrichia A.Rich. = Manettia
Cormylus Raf. = Oldenlandia
Covolia Neck. = Spermacoce
Dasycephala (DC.) Hook.f. = Spermacoce
Decapenta Raf. = Diodia
Dichrospermum Bremek. = Spermacoce
Diodioides Loefl. = Spermacoce
Dioneiodon Raf. = Diodia
Diphragmus C.Pres. = Tessiera
Diplophragma (Wight & Arn.) Meisn. = Hedyotis
Duvaucellia Bowdich = Kohautia
Dyctiospora Reinw. ex Korth. = Oldenlandia
Ebelia Rchb. = Diodia
Edrastenia Raf. = Edrastima
Eionitis Bremek. = Oldenlandia
Endlichera C.Presl = Emmeorhiza
Endolasia Turcz. = Manettia
Endopogon Raf. = Diodella
Ereicoctis (DC.) Kuntze = Arcytophyllum
Gerontogea Cham. & Schltdl. = Oldenlandia
Gonotheca Blume ex DC. = Leptopetalum
Gouldia A.Gray = Kadua
Gruhlmania Neck. = Spermacoce
Guagnebina Vell. = Manettia
Hemidiodia K.Schum. = Spermacoce
Heymia Dennst. = Dentella
Hypodematium A.Rich. = Spermacoce
Jurgensia Raf. = Spermacoce
Karamyschewia Fisch. & C.A.Mey. = Oldenlandia
Lippaya Endl. = Dentella
Listeria Neck. = Oldenlandia
Lygistum P.Browne = Manettia
Macrandria (Wight & Arn.) Meisn. = Hedyotis
Mallostoma H.Karst. = Arcytophyllum
Metabolos Blume = Hedyotis
Metabolus A.Rich. = Hedyotis
Mitratheca K.Schum. = Oldenlandia
Nacibea Aubl. = Manettia
Neosabicea Wernham = Manettia
Octodon Thonn. = Spermacoce
Panetos Raf. = Houstonia
Paragophyton K.Schum. = Spermacoce
Peltospermum Benth. = Sacosperma
Pleiocraterium Bremek. = Hedyotis
Plethyrsis Raf. = Richardia
Poederiopsis Rusby = Manettia
Poiretia J.F.Gmel. = Houstonia
Pseudrachicallis H.Karst. = Arcytophyllum
Pterostephus C.Pres. = Spermacoce
Richardsonia Kunth = Richardia
Sarissus Gaertn. = Hydrophylax
Schiedea Bartl. = Richardia
Schizangium Bartl. ex DC. = Mitracarpus
Sclerococcus Bartl. = Hedyotis
Scleromitrion (Wight & Arn.) Meisn. = Hedyotis
Spermacoceodes Kuntze = Spermacoce
Staurospermum Thonn. = Mitracarpus
Stelmanis Raf. = Oldenlandia
Stelmotis Raf. = Oldenlandia
Symphyllarion Gagnep. = Hedyotis
Tardavel Adans. = Spermacoce
Teinosolen Hook.f. = Arcytophyllum
Terrellianthus Borhidi = Arcytophyllum
Thecagonum Babu = Leptopetalum
Thecorchus Bremek. = Oldenlandia
Theyodis A.Rich. = Oldenlandia
Triodon DC. = Diodia
Vanessa Raf. = Manettia
Wiegmannnia Meyen = Kadua

References

"World Checklist of Rubiaceae". Retrieved 21 April 2016.
Bremer B, Manen JF (2000). "Phylogeny and classification of the subfamily Rubioideae (Rubiaceae)". Plant Systematics and Evolution. 225 (1–4): 43–72. doi:10.1007/BF00985458. S2CID 44614735.
Neupane S, Dessein S, Wikström N, Lewis PO, Long C, Bremer B, Motley TJ (2015). "The Hedyotis-Oldenlandia complex (Rubiaceae: Spermacoceae) in Asia and the Pacific: Phylogeny revisited with new generic delimitations". Taxon. 64 (2): 299–322. doi:10.12705/642.8.
Wikström N, Neupane S, Kårehed J, Motley TJ, Bremer B (2013). "Phylogeny of Hedyotis L. (Rubiaceae: Spermacoceae): Redefining a complex Asian-Pacific assemblage" (PDF). Taxon. 62 (2): 357–374. doi:10.12705/622.2.
Robbrecht E, Manen JF (2006). "The major evolutionary lineages of the coffee family (Rubiaceae, angiosperms). Combined analysis (nDNA and cpDNA) to infer the position of Coptosapelta and Luculia, and supertree construction based on rbcL, rps16, trnL-trnF and atpB-rbcL data. A new classification in two subfamilies, Cinchonoideae and Rubioideae". Systematic Geography of Plants. 76: 85–146.
Bremer B (2009). "A review of molecular phylogenetic studies of Rubiaceae". Annals of the Missouri Botanical Garden. 96: 4–26. doi:10.3417/2006197. S2CID 53378010.
Bremer B, Eriksson E (2009). "Time tree of Rubiaceae: phylogeny and dating the family, subfamilies, and tribes". International Journal of Plant Sciences. 170 (6): 766–793. doi:10.1086/599077. S2CID 49332892.

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