Takhtajan Armen. Flowering Plants

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564 Subclass VIII. LAMIIDAE

more or less sessile, unequally

2-lobed stigma; ovary 1-locular,

superior, with free-central pla-

centa and usually anatropous,

usually numerous (two in

Utricularia subgenus Biovularia)

ovules. Fruits usually capsules

opening by 2–4 valves or irregu-

larly or circumscissile, in subge-

nus Biovularia of the genus

Utricularia indehiscent and

1-seeded. Seeds small, exotestal

cells variously thickened;

embryo green, scarcely differen-

tiated, endosperm absent.

Contain iridoid glycosids (glob-

ularin, globulariscin, scutellari-

osid, aucubin, etc.), ﬂ avones,

p-cumarid and rarely caffeic

acid, n = 7–16, 21–24, 32

(Pinguicula).. . . . . . . . . . . . . . . .

. . . . . . .22. lentibulariaceae.

1. BUDDLEJACEAE

K. Wilhelm 1910. 5/c.100. Tropical, subtropical, and

warm-temperate regions of Africa, America, and Asia.

Emorya (1) from Texas to northern Mexico; Nicodemia –

Madagascar.

Gomphostigma, Buddleja, Nicodemia, Emorya,

Chilianthus

Show many similarities with the Loganiaceae, but dif-

fer from them in the absence of true stipules (only “auric-

ulae”) and of intraxylary phloem, in glandular, stellate, or

lepidite indumentum, and chemically (Buddleja has been

reported to contain an iridoid group typical of Lamiales,

but not of Rubiales, see Jensen [2000]). In general the

Buddlejaceae are nearer to the Scrophulariaceae than to

the Rubiales (Wagenitz 1959; Takhtajan 1966, 1980,

1987, 1997; Punt and Leenhouts 1967; Cronquist 1981,

1988; Dahlgren 1980, 1983, 1989). This is supported by

serological studies (Piechura ﬁ de Cronquist 1981: 947),

embryology, anatomy, and molecular date.

2. POLYPREMACEAE

L. Watson ex Doweld et Reveal 2007. 1/1. Southern USA,

Mexico, Central and South America, West Indies.

Polypremum

Related to the Buddlejaceae, but differ in regard to

the absence of endosperm haustoria, early develop-

ment of the proembryo, and a chromosome number

(Wagstaff 2004).

3. TETRACHONDRACEAE

Wettstein 1924. 1/2. New Zealand and Patagonia and

Tierra del Fuego.

Tetrachondra

Tetrachondra is a reduced and rather isolated plant

with uncertain afﬁ nity.

4. CALCEOLARIACEAE

Olmstead 2001. 3/307. Tropical and Western temper-

ate South America, Brazil, also New Zealand (some

Jovellana).

Calceolaria, Jovellana, Stemotria (including

Porodittia)

Related to the Gesneriaceae (see Weber 1972, 1982,

Andersson and Molau 1980).

5. STILBACEAE

Kunth 1831 (including Retziaceae Bartling 1830).

7/28. Cape Province of South Africa; Nuxia (15):

southern Arabia to tropical Africa, Mascarenes, and

South Africa.

5.1 NUXIOIDEAE

Stamens four. Shrubs or trees. Leaves opposite, rarely

alternate, with stipular ring. Flowers 4-merous, in thyr-

soid inﬂ orescence, either solitary on terminal branches

or aggregated into compact heads. Ovary 2-locular;

ovules numerous in each locule, n = 19. – Nuxia.

5.2 STILBOIDEAE

Stamens four. Shrubs or shrublets. Flowers 5-merous,

entomophilous, in dense terminal spikes. Ovary

2- locular, but one locule always empty, septum between

locules sometimes incomplete; fertile locule with two

basal-axile, erect ovules. – Campylostachys,

Euthystachys, Kogelbergia, Tesmophora, Stilbe

(including Xeroplana and Eurylobium).

Superorder LAMIANAE 565

5.3 RETZIOIDEAE

Stamens ﬁ ve, rarely six or seven. Monopodially

branched shrubs. Flowers ornithophilous, on very short

axillary shoots. Ovary basally 2-locular but apically

1-locular, the septum not reaching beyond the placenta,

this situated centrally a little more than halfway up the

locules. Ovules 2–3, each locule with one ascending

and 1–2 pendulous ovules, n = 12. – Retzia.

Have similarities with both Scrophulariaceae –

Manuleae and also with Buddlejaceae (Nuxia is some-

times included in Buddlejaceae).

6. SCROPHULARIACEAE

A.L. de Jussieu 1789 (including Aeginetiaceae Livera

1927, Antirrhinaceae Persoon 1807, Aragoaceae D.

Don 1835, Buchneraceae Lilja 1870, Calceolariaceae

Raf. ex Olmstead 2001, Caprariaceae Martynov 1820,

Chelonaceae Martynov 1820, Digitalidaceae Martynov

1820, Ellisiophyllaceae Honda 1930, Erinaceae Duvau

ex Pfeiffer 1874, Euphrasiaceae Martynov 1820,

Gratiolaceae Martynov 1820, Hebenstreitiaceae

Horaninow 1834, Hemimeridaceae Doweld 2001,

Limosellaceae J.G. Agardh 1858, Linariaceae

Martynov 1820, Lindenbergiaceae Doweld 2001,

Linderniaceae Borsch, K Müller, et Eb. Fischer 2004,

Melampyraceae Hooker and Lindley 1821,

Orobanchaceae Ventenat 1799, Oxycladaceae

Schnizlein 1857–1870, Paulowniaceae Nakai 1949,

Pediculariaceae A.L. de Jussieu 1789, Phelypaeaceae

Horaninow 1834, Rhinanthaceae Ventenat 1799,

Schlegeliaceae Reveal 1996, Selaginaceae Choisy

1823, Sibthorpiaceae D. Don 1835, Verbascaceae

Raﬁ nesque 1821, Veronicaceae Cassel 1817). c.300–

310/5000. Cosmopolitan, but more abundant in tem-

perate regions (especially in the Northern Hemisphere)

and in tropical mountains.

6.1 SCROPHULARIOIDEAE

The two posterior corolla lobes covering the lateral

lobes in bud. Placentation basically axile. Mostly auto-

trophic plants. – verbasceae: Verbascum (including

Celsia, Staurophragma), Rhabdotosperma; scrophu-

larieae: Nathalliella, Oreosolen, Scrophularia

(including Tuerckheimocharis), Antherothamnus;

ALONSOEAE: Alonsoa, Charadrophila; hemimeri-

deae: Basistemon, Diascia, Hemimeris, Diclis,

Nemesia, Colpias; russelieae: Russelia, Amero-

glossum, Dermatobotrys; teedieae: Ranopisoa,

Teedia; freylinieae: Phygelius, Freylinia; leu-

cophylleae: Eremogeton, Leucophyllum (including

Faxonanthus); aptosimeae: Anticharis, Aptosimum,

Peliostomum; manuleeae: Barthlottia, Manuleopsis,

Jamesbrittenia, Lyperia, Sutera (including Chaeno-

stoma, Sphenandra), Manulea, Melano spermum,

Polycarena, Glekia, Trieenea, Phyllopodium,

Zaluzianskya, Reyemia, Glumicalyx, Strobilopsis,

Tetraselago, Globulariopsis, Microdon (including

Agathelpis), Gosela, Cromidon, Selago (including

Walafrida), Pseudoselago, Chenopodiopsis, Dis-

chisma, Hebenstretia, ? Camptoloma; schlegelieae:

Gibsoniothamnus, Schlegelia, Synapsis, Exarata;

paulownieae: Paulownia

(including Shiuyinghua);

wightieae: Brandisia, Wightia; cheloneae:

Chelone, Nothochelone, Chionophila (including

Penstemonopsis), Keckiella, Pennellianthus, Pen-

stemon, Tetranema, Uroskinnera; collinsieae:

Collinsia, Tonella; antirrhineae: Anarrhinum,

Kickxia (including Elatinoides), Linaria, Cymbalaria,

Asarina, Maurandella, Rhodochiton, Lophospermum,

Pseudorontium, Schweinfurthia, Antirrhinum, Pseudo-

misopates, Misopates, Acanthorrhinum, Howelliella,

Neogarrhinum, Sairocarpus, Mohavea, Galvezia,

Chaenorrhinum, Albraunia, Holzneria, Nanorrhi-

num (including Pogonorrhinum), Nuttallanthus,

Maurandya, Epixiphium, Mabrya, Holmgrenanthe,

Gambelia; angelonieae: Angelonia, Monopera;

gratioleae: Amphianthus, Bacopa (including

Moniera, Herpestis, Hydranthelium, Ildefonsia,

Geochorda, Monocardia, Ancistrostylis, Sinobacopa),

Maeviella, Boelckea, Benjaminia, Braunblanquetia,

Sophronanthe, Gratiola (including Fonkia, Tragiola),

Mecardonia, Scoparia, Capraria, Deinostema,

Dopatrium, Hydrotriche, Limnophila, Philcoxia; limo-

selleae: Limosella; stemodieae: Lindenbergia

(including Omania), Adenosma, Otacanthus,

Achetaria, Tetraulacium, Dizygostemon, Stemodia

(including Poarium), Cheilophyllum, Morgania,

Schistophragma, Conobea, Leucospora, Schizosepala,

Stemodiopsis; lindernieae: Artanema, Picria,

Pierranthus, Schizotorenia, Legazpia, Torenia,

Scolophyllum, Hemiarrhena, Hamaegigas, Lindernia

(including Vandellia, Ilysanthes, Trichotaenia),

Craterostigma, Crepidorhopalon, Hartliella, Bampsia;

melospermeae: Monttea, Melosperma. microcar-

peae: Micranthemum (including Amphyolanthus,

Hemisiphonia), Hemianthus, Microcarpaea, Elacho-

loma, Peplidium, Encopella, Psammetes, Dintera,

Bryodes, Bythophyton, Glossostigma; leucocarpeae

566 Subclass VIII. LAMIIDAE

Hemichaena (including Berendtia, Berendtiella),

Leucocarpus; mimuleae: Dodartia, Lancea, Mazus,

Mimetanthe, Mimulus, Mimulicalyx.

6.2 RHINANTHOIDEAE

The posterior corolla lobes covered in the bud by one

or both of the lateral lobes. Placentation basically

axile. Autotrophic, hemiparasitic, or seldom wholly

parasitic plants (Lathraea). – digitaleae: Digitalis,

Isoplexis; veroniceae: Erinus, Lafuentea, Wulfenia,

Wulfeniopsis, Kashmiria, Picrorhiza, Neopicrorhiza,

Veronicastrum, Scrofella, Synthyris, Besseya,

Paederota, Paederotella, Pseudolysimachion, Veronica

(including Cochlidiosperma, Odicardis), Hebe,

Heliohebe, Parahebe, Chionohebe, Derwentia,

Detzneria, Lagotis, ? Campylanthus, Ourisia; sibthor-

pieae: Sibthorpia, Ellisiophyllum; hemiphragmeae:

Hemiphragma; aragoeae: Aragoa; gerardieae:

Agalinis (Gerardia), Anisantherina, Brachystigma,

Aureolaria, Tomanthera, Macranthera, Seymeria,

Seymeriopsis, Dasistoma, Silviella, Esterhazya,

Lamourouxia; escobedieae: Alectra (including

Pseudorobanche), Pseudomelasma, Melasma,

Magdalenaea, Velloziella, Nothochilus, Physocalyx,

Escobedia, Leptorhabdos, Gerardiina, Micrargeria,

Micrargeriella; buchnereae: Sopubia, Parasopubia,

Baumia, Petitmenginia, Sieversandreas, Pseudo-

sopubia, Hiernia, Graderia, Ghikaea, Centranthera,

Buchnera (including Benthamistella, Stellularia),

Tetraspidium, Striga, Pseudostriga, Parastriga,

Cycniopsis, Cycnium, Rhamphicarpa, Rhaphispermum,

Radamaea, Leucosalpa, Buttonia, Thunbergianthus,

Xylocalyx; cymbarieae: Schwalbea, Cymbaria

(including Cymbochasma), Siphonostegia, Lesque-

reuxia, Bungea, Monochasma, Castilleja, Orthocarpus,

Triphysaria, Clevelandia, Gentrya, Cordylanthus,

Ophiocephalus; rhinantheae: Hedbergia, Bartsia

(including Bellardia), Bornmuellerantha, Macro-

syringion, Odontitella, Bartsiella, Odontites (including

Orthanthella

), Nothobartsia, Euphrasia, Omphalothrix,

Pterygiella, Xizangia, Phtheirospermum, Parentucellia,

Pseudobartsia, Pedicularis, Melampyrum, Rhinanthus,

Rhynchocorys, Tozzia, Lathraea; rehmanniae:

Rehmannia, Titanotrichum, Triaenophora (including

Spirostegia); bowkerieae: Anastrabe, Bowkeria,

Brookea, Halleria, Ixianthes.

6.3 OROBANCHOIDEAE

The two posterior corolla lobes covered in bud by lat-

eral lobes. Placentation parietal. Parasitic herbs. –

Harveya, Paraharveya, Hyobanche, Aeginetia,

Boschniakia (including Kopsiopsis), Conopholis,

Christisonia (including Campbellia), Cistanche,

Phelypaea (including Diphelypaea), Gleadovia,

Mannagettaea, Orobanche (Aphyllon), Phacellanthus

(including Tienmuia), Platypholis, Xylanche, Epifagus,

Phelipanche, Necranthus.

The Scrophulariaceae stand close to the

Buddlejaceae, an afﬁ nity that is supported both by

morphology (including embryology) and chemistry.

The subfamily Orobanchoideae is very closely

related to the Rhinantoideae. It is an advanced group

that “represents the ﬁ nal stage of the parasitic tendency

exhibited in Rhinanthoideae” (Wernham 1912) and

can be regarded as a direct continuation of the line of

parasitism developed in the Scrophulariaceae (Boeshore

1920). It probably derived from the Rhinanthoideae

through forms like Striga orobanchoides (Tiagi 1956,

1963, 1970). There are important embryological simi-

larities between these two subfamilies (Tiagi 1950;

Terekhin and Nikiticheva 1981).

7. BIGNONIACEAE

A.L. de Jussieu 1789 (including Crescentiaceae

Dumortier 1829). 105/800. Mainly tropical regions,

especially in South America, with a few species in

subtropical and warm-temperate Asia.

Classiﬁ cation based on E. Fischer, I.Theisen and

L.G. Lohmann (2004).

tecomeae: Incarvillea, Niedzwedzkia, Perian-

thomega, Perichlaena, Pandorea, Tecomanthe,

Podranea, Dinklageodoxa, Campsidium, Campsis,

Markhamia, Spathodea, Newbouldia, Dolichandrone,

Digomphia, Rhigozum, Heterophragma, Santisukia,

Pajanelia, Tecoma, Lamiodendron, Fernandoa,

Radermachera (including Mayodendron), Stereo-

spermum, Pauldopia, Catophractes, Deplanchea,

Tecomella, Astianthus, Chilopsis, Catalpa, Jacaranda,

Spirotecoma, Romeroa, Tabebuia, Delostoma, Argylia,

Neosepicea, Cybistax, Zeyhria, Paratecoma,

Godmania, Sparattosperma, Ekmanianthe; oroxy-

leae: Oroxylum, Millingtonia, Nyctocalos, Hieris;

bignonieae: Spingiphila, Pseudocatalpa, Macfadyena,

Melloa, Dolichandra, Parabignonia, Phryganocydia,

Callichlamys, Manaosella, Gardnerodoxa, Neojo-

bertia, Pleonotoma, Memora, Pyrostegia, Bignonia,

Macranthisiphon, Piriadacus, Glaziovia, Amphilo-

phium, Haplolophium, Stizophyllum, Paragonia,

Superorder LAMIANAE 567

Roentgenia, Clytostoma, Mansoa, Cydista, Fridericia,

Martinella, Cuspidaria, Distictis, Saritaea, Potamo-

ganos, Arrabidaea, Xylophragma, Mussatia, Adeno-

calymma, Anemopaegma, Tanaecium, Leucocalantha,

Lundia, Tynanthus, Pithecoctenium, Spathicalyx,

Distictella, Ceratophytum, Periarrabidaea; eccremo-

carpeae: Eccremocarpus; tourrettieae: Tourrettia;

coleeae: Ophiocolea, Colea, Kigelia, Rhodocolea,

Phylloctenium, Phyllarthron; crescentieae: Parmen-

tiera, Amphi tecna, Crescentia.

Very near to the Scrophulariaceae, especially to

Scrophularieae (Cheloneae), and probably had a

common origin with them. They differ from

Scrophulariaceae mainly in endospermless seeds and

also in vascular anatomy of the ﬂ owers (see Armstrong

1985).

8. GESNERIACEAE

Richard et A.L. de Jussieu ex A.P. de Candolle 1816

(including Belloniaceae Martynov 1820, Besleriaceae

Raﬁ nesque 1838, Cyrtandraceae Jack 1823,

Didymo carpaceae D. Don 1822, Ramondaceae

Godron 1853). c. 145–150/2500–3500. Pantropical,

with only a few species in the Pyrenees and the Balkan

Peninsula.

Classiﬁ cation after A. Weber (2004).

8.1 CORONATHEROIDEAE

Trees, shrubs or subshrubs. Stomata anomocytic.

Nectary adnate to the ovary. Ovary superior. Fruits

capsules or berries, n = 37–45. – Coronanthera,

Depanthus, Fieldia, Lenbrassia, Negria, Rhabdo-

thamnus, Mitraria, Sarmienta, Asteranthera.

8.2 GESNERIOIDEAE

Perennial herbs, subshrubs, shrubs, small trees, vines,

climbers or epiphytes. Stomata anisocytic, scattered or

in groups. Cotyledons remaining equal after germina-

tion. Ovary superior or more or less inferior.

Inﬂ orescences often racemose. Nectary disc annular or

more often represented by 1–5 separate glands, some-

times wanting, n = 6–16. – beslerieae: Besleria,

Gasteranthus, Cremosperma, Anetanthus, Resia,

Reldia, Tylopsacas; nepeantheae: Napeanthus;

gloxinieae: Gloxinia, Anodiscus, Koellikeria,

Monopyle, Diastema, Kohleria, Pearcea, Capanea,

Achimenes, Eucodonia, Smithiantha, Niphaea,

Moussonia, Solenophora, Phinaea, Heppiella,

Goyazia; gesnerieae: Pheidonocarpa, Gesneria,

Rytidophyllum; sinningieae: Sinningia, Paliavana,

Vanhouttea; episcieae: Nautilocalyx, Chrysothemis,

Paradrymonia, Alloplectus, Drymonia, Columnea,

Dalbergaria, Pentadenia, Trichantha, Bucinellina,

Corytoplectus, Neomortonia, Episcia, Alsobia,

Rufodorsia, Oerstedina, Cobananthus, Nematanthus,

Codonanthe, Codonanthopsis; bellonieae: Bellonia,

Lembocarpus, Rhoogeton, Lampadaria, Cremersia,

Cremospermopsis.

8.3 EPITHEMATOIDEAE

Plants usually ﬂ eshy-succulent, sometimes with

strongly asymmetrical leaves. Ovary short, abruptly

narrowed into the style. Fruit more or less globose.

Endosperm absent, n = (8-)10(-12). – Rhynchoglossum,

Gyrogyne, Stauranthera, Loxonia, Epithema,

Whytockia, Monophyllaea (including Moultonia).

8.4 DIDYMOCARPOIDEAE

(CYTRTANDROIDEAE)

Perennial or rarely annual herbs. Cotyledons becoming

unequal after germination. Ovary superior. Inﬂ orescences

various, usually cymose with two ﬂ owers at each dichot-

omy, never a simple raceme. Nectary disc annular, rarely

divided or split on one side, n = (4) 8–13, 14–17, etc. –

Corallodiscus, Leptoboea, Boeica, Rhynchotechum,

Tetraphyllum, Platystemma, Championia, Haberlea,

Ramonda, Jankaea, Streptocarpus, Saintpaulia,

Linnaeopsis, Acanthonema, Trachystigma, Nodonema,

Schizoboea, Hovanella, Colpogyne, Oreocharis (includ-

ing Dasydesmus and Perantha

), Tremacron, Isometrum,

Paraisometrum, Ancylostemon, Petrocosmea, Metape-

trocosmea, Deinocheilos, Lagarosolen, Petrocodon,

Dayaoshania, Dolicholoma, Calcareoboea, Allocheilos,

Opithandra, Thamnocharis, Bournea, Tengia, Conan-

dron, Briggsia, Briggsiopsis, Raphiocarpus, Loxostigma,

Anna, Lysionotus, Cathayanthe, Beccarinda, Chirita,

Chiritopsis, Primulina, Metabriggsia, Didymocarpus,

Gyrocheilos, Didymostigma, Deinostigma, Pseudo-

chirita, Allostigma, Phylloboeae, Hemiboeopsis,

Hemiboea, Didissandra, Ridleyandra, Hexatheca,

Aeschynanthus (including Euthamnus, Oxychlamys),

Micraeschynanthus, Agalmyla (including Dichro-

trichum), Orchadocarpa, Henckelia (including Loxo-

carpus, Codonoboea, Platyadenia), Boea, Paraboea

(including Chlamydoboea), Trisepalum (including

Dichiloboea), Rhabdothamnopsis, Ornithoboea,

Kaisupeea, Senyumia, Spelaeanthus, Emarhendia,

Cyrtandra (including Protocyrtandra), Sepikaea.

568 Subclass VIII. LAMIIDAE

Genera of uncertain familial afﬁ liation: Sanango,

Cubitanthus, Jerdonia.

Very near to the Scrophulariaceae, especially to the

tribe Scrophularieae. According to Burtt (1977), “a

highly advanced, and presumably recent family.”

9. PLOCOSPERMATACEAE

Hutchinson 1973. 1/1. Southern Mexico and Guatemala.

Plocosperma (including Lithophytum)

Have similarities, including two parietal placentas,

with the Gesneriaceae.

10. CARLEMANNIACEAE

Airy Shaw 1965. 1/6. Himalayas (from eastern Nepal

to Bhutan), northeastern India, northern Burma, south-

western China, Laos, northern Vietnam, Sumatra.

Carlemannia, Silvianthus

Hooker (in Bentham et Hooker 1973, Genera

Plantarum 2: 63–64) included Carlemannia and

Silvianthus in Rubiaceae-Hedyotideae. However, later

Solireder (1893) proposed to transfer them into the

Caprifoliaceae. Hallier (1910a, b), however, suggested

that Carlemanniaceae are closely related to

Gesneriaceae. Savolainen et al. (2000) and Bremer

et al. (2001) indicated an inclusion of Carlemanniaceae

into Lamiales.

11. GLOBULARIACEAE

A.P. de Candolle 1805. 2/25. Macaronesia,

Mediterranean, Europe, western Asia, and also Somalia

and Socotra (Poskea, 2).

Globularia, Poskea

Globulariaceae are closely related to the Scrophula-

riaceae-Selagineae, differing from them especially in

their inﬂ orescence as well as in pseudomonomerous

gynoecium and fruit morphology. Probably derived

from the Scrophulariaceae.

12. PLANTAGINACEAE

A.L. de Jussieu 1789. 3/270. Temperate regions and in

tropical mountains; Bougueria (1): Andes.

Plantago, Littorella, Bougueria

Near to and probably derived from the

Scrophulariaceae. Hallier (1903, 1912) included them

in Scrophulariaceae s. 1. and placed them near

Selagineae and Manuleae.

13. CALLITRICHACEAE

Berchtold et J. Presl 1820. 1/45–50. Nearly cosmo-

politan, but mainly in temperate regions.

Callitriche

According to Olmstead et al. (2001), the clade

Callitrichaceae, Globulariaceae, Plantaginaceae, and

Hippuridaceae is support very strong.

14. HIPPURIDACEAE

Vest 1818. 1/1. Temperate and cold regions of the

Northern Hemisphere, Antarctic America and south-

ern Australia.

Hippuris

Hippuridaceae are the closest relative of the

Callitrichaceae (Leins and Erbar 2004).

15. PEDALIACEAE

R. Brown 1810 (including Sesamaceae Berchtold et J.

Presl 1820). 13/75. From tropical and southern Africa

and Madagascar to Malesia, New Guinea, and north-

ern Australia and adjacent islands.

pedalieae: Pedalium, Pterodiscus, Pedaliodiscus,

Harpagophytum, Holubia, Uncarina; sesameae:

Sesamothamnus, Rogeria, Sesamum, Ceratotheca;

pretreeae: Dicerocaryum (Pretrea), Linariopsis,

Josephinia.

Close to the Bignoniaceae and Scrophulariaceae.

16. MARTYNIACEAE

Stapf 1895. 4/17. Subtropical and especially tropical

regions of America.

Martynia, Holoregmia, Craniolaria, Proboscidea

(including Ibicella).

Have many similarities with Pedaliaceae, including

wood anatomy, but comparison of pollen suggested

Superorder LAMIANAE 569

that Martyniaceae and Pedaliaceae should be main-

tained as separate families (Stapf 1895; Bretting and

Nilsson 1988).

17. TRAPELLACEAE

Honda et Sakisake 1930. 1/2. Eastern Asia.

Trapella

Very near to the Pedaliaceae. Previously only a few

botanists, including Airy Shaw (in Willis 1973) and

Dahlgren (1980, 1983, 1989), accepted Trapellaceae

as a distinct family. However, the Trapellaceae differ

not only in pollen, ovule, ovary, and fruit structure but

also in such embryological features as persistence of

antipodals, simultaneous differentiation of micropylar

and chalazal haustorium, and a four-celled micropylar

haustorium and chalazal haustorium of two elongated

cells (John et al. 1992: 774).

18. MYOPORACEAE

R. Brown 1810 (including Bontiaceae Horaninow

1834). 4/210. Mainly Australia, but Myoporum (32)

also extends to New Guinea, eastern Asia, New

Zealand, Paciﬁ c islands, and Mascarene Islands;

Bontia (1) occurs in the West Indies and northern South

America, and Androya (1) is endemic to Madagascar.

Eremophila (Pholidia, Stenochilus), Myoporum,

Bontia, Androya

Closely related to the Scrophulariaceae –

Scrophularioideae. According to palynological data

(Niezgoda and Tomb 1975) Myoporaceae related to

the scrophularien tribe Leucophylleae. In addition

Karrfalt and Tomb (1983) found that air spaces of

Leucophyllum are homologous with the secretory cavi-

ties of Bontia (Myoporaceae).

19. OFTIACEAE

Takhtajan et Reveal 1993 (Spielmanniaceae J. Agardh

1858, nom.illeg.). 1/3. South Africa

Oftia

Oftia (Spielmannia) has been placed in various

families, such as Verbenaceae (Endlicher 1841; Lindley

1853; and some others), Scrophulariaceae (Baillon

1888; Thorne 1992, 2000), and Myoporaceae (Bentham

and Hooker 1876; Takhtajan 1966; Cronquist 1981;

and many others), or separated into a family of its own

(Takhtajan 1987, 1997). From the Myoporaceae the

genus Oftia differs in the presence of internal phloem

and in primary vascular tissue. Besides, it differs from

the Myoporaceae in the absence of the secretory cavi-

ties and not gland-dotted leaves, and from the

Scrophulariaceae in absent endosperm. However, the

Oftiaceae have some links to both the Scrophulariaceae

and Myoporaceae.

20. ACANTHACEAE

A.L. de Jussieu 1789 (including Justiciaceae Raﬁ nesque

1838, Mendonciaceae Bremekamp 1954, Nelsoniaceae

Sreemadhavan 1977, Thomandersiaceae Sreemadhavan

1977, Thunbergiaceae Lilja 1870). 220–240/3500–

4000. Mainly tropical regions, especially southern and

Southeast Asia, Africa, Brazil, and Central America; a

few genera extend into warm-temperate regions; some

species of Acanthus occur in the Mediterranean region

and in western Asia.

20.1 NELSONIOIDEAE

Ovary 2-locular with numerous ovules or (Elytraria)

1-locular with 6–10 ovules on parietal placentas;

ovules with endothelium. Jaculators usually wanting.

Fruits capsular. Seeds with more or less developed oily

and ruminate endosperm. Leaves alternate. Cystoliths

wanting; descending cochlear aestivation. – Staurogyne,

Nelsonia, Elytraria, Anisosepalum, Gynocraterium,

Ophiorrhiziphyllon, Saintpauliopsis.

20.2 THUNBERGIOIDEAE. (including

Mendoncioideae)

Ovary 2-locular with two ovules per locule or less

often one of the locules aborted. Jaculators wanting.

Fruits capsules or drupes. Seeds without endosperm.

Leaves usually opposite. Cystoliths present. – thun-

bergieae: Thunbergia, Meyenia, Pseudocalyx; mend-

oncieae: Mendoncia, Anomacanthus

20.3 ACANTHOIDEAE

Ovary 2-locular with two to many ovules per locule.

Jaculators usually present. Seeds without endosperm.

Leaves opposite. Cystoliths present or absent

(Acantheae). – acantheae: Acanthus, Acanthopsis,

Achyrocalyx, Blepharis, Crossandra, Crossndrella,

570 Subclass VIII. LAMIIDAE

Gynarospermum, Cyphacanthus, Encephalosphaera,

Geissomeria, Holographis, Neriacanthus, Oropho-

chilus, Rhombochlamys, Salpixantha, Sclerochiton,

Streptosiphon, Strobilacanthus, Xantheranthemum;

aphelandreae: Stenandrium, Aphelandra; ruel-

lieae: Blechum, Acanthopale, Apassalus, Benoicanthus,

Bravaisia, Brunoniella, Calacanthus, Clarkeasia,

Dischistocalyx, Duosperma, Sanchezia, Ruellia,

Dyschoriste, Echinacanthus, Epiclastopelma,

Eranthemum, Eremomastax, Eusiphon, Hemigraphis,

Heteradelphia, Hygrophila, Ionacanthus, Kosmosiphon,

Leptosiphonium, Louteridium, Lychniothyrsus, Mellera.

Mimulopsis, Pararuellia, Petalidium, Phaulopsis,

Physacanthus, Polylychnis, Pseudoruellia, Ruelliospis,

Satanocrater, Sautiera, Spirostigma, Stenosipho-

nium, Stenothyrsus, Strobilanthes, Strobilanthopsis,

Suessenguthia, Trichanthera, Trichosanchezia,

Zygoruellia; lepidagathidae: Lepidagathis; andro-

graphinae: Andrographis, Cystacanthus, Diotacanthus,

Graphandra, Gymnostachyum, Haplan thodes,

Indonesiella, Phlogacanthus; Afroﬁ t tonia, Ambongia,

Ancistranthus, Angkalanthus, Anisacanthus,

Aphanosperma, Asotheca, Asystasia, Ballochia,

Brachystephanus, Calycacanthus, Carlowrightia,

Celerina, Centrilla, Cephalocanthus, Chalarothyrsus,

Chamaeranthemum, Chileranthemum, Chlamydo-

cardia, Chlamydostachya, Chorisochora, Clinacanthus,

Clistax, Codonacanthus, Conocalyx, Cosmianthemum,

Cyclacanthus, Cylindrosolenium, Danguya, Dasytropis,

Dichazothece, Dicladanthera, Dicliptera, Ecbolium,

Filetia, Fittonia, Forcipella, Glossochilus, Grapto-

phyllum, Gypsacanthus, Harpochilus, Henrya,

Herpetacanthus, Hoverdenia, Hypoestes, Ichtyostoma,

Isoglossa, Isotheca, Jadunia, Juruasia, Justicia,

Kalbreyeriella, Linariantha, Mackaya, Marcania,

Megalochlamys, Megalostoma, Megaskepasma,

Mellitacanthus, Metarungia, Mexacanthus, Mirandea,

Monechma, Monothecium, Odontonema, Oplonia,

Oreacanthus, Pachystachys, Pelecostemon, Peristrophe,

Phialacanthus, Podorungia, Poikilacanthus, Polulina,

Pranceacanthus, Pseu deranthemum, Pseudodicliptera,

Psilanthele, Ptyssiglottis, Pulchranthus, Razisea,

Rhinacanthus, Ritonia, Rungia, Ruspolia, Ruttya,

Samuelssonia, Sapphoa, Schaueria, Sebastiano-

Schaueria, Spathacanthus, Sphinctacanthus, Steno-

stephanus, Streblacanthus, Tessmanniacanthus,

Tetramerium, Thysanostigma, Trichaulax, Trichocalyx,

Xerothamnella, Yeatesia; Barleria, Barleriola,

Borneacanthus, Boutonia, Chroesthes, Crabbea,

Hulemacanthus, Lepidagathis, Lophostachys.

whitfieldieae: Whitﬁ eldia, Chlamydacanthus,

Lankesteria.

insertae sedis: Acanthostelma, Acanthura,

Aphelandrella, Camarotea, Dolichostachys, Golaea,

Idiacanthus, Kudoacanthus, Lasiocladus, Leandri-

ella, Morsacanthus, Neuracanthus, Perenideboles,

Pericalypta, Sphacanthus, Vavara, Vindasia.

Near to the Scrophulariaceae. Nelsonioideae are in

some respects a connecting link between the two

families.

21. AVICENNIACEAE

Endlicher 1841. 1/8. Mostly tropical coasts mainly

within the mangrove zone of tropical sheltered coast-

lines, but also in subtropical and even warm-temperate

tidal habitats where Avicennia occurs as the exclusive

tree or shrub form, making it the most widely distrib-

uted of any mangrove. Three species of Avicennia

occur in the Atlantic, Caribbean, and eastern Paciﬁ c,

while ﬁ ve other species occur in the Old World

( Tom-Unson 1986; Duke 1991).

Avicennia

According to Schwarzbach and McDade (2002),

“sequence data from these two chloroplast and one

nuclear regions consistently place Avicenia with

Acanthaceae s.l. They analyzed that resolve relationships

further place Avicennia as sister to Thunber gioideae”.

However, from the Acanthaceae they differ in the second-

ary anomalous growth, in leaves with salt glands on both

sides, orthotropous ovules and fruit morphology.

22. BYBLIDACEAE

Domin 1922. 1/6. The southernmost part of New

Guinea, northern and southwestern Australia.

Byblis

According to recent studies, both molecular and

morphological, the Byblidaceae are related to the

Lentibulariaceae (see especially Albert et al. 1992;

Conran and Carolin 2004).

23. LENTIBULARIACEAE

Richard 1808 (including Pinguiculaceae Dumortier

1829, Utriculariaceae Hoffmannsegg and Link 1809).

3/320. Cosmopolitan.

Superorder LAMIANAE 571

Pinguicula, Genlisea, Ultricularia (including

Polypompholyx and Biovularia)

Near to and derived from the Scrophulariceae.

Hallier (1903a, b, 1905, 1908, 1912) included Lenti-

bulariaceae in Scrophulariaceae s.1., deriving them

from Gratioleae (see also Casper 1963). Embryological

data suggest the derivation from the Scrophulariaceae

(Khan 1970; Johri et al. 1992). Floral morphology and

phytochemistry (iridoid glycosids) indicate a close

relationship to Scrophulariaceae.

24. VERBENACEAE

J. Saint-Hilaire 1805 (including Durantaceae J.G.

Agardh 1858, Lantanaceae Martynov 1820, Petreaceae

J.G. Agardh 1858, Viticaceae A.L. de Jussieu 1789).

50/1800–2500. Tropical and subtropical regions with a

few species in warm-temperate countries.

24.1 VITICOIDEAE

Leaves opposite or rarely verticillate, simple or some-

times 3–7-foliolate. Inﬂ orescence cymose, corymbose,

umbellate, or occasionally ﬂ owers solitary. – tejsman-

niodendreae: Tejsmanniodendron, Garrettia,

Hymenopyramis; callicarpeae: Callicarpa (includ-

ing Geunsia); tectoneae: Petitia, Tectona; viticeae:

Pseudocarpidium, Cornutia, Adelosa, Viticipremna,

Premna (including Pygmaeopremna), Tsoongia,

Paravitex, Petraeovitex, Vitex (including Neorapinia),

Gmelina.

24.2 VERBENOIDEAE

Leaves opposite or rarely alternate, entire, lobed to

dissected, sometimes much reduced, scale-like, or

spinose. Flowers in various types of racemose inﬂ o-

rescences, often with involucre of colored bracts. –

verbeneae: Verbena (including Stylodon),

Glandularia, Junellia, Urbania, Hierobotana; lanta-

neae: Lantana, Neosparton, Xeroaloysia, Nashia,

Lippia (including Burroughsia), Phyla, Aloysia,

Acantholippia, Stachytarpheta (including Ubochea),

Bouchea, Chascanum (including Svensonia), Diostea,

Lampaya; priveae: Priva, Pitraea (including

Castelia), Dipyrena; petreae: Petrea, Xolocotzia;

casselieae: Casselia, Parodianthus, Tamonea;

duranteae (citharexyleae): Citharexylum,

Rehdera, Verbeno xylum, Rhaphithamnus, Baillonia,

Recordia, Duranta.

Insertae sedis: Coelocarpum.

25. PHRYMATACEAE

Schauer 1847. 1/1. Himalayas, eastern Asia, south-

eastern North America.

Phryma

Closely related to the Verbenaceae-Verbeneae,

especially to Verbeneae (see Whipple 1972; Chadwell

et al. 1992; Wagstaff 1992; Cantino 1992a, b; Ramana

et al. 2000). In overall morphology closely resembles

Stachytarpheta, but differs in having trichomes and in

pollen morphology.

26. CYCLOCHEILACEAE

Marais 1981. 2/4. Eastern and northeastern tropical

Africa and Yemen.

Cyclocheilon, Asepalum

A distinct family probably derived from the ver-

benaceous stock.

27. NESOGENACEAE

Marais 1981. 1/9. Tanzania, Madagascar, Seychelles

and Mascarene Is., Malesia, Oceania (Tuamotu

Arch.).

Nesogenes

Close related to the Cyclocheilaceae.

28. SYMPHOREMATACEAE

Wight 1849. 3/35. Southern, Southeast, and eastern

Asia, Malesia.

Symphorema, Sphenodesme, Congea

Related to the Verbenaceae.

29. LAMIACEAE

Martynov 1820 or Labiatae A.L. de Jussieu 1789, nom.

altern. (including Aegiphilaceae Raﬁ nesque 1838,

Chloanthaceae Hutchinson 1959, Dicrastylidiaceae

Drummond ex Harvey 1855, Glechomaceae Martynov

1820, Melittdaaceae Martynov 1820, Menthaceae

Burnett 1835, Nepetaceae Horaninow 1834,

Salazariaceae F.A. Barkley 1975, Salviaceae

Raﬁ nesque 1837, Scutellariaceae Caruel 1894,

Siphonanthaceae Raﬁ nesque 1838). Over

572 Subclass VIII. LAMIIDAE

230–234/c.7000. Cosmopolitan, but especially diversi-

ﬁ ed in the Mediterranean, Irano-Turanian, and Eastern

Asiatic Regions.

29.1 AJUGOIDEAE

Shrubs, subshrubs, or herbs, sometimes aromatic or

ill-scented. Leaves opposite. Flowers more or less

zygomorphic. Calyx 5-lobed or dentate. Corolla often

with deeply cleft tube (Teucrium, Teucridium,

Monochilus, and some species of Spartothamnella),

the corolla limb therefore seemingly unilabiate. Pollen

grains 3-colpate or rarely 3-porate, often operculate

(Teucrium and related genera), with granular exine

structure and often more or less branched columella

(profusely branched in Teucrium), suprareticulate,

tectate-perforate to microreticulate, with supratectal

spinules or spines or verrucate. Style terminal to

gynobasic; ovary mostly shallowly lobed, completely

or incompletely 4-locular, with one ovule per locule

(locellus). Nutlets with lateral-ventral attachment, the

usually large surface of contact often more than half

the height of the ovary. Seeds with straight embryo

and straight and short radicle; endosperm scanty or

none. Terpenoids and rosmarinic acid absent, iridoid

glycosides and acteosides present, n = 7, 10, 13, 14, 16

+. – clerodendreae: Clerodendrum (including

Kalaharia, Tetraclea), Rotheca, Faradaya, Teucridium,

Oncinocalyx; ajugeae: Ajuga; monochileae:

Aegiphila, Amasonia, Monochilus, Amethystea; teu-

crieae: Teucrium, Spartothamnella; caryop-

terideae: Caryopteris, Glossocarya, Hosea, Huxleya,

Karomia, Oxera, Rubiteucris, Discre-

titheca, Pseudocaryopteris, Tripora, Schnabelia,

Trichostema.

29.2 PROSTANTHEROIDEAE

Shrubs, subshrubs, or less often trees, non-aromatic or

occasionally with distal branches and leaves aromatic,

usually provided with various types of trichomes, often

tomentose. Leaves opposite or verticillate, rarely alter-

nate, entire or dentate, often densely clothed with stel-

late hairs. Flowers in terminal or axillary inﬂ orescences,

often spicate, capitate or corymbose-paniculate. Calyx

lobed or dentate. Corolla actinomorphic or zygomor-

phic, lobed to truncate. Stamens 2–8, mostly four.

Pollen grains 3-colpate or sometimes 6(-8)-colpate,

tectate-perforate to microreticulate, with or without

spinulose ornamentation, often with ectexine elements

on the colpi membrane. Nectary disc wanting. Style

terminal or gynobasic. Ovary 2-locular, often shal-

lowly 4-lobed, with usually two ovules per carpel.

Fruits indehiscent, often 1-seeded or separated into

four mericarps. Seeds with straight embryo and usu-

ally with copious or scanty endosperm. – chloanthae:

Chloanthes, Hemiphora, Cyanostegia, Dicrastylis,

Mallophora, Physopsis, Newcastelia, Lachnostachys,

Pityrodia, Brachysola; westringieae (prostan-

thereae): Hemiandra, Hemigenia, Microcorys,

Prostanthera (including Eichlerago), Wrixonia,

Westringia.

29.3 WENCHENGIOIDEAE

Subshrubs. Leaves alternate, entire. Flowers zygomor-

phic. Calyx 2-lipped, 5-dentate. Corolla 5-lobed, three

lower lobes much larger. Stamens four, subequal.

Pollen grains 3-colpate, suprareticulate, with simple

columellae. Nectary disc absent. Style subterminal,

shortly 2-lobed. Ovary shallowly lobed, 4-locular, with

one ovule per locule. Fruit of nutlike mericarps attached

to the receptacle by means of slender stalks. Seeds

with spatulate embryo, endosperm present. –

Wenchengia.

29.4 SCUTELLARIOIDEAE

Herbs, subshrubs, or rarely shrubs. Leaves opposite,

entire or dentate. Calyx 2-labiate with entire, rounded

lips that become tightly appressed after anthests,

enclosing the developing fruit inside. Corolla with

4-lobed upper lip and 1-lobed lower lip. Nectary disc

tubular, elongate. Anterior stamens dimidiate. Pollen

grains mostly inoperculate, suprareticulate to psilate,

tectate-perforate to microreticulate, with simple colu-

mellae. Style mostly attached to the ovary lobes above

the apex of the disc extension, rarely (some species of

Scutellaria) truly gynobasic. Nutlets with basal attach-

ment, the surface of contact small. Seeds more or less

transverse. Embryo with a bent radicle on one cotyle-

don. – Scutellaria (including Perilomia, Salazaria,

Cruzia, Harlanlewisia), Renschia, Tinnea,

Holmskioldia.

29.5 LAMIOIDEAE

Herbs, subshrubs, or rarely shrubs, mostly non-aro-

matic or slightly aromatic. Leaves opposite, entire,

dentate or lobed. Inﬂ orescence thyrsoid or racemelike,

rarely ﬂ owers solitary. Calyx of various shape. Corolla

bilabiate to nearly actinomorphic. Stamens four,

ascending or spreading and projecting straight

Superorder LAMIANAE 573

forward, often long exserted from the corolla; anthers

opening longitudinally, rarely by valves (Galeopsis).

Pollen grains mostly 3-colpate, but with four colpi

arranged in two pairs in Haplostachys, tetrazonocol-

pate in one species of Marrubium and many species

of Sideritis and hexapantocolpate in Sideritis sect.

Empedoclia (Abu-Asab and Cantino 1992), usually

tectate-perforate to microreticulate, mostly supraretic-

ulate, usually with simple columellae. Disc lobes,

when present, alternate with ovary lobes. Style gynoba-

sic. Ovary 4-locular, deeply 4-lobed; ovule solitary in

each locule. Nutlets with small basal attachment,

sometimes (Prasieae) with ﬂ eshy exocarp. Seeds with

spatulate embryo with short, straight superior radicle;

endosperm several-layered. Often present iridoid gly-

cosides and laballenic fatty acid, n = 6. – pogoste-

meae: Pogostemon (including Eusteralis, Anuragia),

Dysophylla, Colebrookea, Eurysolen, Leucosceptrum,

Comanthosphace, Rostrinucula, Anisomeles, Suzukia;

prasieae: Gomphostemma, Bostrychanthera,

Stenogyne, Phyllostegia, Haplostachys, Prasium;

marrubieae: Acrotome, Paralamium, Thuspeinantha,

Hypogomphia, Marrubium, Lagopsis, Sideritis;

lamieae: Physostegia, Chelonopsis, Synandra,

Macbridea, Brazoria, Warnockia, Melittis, Alajja,

Eriophyton, Loxocalyx, Ajugoides, Matsumurella,

Lamium (including Galeobdolon), Orvala, Wied-

emannia, Galeopsis, Paraphlomis, Lamiophlomis,

Phlomis, Phlomoides, Eremostachys, Paraeremo-

stachys, Pseuderemostachys, Pseudomarrubium,

Stachyopsis, Leonurus, Chaiturus, Panzerina

(Panzeria), Lagochilus, Moluccella, Otostegia, Iso-

leucas, Ballota, Sulaimania, Roylea, Metas tachydium

(Metastachys), Stachys, Epimeredi, Colquhounia,

Craniotome, Achyrospermum, Microtoena, Phlomido-

schema, Chamaesphacos, Notochaete, Leonotis,

Leucas (Physoleucas).

29.6 NEPETOIDEAE

Shrubs and herbs, rarely trees, commonly aromatic.

Leaves simple, entire or lobed, sometimes compound.

Corolla zygomorphic, sometimes weakly so, often

strongly 2-lipped. Stamens four or two, ascending or

declinate. Pollen grains usually 3-celled and typically

6(8–12)-colpare, predominantly suprareticulate. Disc

usually well developed, often 4-lobed. Style gynoba-

sic. Seeds without endosperm (except Bystropogon).

Pericarp usually dry, rarely ﬂ eshy; exocarp with

mucilaginous cells producing hygroscopic spiral

ﬁ brils. Embryo erect, investing (except Catoferia

which has bent embryo). Rich in volatile terpenoids

and rosmarinic acid; nepetoidin A and B (caffeic acid

esters) present. Iridoid glycosides and acteosides usu-

ally absent. Seed oils highly unsaturated, n = 6 +. –

elsholtzieae: Elsholtzia, Perilla, Perillula,

Collinsonia (including Micheliella, Keiskea), Mosla;

satureieae: Mentha (including Preslia), Lycopus,

Hyssopus, Pycnanthemum, Origanum (including

Majorana), Amaracus, Bystropogon, Minthostachys,

Thymus, Zataria, Monardella, Cyclotrichium,

Obtegomeria, Thymbra (including Coridothymus),

Cunila, Piloblephis, Ziziphora, Melissa, Heterolamium,

Conradina, Micromeria, Gontscharovia, Cuminia,

Satureja (including Euhesperida), Ceratominthe,

Saccocalyx, Kurzamra, Dicerandra, Clinopodium,

Calamintha, Acinos, Antonina, Hedeoma, Poliomintha,

Hesperozygis, Rhododon, Stachydeoma, Pogogyne,

Eriothymus, Pentapleura; prunelleae: Prunella,

Cleonia; meriandreae: Zhumeria, Meriandra,

Dorystoechas, Perovskia; lepechinieae: Sphacele,

Lepechinia; hormineae: Horminum; nepeteae:

Dracocephalum, Lophanthus, Hymenocrater,

Agastache, Brittonastrum, Meehania, Glechoma,

Schizonepeta, Cedronella, Nepeta (including Pitardia),

Kudrjaschevia, Drepanocaryum, Lallemantia,

Monarda, Marmoritis (including Phyllophyton),

Blephilia, Chaunostoma, Neoeplingia, Rhabdocaulon,

Hoehnea; glechoneae: Glechon, Acanthomintha;

salvieae: Salvia (including Ramona, Schraderia,

Arischrada); rosmarineae: Rosmarinus; catofer-

ieae: Catoferia; ocimeae: Eriope (including

Eriopidion), Hypenia, Hyptis, Hyptidendron,

Marsypianthes, Peltodon, Rhaphiodon, Asterohyptis,

Aeollanthus (including Icomum, Bovonia), Alvesia

(including. Plectranthastrum), Pycnostachys,

Anisochilus (including Stiptanthus), Leocus,

Plectranthus (including Rabdosia, Rabdosiella,

Neohyptis, Solenostemon, Coleus, Englerastrum,

Ascocaridion, Neomuellera, Symphostemon,

Perrierastrum, Holostylon, Capitanya,), Thorncroftia,

Tetradenia, Isodon (including Amethystanthus,

Skapanthus), Siphocranion, Hanceola, Isodictyophorus,

Hoslundia, Syncolostemon, Platostoma (including

Mesona, Ceratanthus, Octomeron), Basilicum,

Benguellia, Endostemon (including Puntia),

Hemizygia, Ocimum (including Becium,

Erythrochlamis, Nautochilus), Orthosiphon, Fuerstia,

Geniosporum, Haumaniastrum (including