[Plantaginales+Solanales]


PLANTAGINALES Juss. ex Bercht. et J. Presl

Bercht. et J. Presl, Přir. Rostlin: 241. Jan-Apr 1820 [‘Plantagineae’]

Lamiales Bromhead in Mag. Nat. Hist., ser. 2, 2: 210. Apr 1838; Verbenopsida Brongn., Enum. Plant. Mus. Paris: xx, 64. 12 Aug 1843 [’Verbenineae’]; Lamianae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnoliph.]: 405. 4 Feb 1967

Habit Usually bisexual (sometimes gynomonoecious, gynodioecious; rarely monoecious, dioecious, polygamomonoecious, or functionally dioecious), usually perennial, biennial or annual herbs, evergreen shrubs or suffrutices (sometimes evergreen or deciduous trees, shrubs, suffrutices or lianas). Many species are xerophytic. Some species are epiphytes. Four known lineages have evolved carnivory and/or insect-trapping mechanisms, usually by foliar mucilage glands.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Secondary lateral growth normal, anomalous (via concentric cambia or cylindrical cambium) or absent. Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits alternate, simple and/or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements fibre tracheids or libriform fibres (sometimes tracheids) with simple or bordered pits, septate or non-septate (often also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, unilateral or banded, or absent (rarely apotracheal diffuse or diffuse-in-aggregates). Wood elements sometimes storied. Tyloses sometimes abundant. Secondary phloem often stratified into hard fibrous and soft parenchymatous layers. Intraxylary phloem rarely present. Sieve tube plastids usually Ss type (sometimes Pc or Pcs type). Nodes usually 1–3:1–3, unilacunar to trilacunar with one to three leaf traces (rarely >3:>3, multilacunar with more than three traces). Schizogenous secretory canals with oils or resins rarely present. Heartwood sometimes with gum-like substances. Medulla sometimes with crystal inclusions. Silica bodies sometimes present in wood ray cells. Styloids, druses, crystal sand and acicular or prismatic calciumoxalate crystals sometimes frequent (rarely rhomboidal crystals, sphaerites or raphides).

Trichomes Hairs unicellular or multicellular, simple or branched, furcate, stellate, dendritic, peltate or lepidote (rarely moniliform); usually multicellular (sometimes unicellular) stalked or sessile glandular hairs often frequent; head of glandular hairs with vertical or horizontal divisions.

Leaves Usually opposite (sometimes verticillate or alternate/spiral), simple or pinnately or palmately compound (rarely bipinnate), entire or pinnately or palmately lobed, often coriaceous, often ericoid (sometimes scale-like), with conduplicate or involute (sometimes curved, rarely circinnate or flat) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular; petiole sometimes also with edge bundles or adaxial bundles. Venation pinnate or palmate, usually eucamptodromous to semicraspedodromous (sometimes brochidodromous or parallelodromous), or leaves one-veined. Stomata usually diacytic, paracytic or anomocytic (sometimes diallelocytic, anisocytic, helicocytic, tetracytic or cyclocytic). Cuticular wax crystalloids as rodlets or platelets (rarely threads). Domatia as pits, pockets or hair tufts, or absent. Crystal inclusions of various kinds often present. Mesophyll often with sclerenchymatous idioblasts. Mesophyll cells often with calciumoxalate as prismatic crystals (sometimes druses, sphaerites, styloids, raphides, acicular crystals or crystal sand). Leaf margin entire, serrate, crenate or lobate. Leaf teeth often with glandular apex, one accessory vein proceeding into tooth, the other running above tooth.

Inflorescence Terminal or axillary, panicle, cincinnus, raceme-like, spicate, corymb, umbel-like or capitate thyrse, often as whorls of dichasial and/or monochasial partial inflorescences, or racemes, spikes, umbels or heads (flowers sometimes solitary or paired, axillary). Bracts and floral prophylls (bracteoles) rarely absent.

Flowers Usually zygomorphic (rarely resupinate; sometimes actinomorphic). Usually hypogyny (sometimes half epigyny, rarely epigyny). Sepals (three to) five (to 16), with imbricate, contorted, valvate or open aestivation, usually persistent, usually connate, calyx often bilabiate. Petals (three to) five (to 16), with imbricate, contorted or ascending-cochlear (sometimes convolute, rarely induplicate, descending-cochlear or open) aestivation, connate into campanulate, hypocrateromorphic, urceolate or infundibuliform, often bilabiate corolla, sometimes spurred. Nectariferous disc intrastaminal, usually annular (sometimes cupular, rarely unilateral), entire or lobed (occasionally consisting of separate glands), often developed on abaxial side only (rarely absent).

Androecium Stamens usually two longer and two shorter (sometimes two, rarely five, six or up to 16), haplostemonous, antesepalous, alternipetalous. Filaments usually free from each other (rarely connate), usually adnate to corolla tube (epipetalous). Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (sometimes disporangiate or synthecal), usually extrorse or introrse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Placentoid often present. Tapetum secretory or amoeboid-periplasmodial. Staminodia one to three or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 2–3(–9)-colpate, -porate, -colporate or -colporoidate (sometimes syncolpate, sometimes with few or numerous pseudocolpi; rarely monocolpate, pororate, pantoporate, inaperturate, pericolpate, stephanocolporate or spiraperturate), usually shed as monads (rarely as tetrads), bicellular or tricellular at dispersal. Exine tectate or semitectate (rarely intectate), with usually columellate (rarely granular) infratectum, reticulate, microperforate, perforate, punctate, spinulate, echinate, gemmate, rugulate, foveolate, areolate, psilate, striate, granulate, or smooth. Pollen tube with or without callose plugs.

Gynoecium Pistil composed of two (to five) connate carpels. Ovary usually superior (sometimes semi-inferior, rarely inferior), bilocular (to quinquelocular; sometimes unilocular or pseudomonomerous), sometimes with locules divided by secondary septa. Style single, usually terminal (sometimes gynobasic), simple or bilobate. Stigma capitate, clavate, fusiform, bilobate or quadrilobate, or stigmas punctate, papillate or non-papillate, usually Dry or Wet type. Pistillodium absent.

Ovules Placentation usually basal to axile (sometimes apical, free central or intrusively parietal). Ovules (one or) two to numerous per carpel, anatropous, hemianatropous, amphitropous or campylotropous (rarely orthotropous), ascending or pendulous, apotropous or epitropous, unitegmic, tenuinucellar (rarely reduced tenuinucellar). Hypostase often present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Antipodal cells sometimes proliferating, sometimes persistent. Endosperm development ab initio usually cellular (rarely nuclear). Endosperm haustorium chalazal and/or micropylar (often large; sometimes absent). Embryogenesis onagrad, asterad or solanad (occasionally chenopodiad or caryophyllad).

Fruit A loculicidal or septicidal capsule (sometimes with replum; rarely a pyxidium), a drupe, or a schizocarp with usually four one-seeded (sometimes two two-seeded) usually nutlike (rarely drupaceous) mericarps, with persistent and sometimes accrescent calyx (rarely a samara).

Seeds Aril usually absent. Exotestal cells often elongate, sometimes palisade, usually with radial and often inner walls thickened. Hypodermal cells sometimes sclerenchymatous. Endotesta? Perisperm not developed. Endosperm usually sparse or absent (sometimes copious, rarely ruminate), oily. Suspensor often long and narrow. Embryo usually straight (sometimes curved), oily (rich in linolic acid or linolenic acid), well differentiated, usually without chlorophyll. Cotyledons two, often incumbent. Radicula usually directed downwards. Germination phanerocotylar or cryptocotylar.

Cytology x = (3–)5–13, 15 – Protein bodies (of various types) often present in mesophyll cell nuclei.

DNA Often a deletion in the plastid gene matK. Mitochondrial intron coxII.i3 lost. Duplication of the nuclear genes FLO=LFY and DEF=AP3 may have taken place within or prior to the evolution of Orobanchaceae, Paulowniaceae, Phrymaceae, and Verbenaceae, but obviously not Oleaceae and Plantaginaceae, although data are unsufficient.

Phytochemistry Flavonols (kaempferol, quercetin), methyl flavones and oxygenated flavones (8-hydroxyapigenin, apigenin, luteolin, 6- and 8-hydroxyluteolin, 6- and/or 8-hydroxylated flavone glycosides, 6- or 8-hydroxyflavones, or 6-methoxyflavones), Route II iridoids (also C4-decarboxylated iridoids, i.e. iridoid glycosides and glycosides of monoterpenoid lactones), Group I carbocyclic iridoids (aucubin, catalpol, melittoside, globularifolin, daphylloside, monotropein), Group II carbocyclic iridoids (galiridoside, caryoptoside, ipolamiide, lamalbide, ajugol, ajugoside, harpagide, harpagioside, procumbide, lamiol, lamiide, lamioside, antirrhinoside, linarioside), Group III carbocyclic iridoids (cornine, hastatoside), Group IV carbocyclic iridoids (forsythiide, kingiside), Group VII secoiridoids (swertiamarin, gentiopicroside), Group VIII secoiridoids (oleuropein), Group IX secoiridoids (indole alkaloids of corynantheane type), Group X secoiridoids (loganin, ketologanin, desoxyloganin, antirrhide, gardoside, nepeta lactones, iridoid pyridine alkaloids), iridoid aldehydes, iridoid glycosides (usually C10 and C11 iridoid glycosides, rarely C8 iridoid glycosides), ethereal oils consisting of monoterpenoids, sesquiterpenoids, phenylpropanoids, etc., diterpenes, diterpenoids (labdanes, neoclerodanes, abietanes, primaranes, ent-kaurans, etc.), triterpenoids, free terpenic acids, ursolic acid, caffeic acid, ursolic acid and caffeic acid esters (acteosides, caffeoyl phenylethanoid glucosides, e.g. verbascosides, orobanchoside, cornosides, martynoside, oleoside, bartsioside, rosmarinic acid), pyrrolizidine alkaloids as macrocyclic diesters (at least in Castilleja and Melampyrum), syringin, triterpene saponins, coumarins, p-coumaride, betaines, hydroxycinnamic acids (ferulic acids) and their derivatives (disaccharide esters of rosmarinic acid, caffeic acid etc.), coniferin, silicic acid, shikimic acid derived arthroquinones, and cyanogenic compounds (e.g. cyanogenic glycosides) present. Ellagic acid, tannins and proanthocyanidins not found. Carbohydrates usually stored as oligosaccharides (i.a. stachyose, mannitol, hexites, saccharose). Cell walls sometimes containing arabinoxyloglucans. Cornoside and/or verbascoside often replacing iridoid glucosides.

Systematics Awaiting a conservation of the name Lamiales now in common use, I have provisionally chosen one of the oldest legitimate names available, according to the ICBN Art. 16B.1: “In choosing among typified names for a taxon above the rank of family, authors should generally follow the principle of priority.”<

Plantaginales may be sister-group to Solanales, although the bootstrap support is usually low.

Plantaginales with the exception of Plocosperma have the following potential synapomorphies, according to Stevens (2001 onwards): head of glandular hairs having cells with exclusively vertical walls (vertical cell divisions); and flowers tetramerous (frequent reversals to pentamerous). Carlemanniaceae and Oleaceae share the potential synapomorphies tetramerous flowers; two stamens; tricolpate pollen grains; clavate stigma; persistent endothelium; and palisade exotestal cells. Tetramerous flowers are also present in Tetrachondraceae and Calceolariaceae, although here due to parallel evolution.

A deletion in the plastid gene matK has occurred in the immediate ancestor of the remaining Plantaginales (i.e. including Tetrachondraceae and subsequent lineages). An additional potential synapomorphy is: corolla and stamens simultaneously initiated, or androecium initiated prior to corolla. It is not known whether the shift to anthraquinone formation from shikimic acid metabolism occurred in the immediate common ancestor of Tetrachondraceae and the remaining Plantaginales, since Polypremum and Tetrachondra have not been investigated for these substances.

The core Plantaginales, i.e. the clade [[Calceolariaceae+Peltanthera+Gesneriaceae]+the remainder], have the following potential synapomorphies: flowers vertically zygomorphic; flowers pentamerous (a reversal); calyx asymmetrical; corolla bilabiate, with upper lip bilobate and lower lip trilobate (2:3 type); aestivation ascending cochlear (i.e. adaxial lobes outside the remainder); corolla tube formation late; stamens usually four, didynamous; anthers connivent (not in, e.g. Calceolariaceae); pollen tubes without callose; ovules numerous per carpel; endosperm haustoria both chalazal and micropylar; suspensor large; cell nuclei with lamellar protein bodies; presence of shikimic acid derived anthraquinones; presence of 6- and/or 8-hydroxylated (6- and/or 8-oxygenated) flavone glycosides (flavones); and stachyose and other oligosaccharides storage carbohydrates.

The clade [Calceolariaceae+Peltanthera+Gesneriaceae] is characterized by the synapomorphies: relatively soft leaves; cymes with paired flowers; endothelial cells arranged in longitudinal rows; alveolated seeds; longitudinally furrowed (aulacospermous) endosperm; presence of cornoside; and absence of iridoids (Stevens 2001 onwards; Schäferhoff & al. 2010).

The lineages above the [Calceolariaceae+Peltanthera+Gesneriaceae] clade share the features: presence of 6- or 8-hydroxyflavones or 6-methoxyflavones and Route II decarboxylated iridoids (aucubin and catalpol frequent); and usually absence of cornosides.

The clade [Mazaceae+[Phrymaceae+[Paulowniaceae+[Rehmanniaceae+[Cyclocheilaceae+ Orobanchaceae]]]]] have racemose inflorescence; and, sometimes, presence of protein crystal stacks in cell nuclei. Alveolated seeds are present in Rehmanniaceae and Orobanchaceae (also in some Linderniaceae and Scrophulariaceae) (Schäferhoff & al. 2010).

Acanthaceae, Bignoniaceae, Lentibulariaceae, Martyniaceae, Pedaliaceae, Schlegeliaceae, Thomandersiaceae, and Verbenaceae form an almost unresolved polytomy in the analyses by Schäferhoff & al. (2010). Martyniaceae and Schlegeliaceae were recovered as sister-groups with weak support. On the other hand, Oxelman & al. (2005) identified a monophyletic group comprising Martyniaceae, Schlegeliaceae and Verbenaceae, and Wortley & al. (2007) found Schlegeliaceae to be sister to Thomandersia yet with low support.

Capturing prey (often including carnivory) have evolved three or perhaps four times in Plantaginales, i.e. in Philcoxia, Byblis and Lentibulariaceae, and possibly in some Martyniaceae. Carnivory is known from the three genera in Lentibulariaceae and in a single genus, Philcoxia, in Plantaginaceae. In Byblis and possibly in two genera in Martyniaceae, the plant merely traps insects; carnivory has not been demonstrated for these three genera.

Plantaginales incertae sedis

Brookea (4; B. albicans, B. auriculata, B. dasyantha, B. tomentosa; Borneo) consists of shrubs or small trees with densely tomentose opposite leaves. The flower has a zygomorphic corolla and four stamens. – Brookea may belong in Plantaginaceae.

Cubitanthus (1; C. alatus; Bahia in Brazil) is a perennial herb with multicellular hairs, four-winged stem, opposite leaves with serrate margins, solitary axillary flowers without bracteoles, persistent calyx with free sepals, bilabiate corolla limb with trilobite lower lip, four didynamous stamens adnate to corolla base, annular nectariferous disc, superior ovary with parietal placentae, capitate stigma, septicidal bivalvular capsule, and striate seed surface. – It was placed in Gesneriaceae, although it is more similar to Linderniaceae (Perret & al. 2013), Plantaginaceae.

Trungboa (1; T. poilanei; Southeast Asia) possibly belongs in Plantaginaceae.

[Peltanthera+Calceolariaceae+Gesneriaceae]

Peltanthera

1/1. Peltanthera (1; P. floribunda; Guatemala, Panamá, Ecuador, Peru, Bolivia). – Bisexual evergreen small tree or shrub. Phellogen? Vessel elements? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma? Sieve tube plastids S type? Nodes 3:3, trilacunar with three leaf traces. Crystals? Hairs branched moniliform. Leaves opposite, simple, with involute ptyxis. Petiole vascular bundle transection flattened annular; petiole with mid-vein bundles (and sometimes medullary bundles). Stipules and leaf sheath absent. Venation pinnate. Stomata? Cuticular wax crystalloids? Leaf margin serrate. Inflorescence axillary, thyrsoid panicles. Flowers actinomorphic to somewhat zygomorphic. Hypogyny. Sepals five, with valvate aestivation, connate at base. Petals five, with valvate aestivation, connate. Nectariferous disc intrastaminal. Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers basifixed, versatile?, tetrasporangiate, introrse?, longicidal, with confluent thecae. Tapetum? Staminodia absent. Pistil composed of two connate carpels. Ovary superior, bilocular. Style single, entire. Stigma bifid, type? Pistillodium absent. Placentation axile. Ovules numerous per carpel, anatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis? Fruit a loculicidal capsule. Aril absent. Exotestal cells? Endotestal cells? Perisperm not developed. Endosperm? Embryo? Cotyledons? Germination? n = ? DNA? Verbascosides and cornoside derivatives present. – Peltanthera may be sister to [Calceolariaceae+Gesneriaceae] (Soltis & al. 2011, Weber 2013, Refulio-Rodriguez & Olmstead 2014).

Phylogeny of Plantaginales based on DNA sequence data (mainly according to Schäferhoff & al. 2010). Soltis & al. (2011) and Refulio-Rodriguez & Olmstead (2014) recovered the clade [Peltanthera+[Calceolariaceae+Gesneriaceae]] as sister-group to the remainder (i.e. except Plocosperma, Carlemanniaceae and Oleaceae), with a bootstrap support of 97%. Likewise, Andersson (2006) identified Calceolariaceae as sister to Gesneriaceae. Verbenaceae may be closely related to Thomandersia (Refulio-Rodriguez & Olmstead 2014).

ACANTHACEAE Juss.

( Back to Plantaginales )

de Jussieu, Gen. Plant.: 102. 4 Aug 1789 [’Acanthi’], nom. cons.

Acanthales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 246. Jan-Apr 1820 [‘Acanthaceae’]; Acanthineae Link, Handbuch 1: 500. 4-11 Jul 1829; Justiciaceae Raf., Fl. Tellur. 4: 60. med 1838 [’Justicoides’]; Avicenniaceae (Endl.) Miq. in J. G. C. Lehmann, Plant. Preiss. 1: 353. 14-16 Aug 1845 [’Avicennieae’], nom. cons.; Thunbergiaceae (Dumort.) Lilja, Skånes Fl., ed. 2: 979. Apr-Dec 1870; Mendonciaceae Bremek. in Proc. Kon. Ned. Akad. Wetensch., ser. C, 56: 540. 27 Apr 1954; Meyeniaceae Sreem. in Phytologia 37: 412. 22 Oct 1977; Nelsoniaceae (Nees) Sreem. in Phytologia 37: 412. 22 Oct 1977

Genera/species c 203/3.470–>3.570

Distribution Tropical regions, especially in South and Southeast Asia, Africa, Brazil and Central America, some species in warm-temperate regions, mangrove vegetation in tropical and subtropical regions on both hemispheres.

Fossils Fossil pollen grains similar to Acanthaceae have been reported from the Miocene in many places in Africa, South America and Southeast Asia. A fossilized seed from Late Eocene of England and assigned to Acanthus rugatus possibly belong in Acanthaceae. Fossils assigned to Ruellia have been found in Eocene layers in, e.g. Spain and California. Plausible acanthaceous pollen fossils (e.g. Areolipollis insularis) were found in Miocene layers of Nigeria and Mexico.

Habit Usually bisexual (rarely unisexual), usually perennial or annual herbs, evergreen shrubs or lianas (rarely evergreen trees; Avicennia consists of mangrove trees or mangrove shrubs with articulated branchlets). Some representatives are aquatic. Many species are xerophytic. Some species are epiphytic.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes deeply seated). Medulla sometimes with inverted vascular bundles. Primary medullary rays narrow. Secondary lateral growth usually normal (sometimes anomalous, via concentric cambia or cylindrical cambium, occasionally via internal inverted cambium). Endodermis in Andrographis, Barleria and Thunbergia prominent. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres usually with simple (sometimes bordered) pits, septate or non-septate (also acicular fibres and vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually paratracheal scanty, sometimes vasicentric (in Avicennia apotracheal or paratracheal, also aliform, lozenge-aliform or confluent). Intraxylary (concentric) phloem sometimes present. Sieve tube plastids S type, Pc type or Pcs type. Nodes usually 1:1?, unilacunar with one? leaf trace (in Avicennia 3:3?, trilacunar with three? traces), often swollen. Cystoliths – outgrowths of epidermal cell wall, impregnated with calciumcarbonate – abundant as lines (absent in Nelsonioideae, Mendoncia and Anomacanthus). Heartwood in Avicennia with gum-like substances. Calciumoxalate as druses or prismatic crystals present in Avicennia.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, stellate, candelabra-like, dendritic; glandular hairs, stalked to almost sessile, abundant.

Leaves Usually opposite, simple, entire or pinnately lobed, sometimes coriaceous, with ptyxis in Thunbergioideae strongly curved. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate. Stomata diacytic or paracytic. Cuticular wax crystalloids? Mesophyll with or without calciumoxalate as druses or single prismatic crystals (rarely raphides). Leaf margin serrate, crenate or entire. Glandular hairs with ethereal oils. Salt glands present, i.a., in Avicennia. Extrafloral nectaries present on lamina in many species in, e.g. Ruellia.

Inflorescence Terminal or axillary, usually dichasial to monochasial (in Avicennia thyrsoid, spike- or umbel-like) cymose (sometimes spike or raceme, or flowers solitary axillary). Bracts and/or floral prophylls (bracteoles) often large and showy (prophylls absent in Nelsonia), petaloid. Inflorescence often with involucre of large bracts surrounding partial inflorescences. Extrafloral nectaries rarely present on floral prophylls and/or pedicels.

Flowers Zygomorphic (flowers sometimes partially or entirely inverted; in Avicennia almost actinomorphic). Hypogyny. Sepals (three to) five (to 16), with imbricate, contorted, valvate or open aestivation, often with acute apex, persistent, free or connate (in Thunbergioideae usually strongly reduced); when five, then median sepal adaxial (sepals in Neuracanthus connate 3+2). Petals usually five, usually with imbricate quincuncial, contorted or ascending-cochlear (rarely open; in Nelsonioideae descending-cochlear) aestivation, more or less connate into quinquelobate bilabiate corolla (upper lip bilobate, lower lip trilobate, with lobes sometimes narrow; upper lobes sometimes reduced; petals sometimes three or four). Nectariferous disc intrastaminal, annular or as glands (rarely absent).

Androecium Stamens usually two (in Nelsonioideae two adaxial-lateral) or two long and two short (didynamous; rarely two fertile and two staminodial; in Pentstemonacanthus five fertile), haplostemonous, antesepalous, alternipetalous. Filaments free from each other or connate in pairs, adnate to corolla tube (epipetalous). Anthers often connivent, dorsifixed, versatile, tetrasporangiate, introrse to extrorse, longicidal (dehiscing by longitudinal slits); connective sometimes prolonged into appendage; thecae often asymmetrically arranged. Tapetum secretory. Staminodia usually one (adaxial-median; when four fertile stamens) or three (one adaxial-median and two adaxial-lateral; when two fertile stamens); staminodia absent in, e.g., Nelsonioideae and Avicennia.

Pollen grains Microsporogenesis simultaneous. Pollen morphology very varying. Pollen grains 2–3(–9)-colpate, -porate, -colporate or -colporoidate (sometimes syncolpate, sometimes with few or numerous pseudocolpi; rarely pororate, pantoporate, inaperturate or spiraperturate), shed as monads, bicellular or tricellular at dispersal. Exine tectate or semitectate (rarely almost intectate), with columellate infratectum, reticulate, microreticulate, perforate, punctate, spinulate, echinate, gemmate, rugulate, foveolate, psilate, striate or smooth (sometimes with raised tectal areas or ridges/ribs).

Gynoecium Pistil composed of two connate carpels; carpels without septal vascular bundles. Ovary superior, usually bilocular (in Mendoncia unilocular due to pseudomonomery; primary locules in Avicennia divided at base by secondary septa). Style single, simple, narrow. Stigma bifid (adaxial lobe often smaller), trumpet-shaped etc., usually non-papillate, Dry type (sometimes papillate, Wet type). Pistillodium absent.

Ovules Placentation usually axile (rarely intrusively parietal; in Avicennia free central). Ovules usually two to more than ten (rarely one) per carpel, anatropous, hemianatropous, amphitropous or campylotropous, unitegmic, tenuinucellar. Funicle usually modified into hard hooked ejaculator (retinaculum). Integument ? cell layers thick. Micropyle absent in Avicennia. Megagametophyte monosporous, Polygonum type, long and curved (apex of quadrinucleate megagametophyte sometimes expanding and reaching outside micropyle into placenta where egg apparatus develops; an extremely long suspensor pushes back developing embryo into endosperm). Antipodal cells sometimes proliferating (up to four to 18 cells), sometimes persistent (not formed in Avicennia). Endosperm development cellular; endosperm asymmetrical, two haustoria becoming closely adjacent to each other (cf. Lamiaceae-Nepetoideae). Endosperm haustoria chalazal and micropylar. Embryogenesis onagrad or solanad. Polyembryony frequent in some genera.

Fruit Usually a loculicidal explosion capsule (in Mendoncia and closely allied genera a one- or two-seeded drupe) with usually cartilaginous walls and persistent calyx. Funicle usually provided by hook-shaped lignified ejaculators, retinacula, explosively throwing out seeds from dehiscing capsule (appendage in Nelsonioideae and Thunbergioideae as papillae or absent). Retinaculum consisting of persistent and during maturation lignified funicle, forming hook below each seed.

Seeds Aril usually absent. Seed coat exotestal. Exotesta usually palisade, often mucilaginous as wet (hygroscopical hairs). Endotesta? Perisperm not developed. Endosperm usually very sparse or absent (in, e.g., Avicennia and some Nelsonioideae copious, ruminate due to localized asymmetrical growth of endosperm cells, oily). Embryo small to large, straight or curved, well differentiated, usually without chlorophyll (in Avicennia with chlorophyll). Cotyledons two, planoconvex, plicate or crumpled (in Avicennia large), often with amyloid. Germination phanerocotylar. Avicennia viviparous.

Cytology n = 9 (Nelsonioideae); n = 9, 28 (Thunbergioideae); n = 18, 32 (Avicennia); n = very various (Acanthoideae; 13, 15, 30, 40 etc.)

DNA Mitochondrial coxI intron present. Deletion in plastid gene matK?

Phytochemistry Flavonols (kaempferol, quercetin), C8-iridoid glycosides (thunbergioside, stilbericoside; in Thunbergia), caffeic acid esters (verbascosides), alkaloids, saponins, cyanogenic compounds, shikimic acid derived arthroquinones (in Barleria), and quaternary methylammonium compounds present. Ellagic acid and proanthocyanidins not found. Iridoids?

Use Ornamental plants, timber, tanning of leather (Avicennia).

Systematics (under construction) The sister-group relationship of Acanthaceae is unresolved.

Either Elytraria or the entire Nelsonioideae may be sister-group to the remaining Acanthaceae. Avicennia is probably sister to Thunbergioideae (or Acanthoideae?).

An endosperm development similar to that in Acanthaceae occurs in Lamiaceae-Nepetoideae – a parallelism.

A probable topology is [Nelsonioideae+[Acanthoideae+[Thunbergioideae+Avicennia]]].

The systematics below mainly follows McDade & al. (2008).

Phylogeny (simplified) of Acanthaceae based on DNA sequence data (McDade & Moody 1999; Schwarzbach & McDade 2002; McDade & al. 2008; Borg & al. 2008). Neuracanthus as sister to the clade [Whitfieldieae+[Barlerieae+Andrographideae]] is weakly supported.

Nelsonioideae Lindl. ex Pfeiff., Nomencl. Bot. 1(1): 10. ante 8 Dec 1871 [‘Nelsonieae’]

7/105–110. Nelsonia (1; N. canescens; tropical Africa, Madagascar, tropical Asia to tropical Australia), Elytraria (17–21; tropical and subtropical regions on both hemispheres), Saintpauliopsis (1; S. lebrunii; tropical Africa), Anisosepalum (3; A. alboviolaceum, A. humbertii, A. lewallei; Central Africa), ‘Staurogyne’ (c 80; tropical regions on both hemispheres; paraphyletic), Gynocraterium (1; G. guianense; tropical South America; in Staurogyne?), Ophiorrhiziphyllon (2; O. diandrum, O. macrobotryum; Burma, southern China, Thailand, Indochina; in Staurogyne?). – Tropical and subtropical regions on both hemispheres. Herbs. Acicular fibres present. Cystoliths absent. Glandular hairs with bicellular head. Leaves opposite to alternate (spiral). Inflorescence usually a raceme (inflorescence branches in Saintpauliopsis cymose). Bracts spirally arranged. Nelsonia without floral prophylls (bracteoles). Corolla with descending cochlear aestivation (adaxial lobes overlapping abaxial lobes). Stamens two or four. Pollen grains usually tricolpate or tricolporate. Anthers of various shape. Stigma broadly (sometimes unequally) lobate, in Elytraria sensitive. Placentation in Elytraria parietal. Ovules usually six to numerous (rarely four or fewer) per carpel, campylotropous. Funicular obturator present. Endothelium present. Antipodal cells persistent. Endosperm development with cellular central area. Ejaculators (retinacula) rudimentary or absent. Seeds two to numerous, ruminate. Testa dissolved. Endosperm present, oily. n = 9. – It is uncertain whether Nelsonioideae have ejaculators (retinacula), but in that case they are non-functioning.

[Acanthoideae+[Thunbergioideae+Avicennia]]

Medulla sometimes with inverted vascular bundles. Acicular fibres present. Corolla sometimes with descending cochlear aestivation (adaxial corolla lobes outside the others). Ovules usually four (sometimes more) per carpel, collateral. Funicular obturator at least usually absent. Endothelium absent. Endosperm usually absent. Cotyledons often with amyloid (xyloglucans).

Acanthoideae Eaton, Bot. Dict., ed. 4: 33. Apr-Mai 1836 [‘Acanthaceae’] (under construction)

c 190/3.190–>3.280. Tropical regions, especially in South and Southeast Asia, Africa, Brazil and Central America, some species in warm-temperate regions. Usually herbs (rarely shrubs). Nodes often swollen; internode part immediately above node collapsing when dry. Cystoliths usually present (absent in Acantheae). Petiole vascular bundle transection usually arcuate, with bundles arranged in circle (rarely annular). Leaf margin sometimes spinose-serrate. Floral prophylls (bracteoles) often prominent. Calyx lobes often narrow. Corolla sometimes with contorted or descending cochlear aestivation (with abaxial lobes overlapping adaxial lobes); lobes sometimes narrow. Anthers sagittate or with thecae displaced, not opposite; anthers often monothecal (with one theca reduced). Pollen grains very variable, often porate. Stigma usually bilobate, Dry type. Fruit an explosive, almost woody capsule. Seeds several, flattened, sometimes hairy, inserted on ejaculators, retinacula, i.e. hook-shaped lignified modified funicles dispersing seeds when mature. Exotesta palisade, often mucilaginous as moistened (due to hygroscopic hairs). Hypodermal cells sometimes thickened. Cytologically very variable (n = 13, 15, 30, 40 etc.). – A possible topology of this “Retinaculate clade” is the following: [Acantheae+[[Neuracanthus+[Whitfieldia clade+[Barlerieae+Andrographideae]]]+[Ruellieae+Justicieae]]].

Phylogeny (simplified) of Acanthoideae based on DNA sequence data (McDade, Daniel & Kiel 2008).

Acantheae Dumort., Anal. Fam. Plant.: 23. 1829

20/495–505. “Unilabiate Corolla LineageCrossandra (c 55; tropical Africa, Madagascar, the Arabian Peninsula, tropical Asia), Crossandrella (3; C. adamii, C. dusenii, C. laxispicata; tropical Africa), Sclerochiton (19; tropical and southern Africa), Streptosiphon (1; S. hirsutus; Tanzania), Cynarospermum (1; C. asperrimum; the West Indies), Blepharis (125–130; the Mediterranean, tropical regions in the Old World and southwards to South Africa), Acanthopsis (8; Namibia, South Africa), Acanthus (29; warm-temperate to tropical regions in the Old World). – “Bilabiate Corolla LineageStenandrium (c 45; Africa, Madagascar, tropical and subtropical America), Salpixantha (1; S. coccinea; Jamaica), Holographis (16; Mexico), Neriacanthus (3; N. grandiflorus, N. harlingii, N. nitidus; tropical America), Rhombochlamys (1; R. elata; Colombia)?, Encephalosphaera (3; E. lasiandra, E. puberula, E. vitellina; tropical South America), Geissomeria (17; Mexico, Central America, tropical South America), ‘Aphelandra’ (170–175; tropical America; paraphyletic). – Unplaced Acantheae Cyphacanthus (1; C. atopus; Colombia), Orophochilus (1; O. stipulaceus; Peru), Strobilacanthus (1; S. lepidospermum; Panamá)?, Xantheranthemum (1; X. igneum; the Andes in Peru). – Pantropical, few species in warm-temperate regions. Cystoliths absent. Nodes not swollen. Corolla usually with imbricate quincuncial aestivation. Anthers monothecal. Pollen grains colpate or biporate girder pollen. Seeds usually without hygroscopic hairs (present in Blepharis). – Taxa with unilabiate (0:5) and bilabiate (2:3) corolla form different clades (McDade & al 2005).

Phylogeny (simplified) of Acantheae based on DNA sequence data (McDade & al. 2005).

[[Neuracanthus+[Whitfieldia clade+[Barlerieae+Andrographideae]]]+[Ruellieae+Justicieae]]

Cystoliths present. Nodes swollen. Anthers dithecal. Pollen grains porate. Endosperm development often unique, with a central area with free nuclear divisions, cell walls being established subsequently (sometimes with basal apparatus, i.e. area without cell wall development). Seeds usually with hygroscopic hairs.

[[Neuracanthus+[Whitfieldia clade+[Barlerieae+Andrographideae]]]

The “BAWN clade”.

Neuracanthus clade

1/13. Neuracanthus (13; tropical and subtropical East Africa, Madagascar, southern Arabian Peninsula, India to Indochina). – Inflorescence compact, with overlapping strongly-veined bracts (sometimes with spines consisting of modified partial inflorescences). Calyx two-lipped (upper lip trilobate, lower lib bilobate). Pollen grains tricolporate. Exine perforate, with interapertural regions of exine foveolate. Seeds with hygroscopic hairs.

[Whitfieldia clade+[Barlerieae+Andrographideae]]

Whitfieldia clade

8/c 38. Lankesteria (7; L. alba, L. barteri, L. brevior, L. elegans, L. glandulosa, L. hispida, L. thyrsoidea; tropical Africa, Madagascar), Whitfieldia (c 15; tropical Africa), ’Chlamydacanthus’ (4; C. dichrostachyus, C. euphorbioides, C. lindavianus, C. rupestris; tropical East Africa; non-monophyletic), Leandriella (2; L. oblonga, L. valvata; Madagascar), Camarotea (2; C. romiensis, C. souiensis; Madagascar), Zygoruellia (1; Z. benthamii; Madagascar), Forcipella (6; F. bosseri, F. cleistochlamys, F. involucrata, F. longistaminea, F. madagascariensis, F. repanda; Madagascar), Vindasia (1; V. virgata; Madagascar). – Tropical Africa, Madagascar. Corolla with contorted (Lankesteria, Whitfieldia) or ascending-cochlear aestivation. Vascular traces to corolla lobes trifurcating. Stamens four. Pollen grains lenticular, biporate, with typical girder-like band, and circular densely granular areas surrounding pores. Seeds usually covered with concentric rings of ridges of coarse scales, usually without hygroscopic hairs (present in Lankesteria).

[Barlerieae+Andrographideae]

Barlerieae Nees in Martius, Fl. Bras. 9: 7, 65. 1 Jun 1847

12/365–370. Barleria (250–255; tropical regions on both hemispheres), Barleriola (4; B. inermis, B. multiflora, B. satureioides, B. solanifolia; the West Indies), Borneacanthus (6; B. angustifolius, B. grandifolius, B. mesargyreus, B. paniculatus, B. parvus, B. stenothyrus; Borneo), Boutonia (1; B. cuspidata; Madagascar), Chroesthes (6; C. bracteata, C. lanceolata, C. longifolia, C. pubiflora, C. acemiflora, C. silvicola; southern China, Southeast Asia), Crabbea (14; tropical and southern Africa), Hulemacanthus (2; H. densiflorus, H. novoguineensis; New Guinea), Lasiocladus (2; L. anthospermifolius, L. rufopilus; Madagascar), Lepidagathis (c 70; tropical and subtropical regions on both hemispheres), Pericalypta (1; P. biflora; Madagascar), Podorungia (5; P. clandestina, P. decaryi, P. humblotii, P. lantzei, P. serotina; Madagascar), Pseudodicliptera (2; P. coursii, P. longifolia; Madagascar). – Tropical and subtropical regions. Corolla usually with imbricate quincuncial aestivation. Seeds usually with hygroscopic hairs.

Andrographideae Endl., Gen. Plant.: 707. Jan 1839

6/105–110. Andrographis (30–35; tropical Asia), Graphandra (1; G. procumbens; Thailand), Gymnostachyum (c 50; India to Central Malesia), Haplanthodes (4; H. neilgherrensis, H. plumosa, H. tentaculatus, H. verticillatus; India), Haplanthus (4; H. hygrophiloides, H. laxiflorus, H. ovatus, H. rosulatus; India, Bangladesh, eastern Himalayas, Burma, southern China, Southeast Asia, Malesia), Phlogacanthus (c 15; tropical Asia). – Tropical Asia. Corolla with ascending cochlear aestivation. Pollen grains tripororate or colporate. Exine thickened at aperture margins. Ovules usually six or more per ovary. Seeds without hygroscopic hairs.

[Ruellieae+Justicieae]

Ruellieae Dumort., Anal. Fam. Plant.: 23. 1829

c 45/995–1.030. Erantheminae Nees in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 11: 425. 27 Nov 1847. Brunoniella (6; B. acaulis, B. australis, B. linearifolia, B. neocaledonica, B. pumilio, B. spiciflora; New Guinea, northern and eastern Australia, New Caledonia), Leptosiphonium (11; New Guinea), Eranthemum (20–25; tropical Asia), Kosmosiphon (1; K. azureus; Cameroon), Pararuellia (3; P. alata, P. hainanensis, P. sumatrensis; southern China, Southeast Asia, Malesia). – Trichantherinae Benth. et Hook. f., Gen. Pl. 2: 1062, 1064. 1-16 Mai 1876. Louteridium (9; southern Mexico, Central America), Bravaisia (3; B. berlandieriana, B. grandiflora, B. integerrima; tropical America), Trichanthera (2; T. corymbosa, T. gigantea; northern South America), Trichosanchezia (1; T. chrysothrix; eastern Peru), Suessenguthia (4; S. cuscoenis, S. multisetosa, S. trochilophila, S. vargasii; eastern Andes in Peru and Bolivia), Sanchezia (c 55; tropical America, with their largest diversity in the Andes). – Hygrophilinae Nees in N. Wallich, Pl. Asiat. Rar. 3: 75. 15 Aug 1832. ‘Hygrophila’ (c 45; tropical regions on both hemispheres; paraphyletic; incl. Brillantaisia?), Brillantaisia (15–17; tropical Africa, Madagascar; in Hygrophila?). – Mimulopsinae E. Tripp in Int. J. Pl. Sci. 174(1): 111. Jan 2013. ‘Heteradelphia’ (2; H. paulojaegeria, H. paulowilhelmia; tropical West and Central Africa; paraphyletic; incl. Eremomastax?), Eremomastax (1; E. speciosa; tropical West and Central Africa, Madagascar?; in Heteradelphia?), ‘Mellera’ (5; M. lobulata, M. mentiodora, M. nyassana, M. parvifolia, M. submutica; tropical and subtropical Africa; paraphyletic; incl. Ionacanthus?), Ionacanthus (1; I. calcaratus; Madagascar; in Mellera?), Mimulopsis (15–30; tropical Africa, Madagascar). – Petalidiinae Benth. et Hook. f., Gen. Pl. 2: 1062, 1064. 1-16 Mai 1876. Ruelliopsis (2; R. damarensis, R. setosa; tropical and southern Africa), Phaulopsis (18; tropical Africa), Petalidium (c 35; tropical and southern Africa, western India, western Himalayas), Duosperma (26; tropical and southern Africa), Strobilanthopsis (1; S. linifolia; tropical Africa), Sautiera (1; S. tinctorum; Timor), Dyschoriste (c 100; tropical and subtropical regions on both hemispheres). – Strobilanthinae T. Anderson in J. Linn. Soc. London, Bot. 9: 443. 6 Apr 1866. Sinoacanthus (1–3; southern China, northern Vietnam), Hemigraphis (20–25; tropical Asia), ‘Strobilanthes’ (165–170; tropical Asia; paraphyletic). – Ruelliinae Nees in N. Wallich, Pl. Asiat. Rar. 3: 75. 15 Aug 1832. Dischistocalyx (13; tropical Africa), Satanocrater (2; S. paradoxa, S. ruspolii; tropical Africa), Acanthopale (9; tropical regions in the Old World), ‘Ruelliaprimuloides (tropical West and Central Africa), Benoicanthus (2; B. gruicollis, B. tachiadenus; Madagascar; in Ruellia?), Pseudoruellia (1; P. perrieri; Madagascar), ‘Ruellia’ (c 270; tropical and subtropical regions on both hemispheres, eastern United States, Argentina; paraphyletic), Eusiphon (3; E. geayi, E. longissimum, E. longistamineum; Madagascar; in Ruellia?), Polylychnis (1; P. fulgens; northeastern South America; in Ruellia?), Blechum (14; tropical America), Lychniothyrsus (1; L. mollis; Brazil). – Unplaced Ruellieae Calacanthus (1; C. grandiflorus; tropical Asia), Diceratotheca (1; D. bracteolata; northwestern Thailand), Echinacanthus (4; E. attenuatus, E. lofuensis, E. longipes, E. longzhouensis; the Himalayas, China), Physacanthus (3; P. batanganus, P. nematosiphon, P. talbotii; tropical Africa), Spirostigma (1; S. hirsutissima; Brazil)?, Stenothyrsus (1; S. ridleyi; Perak on the Malay Peninsula)? – Pantropical. Cystoliths present. Corolla with sinistrorsely contorted aestivation. Corolla tube usually obliquely-longitudinally divided by lamellar structure, “filament curtain”, formed from synstapetal part of corolla tube plus filaments (decurrent filament ridges and connate filament parts just above adnate portion). Anthers in Bravaisia with basal appendages. Ovules sometimes more than four per carpel. Seeds often with hygroscopic hairs.

Justicieae Dumort., Anal. Fam. Plant.: 23. 1829

c 92/1.170–>1.200. Pseuderanthemum clade: Spathacanthus (3; S. hahnianus, S. hoffmannii, S. parviflorus; Central America), Herpetacanthus (12; Panamá to Brazil), Asystasia (c 55; tropical regions in the Old World), Ruttya (6; R. bernieri, R. fragrans, R. fruticosa, R. ovata, R. speciosa, R. tricolor; tropical and southern Africa, Madagascar, Yemen), Ruspolia (5; R. decurrens, R. humbertii, R. hypocrateriformis, R. paniculata, R. seticalyx; tropical Africa, Madagascar), Pseuderanthemum (45–50; tropical regions on both hemispheres), Wuacanthus (1; W. microdontus; Sichuan, Yunnan), Oplonia (17; Madagascar, tropical America, with their highest diversity in the West Indies), Chileranthemum (3; C. lottiae, C. pyramidatum, C. trifidum; Mexico), Mackaya (2; M. bella, M. indica; Swaziland, northern and eastern South Africa, southern Asia); Isoglossinae: Ptyssiglottis (30–35?; tropical Asia), Isoglossa (c 50; tropical regions in the Old World, the Arabian Peninsula), Brachystephanus (22; tropical Africa, Madagascar), Razisea (5; R. citrina, R. ericae, R. spicata, R. villosa, R. wilburii; Central America), Stenostephanus (c 40; tropical South America); Chlamydocardia (3; C. buettneri, C. lanciformis, C. subrhomboidea; tropical West and Central Africa), Kudoacanthus (1; K. albonervosus; Taiwan); 'Justicia' pro parte, Clinacanthus (2; C. nutans, C. spirei; southern China, Southeast Asia, Malesia), Angkalanthus (1; A. oligophylla; Socotra), Chorisochora (3; C. minor, C. striata, C. transvaalensis; Mpumalanga and Northern Province in South Africa, Socotra), Ecbolium (22; tropical regions in the Old World; in Justicia?), Populina (2; P. perrieri, P. richardii; Madagascar), Megalochlamys (10; southwestern tropical Africa to Namibia and northern South Africa, the Arabian Peninsula), Trichaulax (1; T. mwasumbii; Kenya, Tanzania), Hoverdenia (1; H. speciosa; Mexico), 'Yeatesia' (3; Y. laetevirens, Y. platystegia, Y. viridiflora; southeastern United States to northeastern Mexico; polyphyletic), Streblacanthus (5; S. amoenus, S. cordatus, S. dubiosus, S. monospermus, S. roseus; Central America), 'Pachystachys' (12; tropical America; non-monophyletic), Fittonia (2; F. albivenis, F. gigantea; Peru), Ancistranthus (1; A. harpochiloides; Cuba), Schaueria (8; Brazil), Mirandea (6; M. andradenia, M. grisea, M. huastecensis, M. hyssopus, M. nutans, M. sylvatica; Mexico), Henrya (2; H. insularis, H. tuberculosperma; Central America), Aphanospermum (1; A. sinaloense; northwestern Mexico), Chalarothyrsus (1; C. amplexicaulis; Mexico), Gypsacanthus (1; G. nelsonii; Mexico), 'Anisacanthus' (18; southwestern United States, Mexico; polyphyletic), 'Carlowrightia' (c 25; southwestern United States, Mexico, Central America south to Costa Rica; polyphyletic), Tetramerium (30–35; Central America); Rungia (23; tropical regions in the Old World), Metarungia (3; M. galpinii, M. longistrobus, M. pubinervia; tropical and southern Africa; in Anisotes?); Anisotes (19; tropical Africa, Madagascar); Old World ‘Justicia’ (c 600; warm-temperate to tropical regions in the Old World; paraphyletic); Justicia betonica (tropical and southern Africa, tropical Asia); Diclipterinae: Rhinacanthus (10; tropical regions in the Old World), Peristrophe (c 20; tropical regions in the Old World), Hypoestes (100–105; tropical regions in the Old World; in Justicia?), Dicliptera (220–225; tropical and subtropical regions on both hemispheres; in Justicia?). – Unplaced Justicieae Afrofittonia (1; A. silvestris; tropical West and Central Africa), Ambongia (1; A. perrieri; Madagascar), Ascotheca (1; A. paucinervia; Gabon), Ballochia (3; B. amoena, B. atrovirgata, B. rotundifolia; Socotra), Calycacanthus (1; C. magnusianus; New Guinea), Celerina (1; C. seyrigii; Madagascar), Chameranthemum (4; C. beyrichii, C. durandii, C. tonduzii, C. venosum; tropical America), Chlamydostachya (1; C. spectabilis; tropical East Africa), Codonacanthus (3; C. pauciflorus, C. sanjappae, C. spicatus; northeastern India, southern China, Japan), Conocalyx (1; C. laxus; Madagascar), Cosmianthemum (14; western Borneo), Cyclacanthus (2; C. coccineus, C. poilanei; Southeast Asia), Danguya (1; D. pulchella; Madagascar), Dasytropis (1; D. fragilis; eastern Cuba), Dichazothece (1; D. cylindracea; eastern Brazil), Dicladanthera (2; D. forrestii, D. glabra; western Western Australia), Filetia (9; the Malay Peninsula, Sumatra), Glossochilus (2; G. burchellii, G. parviflorus; southern Africa), Graptophyllum (c 15; tropical regions in the Old World, northern and eastern Australia, Melanesia), Ichthyostoma (1; I. thulinii; southeastern Ethiopia, Somalia)?, Isotheca (1; I. alba; Trinidad), Jadunia (1; J. biroi; New Guinea), Juruasia (2; J. acuminata, J. rotundata; Brazil), Kalbreyeriella (3; K. gigas, K. rioquebradasiana, K. rostellata; Panamá, Colombia), Linariantha (1; L. bicolor; Borneo)?, Marcania (1; M. grandiflora; Thailand)?, Megalostoma (1; M. viridescens; Central America), Melittacanthus (1; M. divaricatus; Madagascar), Monothecium (3; M. aristatum, M. glandulosum, M. leucopterum; tropical Africa to southern India), Oreacanthus (3–4; O. coeruleus, O. mannii, O. sudanicus; Central Africa), Pelecostemon (1; P. trianae; Colombia), Phialacanthus (5; P. griffithii, P. major, P. minor, P. pauper, P. wrayi; the Himalayas to the Malay Peninsula)?, Pranceacanthus (1; P. coccineus; Amazonian Brazil), Psilanthele (1; P. eggersii; Ecuador), Pulchranthus (4; P. adenostachyus, P. congestus, P. surinamensis, P. variegatus; tropical South America), Ritonia (4; R. barbigera, R. humbertii, R. poisonii, R. rosea; Madagascar), Samuelssonia (1; S. verrucosa; Hispaniola)?, Sapphoa (2; S. ekmanii, S. rigidifolia; Cuba)?, Sebastiano-schaueria (1; S. oblongata; Brazil)?, Sphinctacanthus (1; S. griffithii; northeastern India to Burma)?, Tessmanniacanthus (1; T. chlamydocardioides; eastern Peru), Thysanostigma (2; T. odontites, T. siamense; southern Thailand, the Malay Peninsula)?, Trichocalyx (2; T. obovatus, T. orbiculatus; Socotra)?, Vavara (1; V. breviflora; Madagascar), Xerothamnella (2; X. herbacea, X. parvifolia; southern Queensland, northwestern New South Wales, northeastern South Australia)? – Pantropical. Corolla with imbricate quincuncial aestivation. Stamens two. Thecae with different height. Pollen grains usually hexapseudocolpate tricolporate. – A possible topology of Justicieae is [Pseuderanthemum clade+[Isoglossinae+[Tetramerium clade+[[Rungia+Metarungia]+[Duvernoia+[Anisotes+[Old World ‘Justicia’ spp.+[Justicia betonica+[Diclipterinae+New World ‘justicioids’]]]]]]]]] (McDade & al. 2000). Pollen grains in Isoglossinae often “girdle pollen”, i.e. lenticular biporate pollen grains with prominent circumferential band.

Unplaced Acanthoideae

Dolichostachys (1; D. elongata; Madagascar), Gymnophragma (1; G. simplex; Papua New Guinea), Morsacanthus (1; M. nemoralis; Brazil), Perenideboles (1; P. ciliatum; Nicaragua), Sericospora (1; S. crinita; the West Indies), Sphacanthus (2; S. brillantaisia, S. humbertii; Madagascar; Justicieae-Isoglossinae?).

[Thunbergioideae+Avicennia]

Ovules two per carpel. Cotyledons folded. – The support of this clade is not particularly strong (Schwarzbach & McDade 2002; McDade & al. 2008).

Thunbergioideae Kostel., Allg. Med.-Pharm. Fl. 3: 923. Apr-Dec 1834

5/165–170. Thunbergia (c 100; tropical and subtropical regions in the Old World), Meyenia (1; M. hawtayneana; India, Sri Lanka)?, Pseudocalyx (5; P. aurantiacus, P. heterochondros, P. macrophyllus, P. pasquierorum, P. saccatus; tropical Africa, Madagascar), Mendoncia (60–65; tropical Africa, Madagascar, tropical America), Anomacanthus (1; A. congolanus; Congo, Angola). – Pantropical. Usually twining-climbing herbs (sometimes upright). Medulla with inverted vascular bundles, or secondary lateral growth anomalous (inner and inverted cambium developing). Leaves with strongly curved ptyxis. Petiole vascular bundle transection arcuate or annular. Inflorescence with two or more flowers in median plane of leaf axil. Adaxial flowers first mature. Bracts absent. Floral prophylls (bracteoles) large, connate, surrounding flower as epicalyx. Calyx as a rim, or as up to 16 minute lobes; in Thunbergia with extrafloral nectaries. Corolla often with contorted (sometimes imbricate quincuncial) aestivation. Anthers poricidal, with lignified unicellular hairs (bristles), sagittate, sometimes slightly displaced; connective prolonged; basal staminal appendages sometimes present. Endothecium absent. Pollen grains octocolpate or spiraperturate. Connective elongate. Adaxial carpel in Mendoncia and Anomacanthus reduced and sterile. Stigma broadly and shortly bilobate (in Mendoncia and Anomacanthus) to trumpet-shaped (in Thunbergia, Pseudocalyx and Meyenia), with wide and often unequally sized lobes, papillate, Wet type. Ovules two per carpel. Fruit a one- or two-seeded drupe (Mendoncia and Anomacanthus) or a usually four-seeded septifragal capsule, without retinacula (Thunbergia, Pseudocalyx and Meyenia). Cotyledons sometimes (e.g. in Mendoncia) twice folded. n = 9, 28. Iridoids absent in Mendoncia. –– Mendoncia is sister to the clade [Thunbergia+Pseudocalyx] (Borg & al. 2008).

Avicennioideae Miers in London J. Bot. 7: 58. 1848 [‘Avicennieae’]

1/8. Avicennia (8; A. balanophora, A. bicolor, A. germinans, A. integra, A. marina, A. officinalis, A. schaueriana, A. tonduzii; tropical, subtropical and warm-temperate estuarine areas). – Mangrove trees or mangrove shrubs with pneumatophore roots. Wood with anomalous secondary lateral growth from successive cambia. Intraxylary phloem as islands in bands of conjunctive tissue? Nodes 3:3, trilacunar with three leaf traces, swollen. Leaves opposite, entire, somewhat fleshy, with abaxial side covered with clavate hairs, and with salt glands on both sides. Petiole vascular bundle transection annular. Sclereids and colleters present. Flowers in dense thyrsoid spike-like units. Flowers more or less zygomorphic (sometimes actinomorphic). Sepals four (to six), persistent, free or almost free. Petals four (to six). Corolla tube with nectar-secreting glands. Disc small (nectaries as secretory hair tufts?). Stamens four (to six), alternipetalous. Secondary septa sometimes present. Ovary with apically fused locules. Stigma bilobate, with obtuse lobes. Placentation free central to apical (ovules inserted on septa). Ovules sometimes orthotropous? Micropyle absent. Megagametophyte extra-ovular. Endosperm development cellular?, asymmetrical. Micropylar endosperm haustorium extra-ovular, highly branched, reaching placenta. Fruit one-seeded fleshy capsule-like, dehiscing when seed germinates, with persistent green calyx. Seed fairly large. n = 18, 32. Tannins and betaines present. – Avicennia is sister to Thunbergioideae, according to DNA data, although morphological features identify Acanthoideae as its sister-group (Schwarzbach & McDade 2002; McDade & al. 2008).

BIGNONIACEAE Juss.

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de Jussieu, Gen. Plant.: 137. 4 Aug 1789 [’Bignoniae’], nom. cons.

Bignoniales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 246. Jan-Apr 1820 [‘Bignoniaceae’]; Bignoniopsida Nees in Flora 8: 142, 143. 7 Mar 1825 [’Bignoniaceae’]; Bignoniineae Link, Handbuch 1: 503. 4-11 Jul 1829 [‘Bignoniaceae‘]; Crescentiaceae Dumort., Anal. Fam. Plant.: 20, 24. 1829

Genera/species 77–78/870–885

Distribution Mainly tropical regions, with their largest diversity in tropical South America; some species in subtropical and warm-temperate Asia.

Fossils Winged seeds attributable to Bignoniaceae are reported from the Paleocene of North America and Japan.

Habit Bisexual, usually evergreen (rarely deciduous) trees, shrubs or lianas (Argylia, Incarvillea and Niedzwedzkia are perennial herbs, Tourrettia consists of twining perennial herbs). Often with leaf-tendrils. Lenticels often frequent on stems and branches.

Vegetative anatomy Phellogen ab initio superficial or cortical. Secondary lateral growth usually normal (in Bignonieae anomalous, via concentric cambia or cylindrical cambium; secondary xylem sometimes not formed). Cambium storied. Vessel elements with usually simple (rarely scalariform or reticulate) perforation plates; lateral pits alternate, usually bordered (rarely simple) pits. Vestured pits absent? Imperforate tracheary xylem elements often very long libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually homocellular (sometimes heterocellular). Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, unilateral or banded. Wood element often entirely or partially storied. Tyloses often abundant. Secondary phloem often stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids S type. Nodes 1:1 or 1:≥3, unilacunar with one, three or more leaf traces. Sclerenchymatous fibres present in some species. Silica bodies or prismatic calciumoxalate crystals (sometimes styloids or acicular crystals) present in some species.

Trichomes Hairs multicellular, uniseriate or branched, furcate, stellate, dendritic, lepidote; sometimes glandular hairs.

Leaves Usually opposite (sometimes verticillate, rarely alternate, spiral), usually pinnately compound (sometimes bipinnate or more, or palmately compound, or simple/unifoliolate); terminal leaflet in lianas usually modified into tendril), when simple then usually pinnately lobed (sometimes palmately lobed or entire), usually with conduplicate (in, e.g., Pyrostegia involute) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular; petiole sometimes also with edge bundles or adaxial bundles. Venation pinnate or palmate. Stomata usually anomocytic (sometimes paracytic, anisocytic, helicocytic, paracytic, cyclocytic, or diacytic). Cuticular wax crystalloids as clusters of terete rodlets. Domatia as pockets or hair tufts or absent. Leaf margin usually entire (sometimes serrate). Leaf apices, axillary buds and nodes often with extrafloral nectaries.

Inflorescence Terminal or axillary, dichasial, cincinnus or thyrse, raceme etc., or flowers solitary.

Flowers Zygomorphic (rarely almost actinomorphic), usually large. Hypogyny. Sepals five, with open or imbricate aestivation, connate into campanulate calyx (sometimes bilabiate or truncate; in some species of Lundia with calyptra), often with vascularized nectaries on abaxial side. Petals five, usually with imbricate quincuncial (rarely valvate) aestivation, connate into campanulate or infundibuliform corolla, quinquelobate or often bilabiate (upper lip bilobate, lower lip trilobate). Nectariferous disc intrastaminal, usually annular (sometimes cupular or absent).

Androecium Stamens usually two longer and two shorter (didynamous; in Catophractes, Oroxylum and ‘Rhigozum’ five), haplostemonous, antesepalous, alternipetalous; fifth stamen (adaxial, median) usually modified into staminodium or absent (in Catalpa and Paragonia two fertile stamens and three staminodia). Filaments free from each other, adnate to corolla tube (epipetalous). Anthers often head-to-head, usually connate, sometimes confluent, dorsifixed, often versatile, tetrasporangiate or disporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial. Staminodia usually one or three (rarely absent).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate or tricolporate (sometimes monocolpate, tetra- or pentacolpate, inaperturate, pericolpate, spiraperturate or syncolpate), usually shed as monads (sometimes as tetrads, rarely polyads), bicellular at dispersal. Exine tectate, semitectate or intectate, with columellate infratectum, reticulate, psilate, spinulate, or areolate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, usually bilocular (sometimes unilocular or almost quadrilocular due to incomplete secondary septa); often with nectaries on abaxial side. Style single, simple. Stigma capitate or broadly bilobate, sensitive, papillate, Wet type. Pistillodium absent.

Ovules Placentation axile (when ovary bilocular) or intrusively parietal (when ovary unilocular); placentae usually two per locule, sometimes lobed. Ovules usually numerous (rarely few) per carpel (two series of ovules per carpel), usually anatropous or hemianatropous (rarely orthotropous), ascending, unitegmic, tenuinucellar. Integument ? cell layers thick. Megasporangial endothelium present. Hypostase usually present. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes persistent. Endosperm development cellular. Endosperm haustoria chalazal or micropylar and chalazal. Embryogenesis usually ? (in Catalpa onagrad).

Fruit Usually a loculicidal or septicidal capsule (often very large and lignified, often with replum; in Crescentieae and Colea clade usually indehiscent), often with extrafloral nectaries.

Seeds Seeds flat. Aril absent. Testa usually with membranous or suberous wings (in species with capsular fruit); wing cells with usually annular or helical (sometimes reticulate) thickenings. Exotestal cells? Endotesta? Perisperm not developed. Endosperm usually absent. Embryo straight, oily, without chlorophyll. Cotyledons two, usually large, lobate, foliaceous, obcordate, persistent. Germination usually phanerocotylar (sometimes cryptocotylar).

Cytology n = 11, 13–15, (18–)20, (21) (40) – Polyploidy occurring.

DNA Deletion present in plastid gene matK. Mitochondrial coxI intron present in Catalpa.

Phytochemistry Flavonols (quercetin), 6- or 8-hydroxyflavones or 6-methoxyflavones, Route II decarboxylated iridoids, Group I carbocyclic iridoids (catalpol, macfadienoside), Group X secoiridoids (iridoid pyridine alkaloids), iridoid glucosides, iridoid aldehydes, ursolic acid and caffeic acid esters (cornosides in Eccremocarpus, verbascosides), saponins, shikimic acid derived arthroquinones, and naphthoquinones present. Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Carbohydrates stored as stachyose and other oligosaccharides.

Use Ornamental plants, medicinal plants, timber, ropes, calabashes and musical instruments (Crescentia), dyeing substances.

Systematics (according to Olmstead & al. 2009)

The sister-group relationship of Bignoniaceae is unresolved, although Refulio-Rodriguez & Olmstead (2014) recovered the following topology (with weak support): [Bignoniaceae+[[Schlegeliaceae+Lentibulariaceae]+[[Thomandersiaceae+Verbenaceae]+[Lamiaceae+[Mazaceae+[Phrymaceae+[Paulowniaceae+[Rehmanniaceae+Orobanchaceae]]]]]]]].

A probable topology of Bignoniaceae is the following: [Jacarandeae+[Tourrettieae+ [Tecomeae+[Delostoma+[Bignonieae+[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]]]]]]

Jacarandeae Seem. in Ann. Mag. Nat. Hist., ser. 3, 10: 31. Jul 1862

2/50–53. Jacaranda (c 50; Mexico, Central America, the West Indies, tropical South America), Digomphia (3; D. ceratophora, D. densicoma, D. laurifolia; the Guayana Highlands in southeastern Colombia, southern Venezuela, northwestern Brazil and Guyana). – Tropical and subtropical America. Evergreen trees or shrubs. Sepals free or almost free. Staminodium large, with beard-like indumentum. Placentation parietal. Fruit orbicular, angustiseptate. n = 18.

[Tourrettieae+[Tecomeae+[Delostoma+[Bignonieae+[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]]]]]

Tourrettieae G. Don, Gen. Hist. 4: 215, 231. 1837-8 Apr 1838 [‘Tourretieae’]

2/4. Eccremocarpus (3; E. longiflorus, E. scaber, E. vargasianus; Peru, Chile, western Argentina), Tourrettia (1; T. lappacea; Mexico to the Andes in tropical South America). – Mexico to Chile. Herbs twining with leaf-tendrils. Inflorescence a bracteate raceme. Staminodium absent in Tourrettia. Tourrettia with quadrilocular ovary having one row of ovules in each locule. Placentation in Eccremocarpus parietal. Cornosides present in Eccremocarpus.

[Tecomeae+[Delostoma+[Bignonieae+[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]]]]

Leaves once compound. Staminodia sometimes present, simple.

Tecomeae Endl., Gen. Plant.: 711. Jan 1839

11/71–72. Argylia (13; southern Peru, Chile, Argentina); Campsis (2; C. grandiflora: East Asia; C. radicans: southeastern United States), Tecoma (14; tropical and southern Africa, Arizona to northern Argentina, the West Indies), Incarvillea (c 16; Central Asia and the Himalayas to East Asia), Podranea (1; P. ricasoliana; tropical and southern Africa), Deplanchea (c 8; Malesia to New Guinea, northeastern Queensland and New Caledonia), Lamiodendron (1; L. magnificum; New Guinea), Tecomanthe (5–6; T. dendrophila, T. ternatensis: Malesia to New Guinea, New Britain, Solomon Islands; T. volubilis: New Guinea; T. hillii: eastern Queensland, northeastern New South Wales; T. speciosa: Three Kings Islands off New Zealand), ‘Pandorea’ (9; East Malesia to New Guinea, Australia, New Caledonia; polyphyletic), Campsidium (1; C. valdivianum; Chile, Argentina), Astianthus (1; A. viminalis; Mexico to Nicaragua). – Warm-temperate to tropical regions. Distinctive C-4 formyl iridoids present. – Argylia is sister-group to the remaining Tecomeae, with Campsis as successive sister to the rest. In Campsis and species of Tecoma the abaxial calyx nectaries have a cellular structure very similar to that in Thomandersia (Thomandersiaceae; Wortley & al. 2005).

[Delostoma+[Bignonieae+[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]]]

Delostoma clade

1/5. Delostoma (5; D. dentatum, D. gracile, D. hookeri, D. integrifolium, D. lobbii; the Andes).

[Bignonieae+[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]]

Bignonieae Dumort., Anal. Fam. Plant.: 23. 1829 [‘Bignoniaceae’]

20/415–420. Perianthomega (1; P. vellozoi; southeastern Brazil, southeastern Bolivia, northeastern Paraguay); Adenocalymma (84; southern Mexico, the Lesser Antilles, Central America to southeastern Brazil), Stizophyllum (c 20; southern Mexico, Central America, tropical South America); Manaosella (1; M. cordifolia; Venezuela, Brazil, Bolivia), Pleonotoma (16; northern Central America to southern Brazil), Dolichandra (8; southeastern United States, Mexico, the West Indies, Central America, tropical South America), Amphilophium (43; southern Mexico, the West Indies, Central America, tropical South America), Bignonia (28; southeastern United States, Mexico, Central America, tropical South America), Mansoa (12; southern Mexico, Central America, the West Indies, tropical South America), Anemopaegma (45; southern Mexico, Central America, tropical South America), Pyrostegia (1; P. venusta; Mexico, Central America, tropical South America); Callichlamys (1; C. latifolia; southern Mexico, Central America, tropical South America), Martinella (3; M. insignis, M. iquitoensis, M. obovata; southern Mexico, Central America, tropical South America), Pachyptera (5; P. aromatica, P. erythraea, P. incarnata, P. kerere, P. linearis; Central America, tropical South America), Tanaecium (16–18; southern Mexico, the West Indies, Central America, tropical South America), Cuspidaria (16–19; southern Mexico, Central America, tropical South America), Tynanthus (c 20; southern Mexico, Central America, tropical South America), Lundia (c 20; southern Mexico, Central America, tropical South America), Xylophragma (7; X. harleyi, X. heterocalyx, X. myrianthum, X. platyphyllum, X. pratense, X. seemannianum, X. unifoliolatum; southern Mexico, Central America, tropical South America), Fridericia (68; Mexico, Central America, tropical South America). – Southeastern United States to tropical South America. Lianas with leaf-tendrils. Secondary lateral growth anomalous, via concentric cambia or cylindrical cambium (xylem cylinder in principle quadrilobate, hypothetical basal condition), phloem discontinuous. Leaves usually ternate. Stamens in Paragonia two fertile stamens and three staminodia. Fruit usually a septifragal capsule (sometimes also loculicidal) with persistent septum and separate whip-like vascular strands of lignified tissue (vascular bundles opposite septum). – Perianthomega is sister to the remaining Bignonieae. It has biternate leaves, stout simple tendrils representing petioles and three small remnants of leaflets present at their apices. Adenocalymma is successive sister to the remainder.

[[Catalpeae+Oroxyleae]+[Crescentieae+Coleeae]]

[Catalpeae+Oroxyleae]

Catalpeae DC. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 300, Comm. 208. 25-31 Oct 1840

1–2/11. Catalpa (10; East Asia, southeastern United States, the Greater Antilles; incl. Chilopsis?), Chilopsis (1; C. linearis; southwestern United States; in Catalpa?). – East Asia, southeastern United States, the Greater Antilles. Leaves opposite or verticillate (Catalpa) or alternate (spiral; Chilopsis), simple. Stamens in Catalpa two fertile stamens and three staminodia.

Oroxyleae A. H. Gentry ex Reveal et L. G. Lohmann in Phytoneuron 2012-37: 218. 23 Apr 2012

4/6–8. Hieris (1; H. curtisii; Penang in Malaysia), Millingtonia (1; M. hortensis; India to Burma and Southeast Asia, Malesia), Nyctocalos (3–5; N. brunfelsiiflorum, N. cuspidatum, N. jucundum, N. pinnatum, N. thomsonii; northeastern India to Burma, Yunnan and Southeast Asia, West Malesia), Oroxylum (1; O. indicum; Sri Lanka, India and the Himalayas to southern China, Southeast Asia, Malesia to Sulawesi and Timor). – Tropical Asia. Leaves sometimes bicompound. Flowers sometimes actinomorphic. Stamens in Oroxylum five. Fruit a septicidal capsule.

[Crescentieae+Coleeae]

Fruit more or less indehiscent, sometimes berry-like.

Coleeae Bojer ex Reveal in Phytoneuron 2012-37: 217. 23 Apr 20122012

18/140–145. ‘Rhigozum’ (7; R. brevispinosum, R. madagascariense, R. obovatum, R. somalense, R. trichotomum, R. virgatum, R. zambesiacum; northeast tropical and southern Africa, Madagascar; paraphyletic), Spathodea (1; S. campanulata; tropical West and Central Africa), Catophractes (1; C. alexandri; subtropical and southern Africa); Radermachera (c 17; southern China, Southeast Asia, West Malesia), Tecomella (1; T. undulata; the Arabian Peninsula to western India); Kigelia (1; K. africana; tropical and southern Africa), Stereospermum (20–25; tropical Africa, Madagascar, tropical Asia), Newbouldia (1; N. laevis; tropical West and Central Africa), Fernandoa (c 15; tropical Africa, Madagascar, southern China, Southeast Asia to Sumatra), Heterophragma (2; H. quadriloculare: India; H. sulfureum: Southeast Asia), Dolichandrone (10; East Africa, India to northern Australia, New Caledonia, islands in the Pacific), Markhamia (5–6; tropical and southern Africa, tropical Asia), Dinklageodoxa (1; D. scandens; Liberia), Perichlaena (1; P. richardi; Madagascar), Phylloctenium (2; P. bernieri, P. decaryanum; Madagascar), Phyllarthron (c 15; Madagascar, the Comoros), Rhodocolea (c 10; Madagascar), Colea (c 30; Madagascar, the Comoros, Mauritius, the Seychelles). – Tropical Africa, Madagascar, Indian Ocean islands, northwestern and western India, Southeast Asia, Malesia and eastwards to northern Queensland and Southwest Pacific Islands. Leaves sometimes phyllodinous, articulated. Inflorescence cauliflorous. Stamens in Catophractes and ‘Rhigozum’ five. Fruit indehiscent or almost indehiscent. Testa unwinged?

Crescentieae G. Don, Gen. Hist. 4: 216, 232. 1837-8 Apr 1838

12/c 160. Sparattosperma (4; S. catingae, S. leucanthum, S. rosea, S. vernicosum; tropical South America); Cybistax (1; C. antisyphilitica; Amazonian Brazil, eastern Peru), Godmania (5; G. aesculifolia, G. dardanoi, G. luteola, G. macrocarpa, G. uleana; southern Mexico, Central America, tropical America), Zeyheria (2; Z. montana, Z. tuberculosa; Brazil), Ekmanianthe (2; E. actinophylla, E. longiflora; Cuba, Hispaniola), ‘Tabebuia’ (c 75; Mexico, Central America, the West Indies, tropical South America; polyphyletic), Roseodendron (2; R. chryseum: northern Colombia, northwestrn Venezuela; R. donnell-smithii: southern Mexico, Central America), Handroanthus (c 30; Central America, the West Indies, tropical South America), Parmentiera (9; southern Mexico, Central America, northwestern Colombia), Spirotecoma (5–6; S. apiculata, S. guantanamense, S. holguinensis, S. rubriflora, S. spiralis; Cuba, Hispaniola), Amphitecna (19; southern Mexico, Central America, tropical South America), ‘Crescentia’ (6; C. alata, C. amazonica, C. cujete, C. linearifolia, C. mirabilis, C. portoricensis; southern United States, Mexico, Central America, the West Indies, tropical South America; paraphyletic). – Tropical America including Cuba and Hispaniola. Leaves palmately compound (sometimes unifoliolate); in, e.g., Amphitecna and ‘Crescentia’ spiral, simple, phyllodinous. Inflorescence sometimes cauliflorous. Fruit sometimes (e.g. Amphitecna and ‘Crescentia’) indehiscent. Testa sometimes unwinged. – Sparattosperma may be sister to the remaining Crescentieae.

Parsimony strict consensus tree of Bignoniaceae based on DNA sequence data (Olmstead & al. 2009).

Unplaced Bignoniaceae

Neosepicaea (5; N. aurantiaca, N. jucunda, N. leptophylla, N. superba, N. viticoides; the Moluccas, New Guinea, eastern Queensland), Pajanelia (1; P. longifolia; tropical Asia), Paratecoma (1; P. peroba; coastal regions in Brazil), Pauldopia (1; P. ghorta; northeastern India to Southeast Asia), Romeroa (1; R. verticillata; Colombia), Santisukia (2; S. kerrii, S. pagetii; Thailand).

BYBLIDACEAE (Engl. et Gilg) K. Domin

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Domin in Acta Bot. Bohem. 1: 3. 1922, nom. cons.

Byblidales Nakai ex Reveal in Phytologia 74: 175. 25 Mar 1993

Genera/species 1/8

Distribution Northern and Western Australia, southern New Guinea.

Fossils A fossil seed of Byblis has been reported from mid-Eocene strata in the Golden Grove area in South Australia.

Habit Bisexual, evergreen suffrutices, perennial or annual herbs, often with woody rhizome. Probably not insectivorous (leaves apparently absorbing exudates/faeces from carnivorous mirids, cf. Roridula). Mycorrhiza absent.

Vegetative anatomy Mycorrhiza absent. Roots fibrous. Phellogen? Young stem with vascular tissue as separate bundles. Secondary lateral growth absent. Endodermis a starch sheath. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays usually biseriate or triseriate (sometimes uniseriate), homocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty. Sieve tube plastids Ss type. Nodes 1:1 or 1:3, unilacunar with one or three leaf traces.

Trichomes Glandular hairs stalked or sessile, non-vascularized; glands with mucilaginous head usually consisting of a layer of (eight to) 32 cells radiating like an umbrella outwards from centre of head.

Leaves Alternate (spiral), linear to filiform, with flat or circinate, abaxially curved, ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Veins parallelodromous (lamina reduced or absent?). Stomata paracytic. Cuticular wax crystalloids? Lamina covered with mucilage-secreting insect-trapping glandular hairs. Leaf margin entire, with vascularized apical hydathode. Foliar apex a knob-like swelling.

Inflorescence Flowers axillary, solitary. Bracts and floral prophylls (bracteoles) absent.

Flowers Slightly zygomorphic. Hypogyny. Sepals five, with imbricate aestivation, persistent, connate at base. Petals five, with contorted aestivation, connate at base, serrate to almost fimbriate at apex. Nectary absent. Disc absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous, displaced and often bent against one side of flower. Filaments short, subulate, free from each other, twisted, often slightly adnate to petals (epipetalous). Anthers connivent, basifixed, non-versatile, with cone-shaped apex and ephemeral epidermal cells, tetrasporangiate, introrse, poricidal (dehiscing by apical pores or short apical slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–4)-colpate (or tricolporate? or tetra- or hexarugate?), shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, punctitegillate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular. Style single, simple, long, filiform. Stigma twisted, usually punctate to capitate (sometimes slightly bilobate), type? Pistillodium absent.

Ovules Placentation apical-axile. Ovules two to numerous per carpel, anatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent. Endosperm development ab initio cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis onagrad.

Fruit A loculicidal capsule.

Seeds Aril absent. Exotestal cells tangentially elongate, with anticlinal walls not uniformly thickened. Mesotesta sclerenchymatous. Endotesta? Perisperm not developed. Endosperm copious, starchy and aleuroniferous. Embryo straight, elongate, chlorophyll? Cotyledons two, foliaceous. Germination phanerocotylar.

Cytology n = 8, (9) (12) 16 – Protein inclusions present in nucleus?

DNA Deletion in plastid gene matK?

Phytochemistry Insufficiently known. 6- and/or 8-hydroxylated flavone glycosides or 6-methoxy flavones present. Proanthocyanidins, cyanogenic compounds, and naphthoquinones not found. Iridoids? Carbohydrates stored as stachyose and other oligosaccharides.

Use Occasionally as ornamental plants.

Systematics Byblis (8; B. aquatica, B. filifolia, B. gigantea, B. guehoi, B. lamellata, B. liniflora, B. pilbarana, B. rorida; northern and western Australia, southern New Guinea).

The sister-group relationship of Byblis is unresolved. In Schäferhoff & al. (2010) it is part of trichotomy also including Linderniaceae and the remaining Plantaginales “above” Scrophulariaceae.

CALCEOLARIACEAE (D. Don) Olmstead

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Olmstead in Amer. J. Bot. 88: 357. 23 Feb 2001

Genera/species 2/390–395

Distribution Mexico to western South America, the Falkland Islands, New Zealand.

Fossils Unknown.

Habit Bisexual, perennial or annual herbs, suffrutices or shrubs.

Vegetative anatomy Phellogen? Vessel elements with simple? perforation plates; lateral pits? Vestured pits? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma? Sieve tube plastids S type? Nodes? Crystals?

Trichomes Hairs unicellular or multicellular; glandular hairs often present.

Leaves Opposite, simple or pinnately compound, entire or pinnately lobes, sometimes connate at base, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate or palmate. Stomata? Cuticular wax crystalloids? Leaf margin usually serrate or crenate (rarely entire).

Inflorescence Terminal, reduced thyrse with accessory paired flowers; thyrse consisting of one terminal flower and flower from its reduced prophyll (bracteole).

Flowers Zygomorphic. Usually hypogyny (sometimes half epigyny). Sepals four, with valvate aestivation, connate, orthogonally arranged/initiated. Petals four, with ascending cochlear aestivation? (adaxial corolla lobes of buds inserted outside the others), bilabiately connate, diagonally arranged/initiated; corolla bilabiate with large usually inflated saccate lower (abaxial) lip and small upper (adaxial) lip (fusion occurring late during development of each pair of petals; adaxial lip in Calceolaria triandra bipartite); inner side of lower (abaxial) corolla lip usually with elaiophores (oil glands attracting bees and bumblebees; absent in Jovellana) formed by hair cushions. Nectary usually absent (sometimes hair-like). Disc absent.

Androecium Stamens usually two, adaxial-lateral (in Calceolaria triandra three, i.e. two adaxial-lateral and one abaxial-median stamen). Filaments free from each other, adnate to corolla tube (epipetalous). Anthers basifixed, with thecae usually diverging, when dehiscing often confluent (sometimes parallel; sometimes with one theca), non-versatile, tetrasporangiate, dehiscing by apical pore-like slits. Placentoid? Tapetum secretory? Staminodia present or absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolpate?, shed as monads, bicellular at dispersal. Exine?, with ? infratectum, sculpturing?

Gynoecium Pistil composed of two connate carpels. Ovary usually superior (sometimes semi-inferior), bilocular. Style single, simple. Stigma small, capitate or slightly bilobate, type? Pistillodium absent.

Ovules Placentation axile. Ovules numerous per carpel, anatropous?, unitegmic, tenuinucellar. Integument three or four cell layers thick. Endothelium present; endothelial cells elongating (aulacospermous). Megagametophyte monosporous, Polygonum type. Endosperm development ab initio cellular. Endosperm haustoria? Embryogenesis onagrad.

Fruit A septicidal and loculicidal capsule.

Seeds Aril? Seed pedestals present. Testa with usually sinuate (sometimes straight) anticlinal cell walls. Exotesta? Endotesta? Perisperm not developed. Endosperm copious, prominent with longitudinal furrows. Embryo?, chlorophyll? Cotyledons two. Germination?

Cytology n = 8, 9, 15, 16 – Protein bodies in cell nuclei lamellar or absent.

DNA Mitochondrial coxI intron present. Deletion in plastid gene matK?

Phytochemistry Virtually unknown. Shikimic acid derived anthraquinones?

Use Ornamental plants.

Systematics Calceolaria (385–390; central Mexico, Central America, western South America southwards to Patagonia and the Falkland Islands, with their largest diversity in the Andes), Jovellana (8; J. albula, J. guentheri, J. petiolaris, J. punctata, J. sturmii, J. violacea: Chile; J. repens, J. sinclairii: New Zealand).

Calceolariaceae may be part of a clade [Calceolariaceae+Gesneriaceae+Peltanthera] or sister to Gesneriaceae. However, the support for any sister-group relationship among Calceolariaceae, Peltanthera and Gesneriaceae is not very high.

In spite of Calceolaria being one of the most frequently cultivated ornamental plants, the information on anatomy, pollen morphology, embryology, cytology, phytochemistry, etc. is extremely deficient.

CARLEMANNIACEAE Airy Shaw

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Airy Shaw in Kew Bull. 19: 511. 26 Jul 1965

Carlemanniales Doweld, Tent. Syst. Plant. Vasc.: xlviii. 23 Dec 2001

Genera/species 2/5

Distribution Assam, eastern Himalayas, southern China, Southeast Asia, Sumatra.

Fossils Unknown.

Habit Bisexual, shrubs or perennial herbs.

Vegetative anatomy Phellogen ab initio cortical (Carlemannia). Vessel elements with simple perforation plates; lateral pits alternate? Vestured pits? Imperforate tracheary xylem elements? Wood rays heterocellular? Axial parenchyma? Sieve tube plastids S type. Nodes ?-lacunar with ? leaf traces, swollen. Calciumoxalate raphides absent. Palisade parenchyma with stellate (calciumoxalate?) crystals.

Trichomes Glandular hairs peltate, with unicellular stalk and multicellular head; cells in head with exclusively vertical walls.

Leaves Opposite, simple, entire, sometimes distinctly asymmetrical, with ? ptyxis. Stipules and leaf sheath absent. Petiole bases joined by line. Petiole vascular bundles? Venation pinnate, brochidodromous. Stomata anomocytic (Carlemannia) or diacytic (Silvianthus). Cuticular waxes absent. Leaf margin serrate. Extrafloral nectaries present in leaf axils in Carlemannia.

Inflorescence Terminal and axillary, paniculate, corymboid or head-like, cymose.

Flowers Somewhat obliquely zygomorphic to almost actinomorphic. Epigyny. Sepals four or five, with open aestivation, often unequal in size, persistent, connate. Petals four or five, with imbricate or induplicate-valvate aestivation, connate into campanulate or infundibuliform corolla. Nectaries present on ovary apex. Disc cylindrical or conical. Heterostyly present in Silvianthus.

Androecium Stamens two, antesepalous, alternipetalous. Filaments short, free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, connivent around style, versatile?, tetrasporangiate, introrse (to latrorse?), longicidal (dehiscing by longitudinal slits). Placentoid? Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate (Silvianthus) or penta- or hexacolpate (Carlemannia), shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of two connate carpels. Ovary inferior, bilocular. Style single, simple, elongate. Stigma clavate to fusiform, bifid, type? Pistillodium absent.

Ovules Placentation axile to subbasal. Ovules numerous per carpel, anatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustorium? Embryogenesis?

Fruit A dry bifid loculicidal capsule (Carlemannia) or a quadrilobate or quinquelobate fleshy capsule (Silvianthus) with persistent calyx (capsule valves same number as calyx lobes).

Seeds Aril? Exotestal cells palisade?, narrow, polygonal, with all walls thickened (Carlemannia) or with thickened radial walls and inside these cells with unthickened walls (Silvianthus). Endotesta? Perisperm not developed. Endosperm copious, fleshy, oily, ruminate (Silvianthus). Embryo small, chlorophyll? Cotyledons two. Germination?

Cytology n = 15 (Carlemannia), n = 19 (Silvianthus) – Protein bodies present in nucleus?

DNA

Phytochemistry Virtually unknown. Iridoids? Alkaloids not found.

Use Unknown.

Systematics Carlemannia (3; C. congesta, C. griffithii, C. tetragona; Assam, eastern Himalayas, southwestern China, mountains in Southeast Asia to Sumatra), Silvianthus (2; S. bracteatus, S. tonkinensis; Assam, southern China, Southeast Asia).

Carlemanniaceae are sister-group to Oleaceae.

Since Carlemanniaceae are basal in Plantaginales, it is probable that Route II decarboxylated iridoids are absent.

GESNERIACEAE Rich. et Juss. ex DC.

( Back to Plantaginales )

de Candolle, Essai Propr. Méd. Pl., ed. 2: 192. 11 Mai 1816 [’Gessnerieae’], nom. cons.

Belloniaceae Martinov, Tekhno-Bot. Slovar: 67. 3 Aug 1820 [’Bellonides’]; Gesneriales Rich. ex Bercht. et J. Presl, Přir. Rostlin: 252. Jan-Apr 1820 [‘Gesneriae’]; Didymocarpaceae D. Don in Edinburgh Philos. J. 7: 83. 1822 [’Didymocarpeae’]; Cyrtandraceae Jack in Trans. Linn. Soc. London 14: 23. 1823; Gesneriineae Link, Handbuch 1: 505. 4-11 Jul 1829 [‘Gesneriaceae’]; Besleriaceae Raf., Sylva Tellur.: 70. Oct-Dec 1838 [’Beslerides’]; Cyrtandrineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 981. 1846 [‘Cyrtandraceae’]; Ramondaceae Godr. in J. C. M. Grenier et D. A. Godron, Fl. France 2: 506. 1853

Genera/species 148/3.220–3.440

Distribution Tropical and subtropical regions in the Northern and Southern Hemispheres; some species in temperate regions.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious), usually perennial herbs (sometimes lianas, rarely trees, shrubs or annual herbs). Many species are epiphytic. Stem and leaves often more or less succulent. Root fibrous.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Endodermis sometimes prominent. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, bordered pits. Vestured pits? Imperforate tracheary xylem elements usually libriform fibres (in Coronanthera fibre tracheids) with simple or (vestigially) bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric, or absent. Cambium sometimes storied. Wood elements (fibres) occasionally storied. Tyloses present. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace, or 3:3, trilacunar with three traces (rarely 4:4, quadrilacunar, or 5:5, quinquelacunar), sometimes with lateral vascular bundles split at nodes. Schizogenous secretory canals with oils or resins present in some Epithematoideae. Sclereids of various ouline abundant. Calciumoxalate frequent (sometimes as rhomboidal crystals or raphides). Cortical cells often with druses and crystals.

Trichomes Hairs usually simple, uniseriate (rarely multiseriate, rarely branched), unicellular or multicellular, often with thickened terminal cells (sometimes calcified/silicified); stalked multicellular (sometimes lepidote) glandular hairs sometimes frequent. Extrafloral nectaries rarely present on petiole.

Leaves Usually opposite (rarely verticillate, spiral or distichous; sometimes seemingly alternate due to anisophylly, especially in Epithematoideae), usually simple (rarely pinnately compound), usually entire (rarely lobed; rarely with one cotyledon developing into solitary large leaf), often coriaceous, with involute ptyxis. Stipules and leaf sheath absent. Leaves connate pairwise at base. Petiole vascular bundle transection arcuate; bundles variously arranged. Venation pinnate. Stomata usually anomocytic (often large; sometimes anisocytic, helicocytic or paracytic). Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts. Secretory cavities with oils or resins present in some epithematoid genera. Leaf margin serrate, crenate, lobed or entire.

Inflorescence Usually axillary (sometimes terminal), thyrsoid; flowers often paired, or solitary axillary (in Chirita, Didymocarpus, and Streptocarpus sometimes epiphyllous). Accessory flowers often present.

Flowers Usually zygomorphic (rarely inverted 180o; rarely actinomorphic). Hypogyny, half epigyny or epigyny. Sepals (four or) five, usually with valvate (rarely imbricate) aestivation, usually connate into tube (calyx sometimes bilabiate or trilabiate; sepals sometimes free). Petals (four or) five, with usually descending cochlear aestivation, adaxial petals usually posterior, connate into infundibuliform, tubular, campanulate or hypocrateriform and usually bilabiate (two upper and three lower lobes) corolla (sometimes quinquelobate), sometimes with spur. Nectariferous disc intrastaminal, annular or cupular, sometimes unilateral or consisting of separate glands (disc sometimes absent); nectary vascularized from androecial trace.

Androecium Stamens usually two longer and two shorter (didynamous), fifth (adaxial) stamen staminodial or absent (sometimes two fertile stamens and three staminodia; rarely all five stamens fertile), haplostemonous, antesepalous, alternipetalous. Filaments usually free from each other, adnate to corolla tube (epipetalous). Anthers usually basifixed (rarely dorsifixed), usually connivent (sometimes connate, rarely separate) pairwise or all together, versatile?, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical, rarely basal, pores). Tapetum secretory, with binucleate to quinquenucleate cells. Staminodia one to three or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colpor(oid)ate or (2–)3(–6)-colpate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, usually perforate or reticulate, verrucate, scabrate, psilate or smooth.

Gynoecium Pistil composed of two connate carpels. Ovary superior, inferior or semi-inferior, usually unilocular (sometimes secondarily bilocular by ingrowth of placentae; rarely primarily bilocular or with one carpel sterile, pseudomonomerous). Style single, simple, narrow. Stigma capitate or broadly bifid to trumpet-shaped, papillate, Dry or Wet type. Pistillodium?

Ovules Placentation usually intrusively parietal (rarely axile and ovary thus primarily bilocular). Ovules numerous per carpel, usually anatropous (rarely orthotropous), unitegmic, tenuinucellar (reduced, with meiocyte semi-inferior). Integument three to five cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids sometimes with long narrowing tips, entirely penetrating micropyle. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis onagrad.

Fruit Usually a loculicidal and/or septicidal capsule (sometimes fleshy, rarely irregularly dehiscing or a pyxidium; sometimes berry- or nut-like).

Seeds Aril present or absent. Exotestal cells elongate, with thickened walls. Endotestal cells degenerating or persistent. Perisperm not developed. Endosperm copious (multi-layered in neotropical Gesneriaceae) or sparse (uni-layered in paleotropical Gesneriaceae), oily, or absent. Embryo straight, without chlorophyll. Cotyledons two, often with non-uniform growth (in some species a single cotyledon, macrocotyledon, continuing its growth and being single leaf of plant); radicula sometimes with limited growth. Germination phanerocotylar.

Cytology n = 4, 7–18(–21, 24, 28, 30, 32, 36, 48, 64) – Polyploidy occurring. Protein bodies in cell nucleus lamellar?

DNA Deletion in plastid gene matK. Nuclear gene GCyc duplicated. Mitochondrial coxI intron present in Drymonia and Nematanthus.

Phytochemistry Flavones, 6- and/or 8-hydroxylated flavone glycosides, aurones and chalcones (paleotropical Gesneriaceae), 3-desoxyanthocyanins (neotropical Gesneriaceae), tannins, caffeic acid, caffeic acid esters (cornosides, verbascosides), naphthoquinones, and shikimic acid derived anthraquinones present. Flavonols, iridoids, ellagic acid, proanthocyanidins, alkaloids, saponins, and cyanogenic compounds not found. Carbohydrates stored as stachyose and other oligosaccharides.

Use Ornamental plants, medicinal plants.

Systematics The clade [Calceolariaceae+Gesneriaceae] is sister to the remaining Plantaginales “beyond” Tetrachondraceae. – For information on Peltanthera, see Plantaginales incertae sedis.

Sanangoideae A. Weber, J. L. Clark et Mich. Möller in Selbyana 31(2): 83. Dec 2013

1/1. Sanango (1; S. racemosum; eastern Peru) was sister to the remaining Gesneriaceae, according to Perret & al. (2013).

Analyses of DNA sequences identify Sanango racemosum outside (alternatively at the base of) Gesneriaceae (e.g. Perret & al. 2013). In ndhF analyses by Smith, Brown & al. (1997) it is recovered as sister to Gesneria. It is a small tree with opposite leaves. The hypogynous flowers are, often pairwise, arranged in terminal thyrses. The sepals are connate at base, and the petals have cochlear aestivation and are connate into a curved tube. The nectary is cup-shaped. The stamens are four, with filaments adnate to base of corolla tube. The anthers are versatile, introrse and dehisce along a hippocrepomorphic line. A single small staminodium is present. The ovary is incompletely septate in upper part and completely septate in lower part. The stigma is capitate, bilobate, and with lobes laterally adnate to style and directed downwards. The placentation is axile. The fruit is a capsule, at first septicidal, later loculicidal, with long persistent style. n = 16. Sananga has sanangoside, a caffeoyl phenylethanoid glycoside (also suggesting at least a position within Plantaginales), whereas iridoids have not been found.

Gesnerioideae Burnett, Outlines Bot.: 959, 1095, 1108. Feb 1835 [’Gesneridae’]

77/1.380–1.400. Mainly neotropical, some genera in East Asia and southwestern Pacific. Testa usually without surface ornamentation. Testal cells usually much elongated, usually spirally arranged. Endosperm copious. Nuclear gene GCyc2 absent (lost). 3-desoxyanthocyanins present. Chalcones and aurones not found. – Napeanthus, Titanotrichum, Besleria, Cremosperma, Anetanthus, Gasteranthus, and Reldia were identified as basal to the remaining Gesnerioideae in analyses by, e.g., Zimmer & al. (2002) and Perret & al. (2013).

Titanotricheae Yamaz. ex W. T. Wang, Fl. Reipubl. Popularis sin. 69: 577. 1990

1/1. Titanotrichum (1; T. oldhamii; southeastern China, southern Japan, the Ryukyu Islands, Taiwan). Stomata sometimes anomocytic. Vegetative propagules (‘bulbils’) formed after flowering from elongated inflorescence apex. Testa with striate-reticulate surface, and isocotylous cotyledons. n = 20. – Titanotrichum may be sister to Napeanthus (Perret & al. 2013).

Napeantheae Wiehler in Selbyana 6: 151. 31 Aug 1983

1/>20. Napeanthus (>20; Central America, tropical South America). Flowers almost actinomorphic, without nectaries. n = 16.

Beslerieae Bartl., Ord. Nat. Pl.: 175. Sep 1830 [’Besleriea’]

9/>275. Besleriinae G. Don 1837-1838 [‘Beslerieae’]. Besleria (>200; tropical and subtropical America, with their largest diversity in the northern Andes), Gasteranthus (c 35; southern Mexico, Central America, tropical South America), Reldia (5; R. alternifolia, R. calcarata, R. grandiflora, R. minutiflora, R. multiflora; Panamá to northern Peru), Cremosperma (25–30; Panamá to the Andes in Peru). – Anetanthinae A. Weber et J. L. Clark in Selbyana 31(2). Dec 2013. Anetanthus (3; A. gracilis, A. parviflorus, A. rubra; central Colombia, southeastern Brazil, Peru, Bolivia), Resia (3; R. ichthyoides, R. nimbicola, R. umbratica; Colombia, Venezuela), Shuaria (1; S. ecuadorica; Ecuador); Cremospermopsis (2; C. cestroides, C. parviflora; Colombia), Tylopsacas (1; T. cuneata; the Guayana Highlands). – Besleria has n = 16.

Coronanthereae Fritsch in Engler et Prantl, Nat. Pflanzenfam. IV, 3b: 143. Mai 1893

9/21–29. Coronantherinae Fritsch in Engler et Prantl, Nat. Pflanzenfam. IV, 3b: 143. May 1893. Coronanthera (13–20; New Caledonia, Solomon Islands, eastern Queensland; paraphyletic?), Rhabdothamnus (1; R. solandri; North Island in New Zealand). – Mitrariinae Hanst. in Linnaea 26: 198, 199. 1854 [‘Beslerieae subtr. Mitrarieae’]. Asteranthera (1; A. ovata; southern Chile, southwestern Argentina), Mitraria (1; M. coccinea; southern Chile, southwestern Argentina), Sarmienta (1; S. scandens; southern Chile), Fieldia (1; F. australis; southeastern Queensland, eastern New South Wales, eastern Victoria). – Negriinae V. L. Woo, J. F. Smith et Garn.-Jones in Int. J. Pl. Sci. 172(3): 454. 2011. Depanthus (1–2; D. glaber, D. pubescens; New Caledonia), Lenbrassia (1; L. australiana; northeastern Queensland), Negria (1; N. rhabdothamnoides; Lord Howe). – Eastern Australia, Solomon Islands, New Caledonia, Lord Howe, New Zealand, southern South America. Trees, shrubs or perennial herbs. Stomata usually anomocytic (sometimes paracytic). Flowers usually zygomorphic (occasionally actinomorphic). Corolla sometimes fringed. Nectaries embedded in ovary wall, vascularized from staminal traces. Stamens sometimes two (adaxial pair) or five. Capsule usually septicidal (sometimes loculicidal-septicidal or fruit a berry with fleshy placentae). n = 37(–45). Nuclear gene GCyc duplicated. – The Australian-southern South American clade.

Gesnerieae Dumort., Anal. Fam. Plant.: 30. 1829 [‘Gesnereae’]

57/1.065–>1.075. Gesneriinae Oerst., Centralamer. Gesner.: 10. 1858 [‘Gesnereae’]. Bellonia (2; B. aspera: Hispaniola; B. spinosa: Cuba), Gesneria (45–50; the West Indies), Pheidonocarpa (1; P. corymbosa; Cuba, Jamaica; in Gesneria?), Rhytidophyllum (c 20; the West Indies; in Gesneria?). – Gloxiniinae G. Don, Gen. Hist. 4: 644, 645. 1837– 8 Apr 1838 [‘Gloxinieae’]. Chautemsia (1; C. calcicola; Minas Gerais in Brazil), Gloxinia (3; G. erinoides, G. perennis, G. xanthophylla; Costa Rica, tropical South America), Seemannia (4; S. gymnostoma, S. nematanthodes, S. purpurascens, S. sylvatica; the Andes from Ecuador to northern Argentina), Gloxinella (1; G. lindeniana; Cajamarca in Peru), Monopyle (17; Guatemala to Bolivia), Diastema (>20; Mexico, Central America, the Andes in Venezuela to Bolivia), Nomopyle (2; N. dodsonii, N. peruviana; Ecuador, Peru), Kohleria (17; Mexico, Central America, Trinidad, tropical South America to Peru and the Guianas, with their largest diversity in Colombia), Moussonia (11; southern Mexico to Panamá; in Kohleria?), Capanea (2; C. humboldtii, C. oerstedii; tropical America; in Kohleria?), Gloxiniopsis (1; G. racemosa; the Andes in Colombia), Amalophyllon (c 15; southern Mexico, Central America, the Andes in Venezuela to Peru), Pearcea (17; the Andes from northern Colombia to northwestern Bolivia), Eucodonia (2; E. andrieuxii, E. verticillata; central and southern Mexico), Smithiantha (8; Mexico, Guatemala), Niphaea (2; N. mexicana, N. oblonga; southern Mexico, Guatemala), Solenophora (16; Central America), Phinaea (3; P. albolineata, P. multiflora, P. pulchella; Mexico, Central America, Cuba, Hispaniola, northwestern South America to Venezuela and Peru), Heppiella (4; H. repens, H. ulmifolia, H. verticillata, H. viscida; the Andes in western Venezuela and Peru), Mandirola (3; M. ichthyostoma, M. multiflora, M. rupestris; Brazil), Goyazia (2; G. petraea, G. rupicola; central Brazil), Achimenes (24; southern Mexico, Central America, the West Indies, Colombia). – Columneinae Hanst. in Linnaea 26: 198, 199. Apr 1854 [‘Columneae’]. The Guayana Shield clade: Pagothyra (1; P. maculata; Venezuela, Guyana), Rhoogeton (1–4; R. viviparus; Guyana), Lembocarpus (1; L. amoenus; Suriname, French Guiana), Cremersia (1; C. platula; French Guiana); ‘Episcia’ (9; Nicaragua to tropical South America; paraphyletic), Alsobia (2; A. dianthiflora, A. punctata; southern Mexico, Belize, Guatemala, Costa Rica; in Episcia?), Cobananthus (1; C. calochlamys; Central America), ‘Nematanthus’ (c 30; southern and southeastern Brazil; paraphyletic; incl. Codonanthe?), Codonanthe (>25; tropical America, with their highest diversity in Brazil; in Nematanthus?), Crantzia (2; C. cristata, C. tigrina; the West Indies), Drymonia (>140; tropical America, with their highest diversity in Colombia and Ecuador; paraphyletic; incl. Neomortonia?), Neomortonia (2; N. nummularia, N. rosea; Central America, western Colombia; in Drymonia?), Alloplectus (10; Guatemala, Costa Rica, Panamá, the West Indies, the Andes in Colombia, northern Ecuador and Peru), Glossoloma (>20; southern Mexico, Central America, tropical South America to Bolivia), Columnea (>250; Central America, the West Indies, northern tropical South America), Paradrymonia (c 10; Central America, northern tropical South America), Centrosolenia (15; the Guayana Highlands), Chrysothemis (9; southern Mexico, Central America, the Lesser Antilles, northern tropical South America), ‘Nautilocalyx’ (>30; tropical America; paraphyletic), Trichodrymonia (c 50; southern Mexico, Central America, the Andes, tropical South America); unplaced Columneinae: Christopheria (1; C. xantha; Guyana, French Guiana), Corytoplectus (15; Panamá to coastal Venezuela and Bolivia, the Guayana Highlands), Lampadaria (1; L. rupestris; Guyana), Lesia (1; L. savannarum; Colombia to Suriname and Peru), Oerstedina (2; O. cerricola, O. suffrutescens; Central America), Pachycaulos (1; P. nummularium; southern Mexico to Peru), Rufodorsia (4; R. congestiflora, R. intermedia, R. major, R. minor; Costa Rica, Panamá). – Sphaerorrhizinae A. Weber et J. L. Clark in Selbyana 31(2): 85. Dec 2013. Sphaerorrhiza (2; S. burchellii, S. sarmentiana; Brazil). – Ligeriinae Hanst. in Linnaea 26: 198, 199. Apr 1854 [‘Ligerieae’]. Sinningia (c 75; Central America, tropical South America, with their highest diversity in eastern and southern Brazil), Vanhouttea (8; southeastern Brazil; in Sinningia?), Paliavana (5; P. gracilis, P. plumerioides, P. prasinata, P. racemosa, P. schiffneri; eastern and southeastern Brazil). – Southern Mexico, Central America, the West Indies, South America. Lateral vascular bundles in Episcieae dividing at nodes. Nodes sometimes 3:3, trilacunar with three leaf traces. Calciumoxalate styloids and raphides sometimes present. Leaves rarely spiral (Gesneria etc.). Petiole vascular bundle transection sometimes deeply arcuate to annular; wing bundles sometimes present. Stomata sometimes present on raised mounds. Flowers rarely resupinate. Hypogyny or epigyny. Nectaries vascularized from large number of vascular bundles in ovary wall. Fruit sometimes with fleshy placentae or funicles; rarely a berry. n = (8) 9 (10) 11 (12) 13–14 (16). – The Neotropical clade. – As indicated above, Gesnerieae may be paraphyletic.

[Didymocarpoideae+Epithemateae]

Nectaries vascularized from staminal traces. Ovary wall not very vascularized. Endosperm inconspicuous. Cotyledons unequal (one of them accrescent). Chalcones and aurones present. 3-desoxyanthocyanins not found. – The Paleotropical clade.

Didymocarpoideae Arn., Botany: 121. 9 Mar 1832 [‘Didymocarpeae’] (Trichosporeae Nees in Flora 8: 143. 7 Mar 1825)

62/1.760–1.980. Jerdoniinae A. Weber et Mich. Möller in Selbyana 31(2): 86. Dec 2013. Jerdonia (1; J. indica; southwestern India). – Corallodiscinae A. Weber et Mich. Möller in Selbyana 31(2): 87. Dec 2013. Corallodiscus (6; C. bhutanicus, C. cooperi, C. forrestii, C. grandis, C. kingianus, C. lanuginosus; eastern Himalayas, northern and northeastern India, southern and southwestern China, northern Thailand). – Tetraphyllinae A. Weber et Mich. Möller in Selbyana 31(2): 87. Dec 2013. Tetraphyllum (>3; T. bengalense, T. confertiflorum, T. roseum; northeastern India, Bangladesh, Burma, Thailand). – Leptoboeinae C. B. Clarke in J. D. Hooker, Fl. Brit. India 4: 337. Jan 1884 [‘Leptoboeae’]. 5(6)/42(43). Boeica (c 12; eastern Himalayas to southern China, Burma, northern Vietnam and the Malay Peninsula), Rhynchotechum (13–15; southern Himalayas to southern China, Southeast Asia, Malesia to New Guinea), Leptobaea (2; L. glabra, L. multiflora; eastern Himalayas to Yunnan, Burma and Thailand), Beccarinda (c 8; Assam, Burma, southern China, Hainan, Vietnam, Sumatra), Platystemma (1; P. violoides; the Himalayas, southwestern China), Championia (1; C. reticulata; Sri Lanka)? – Ramondinae DC. ex Meisn., Pl. Vasc. Gen.: Tab. Diagn. 302, Comm. 212. 25–31 Oct 1840 [‘Ramondieae’]. Haberlea (1; H. rhodopensis; the Rhodopi Mountains on southern Balkan Peninsula), Ramonda (3; R. myconi: the Pyrenees, northeastern Spain; R. nathaliae, R. serbica: southern Balkan Peninsula; incl. Jancaea?), Jancaea (1; J. heldreichii; Mount Olimbos in central Greece; in Ramonda?). – Litostigminae A. Weber et Mich. Möller in Selbyana 31(2): 87. Dec 2013. Litostigma (2; L. coriaceifolium, L. crystallinum; southwestern China). – Streptocarpinae Ivanina in Bot. Žurn. (Moscow et Leningrad) 50: 33. 1 Feb 1965. Streptocarpus (c 150; tropical and southern Africa, Madagascar; incl. Acanthonema, Colpogyne, Hovanella, Linnaeopsis, Nodonema, Saintpaulia, Schizoboea and Trachystigma?). – Didissandrinae A. Weber et Mich. Möller in Selbyana 31(2): 87. Dec 2013. Didissandra (8; West Malesia), Tribounia (2; T. grandiflora, T. venosa; Thailand). – Loxocarpinae A. DC. in A. P. et A. L. P. P. de Candolle, Prodr. 9: 277. 1 Jan 1845 [‘Loxocarpeae’]. Boea (17; northeastern India, southern China, northern Thailand, Vietnam, Central and East Malesia to New Guinea, northeastern Queensland, the Bismarck Archipelago, Solomon Islands), ‘Streptocarpus’ pro parte (Asia), Rhabdothamnopsis (1; R. sinensis; southern China), Senyumia (1; S. minutiflora; Pahang on the Malay Peninsula), Emarhendia (1; E. bettiana; Pahang on the Malay Peninsula), Spelaeanthus (1; S. chinii; Pahang on the Malay Peninsula, Batu Luas), Paraboea (95–100; Burma, southern China, Thailand to New Guinea), Middletonia (4; M. evrardii, M. monticola, M. multiflora, M. regularis; India, Bhutan and Burma to southern China and Southeast Asia), Kaisupeea (3; K. cyanea, K. herbacea, K. orthocarpa; Burma, Thailand, southern Laos), Loxocarpus (20–25; peninsular Thailand, the Malay Peninsula, Sumatra, Borneo), Damrongia (c 10; southern Burma, Thailand, the Malay Peninsula, Sumatra), Somrania (3; S. albiflora, S. flavida, S. lineata; southern Thailand), Ornithoboea (11; southern China, eastern Burma, Thailand, Vietnam, northern Malay Peninsula), Orchadocarpa (1; O. lilacina; the Malay Peninsula). – Didymocarpinae G. Don, Gen. Hist. 4: 644, 658. 1837–8 Apr 1838 [‘Didymocarpeae’]. Codonoboea (80–85; Southeast Asia), Microchirita (c 35; India, Burma, southern China, Thailand, Indochina, West Malesia), Henckelia (c 55; southern India, Sri Lanka, southern and eastern China, peninsular Thailand, Laos, the Malay Peninsula, Malesia to New Guinea), Didymostigma (3; D. leiophyllum, D. obtusum, D. trichanthera; southeastern China including Hainan), Billolivia (9; Indochina), Conandron (1; C. ramondioides; eastern China, southern Japan, Taiwan), Ridleyandra (>20; the Malay Peninsula, Borneo), Hexatheca (3–4; H. dolichopoda, H. fulva, H. johannis-winkleri; Borneo), Cyrtandra (450–600; the Nicobar Islands and southern Thailand to Malesia, islands in southern Pacific to the Hawaiian Islands; paraphyletic?), Sepikea (1; S. cylindrocarpa; the Sepik area on New Guinea), Metapetrocosmea (1; M. peltata; Hainan), Aeschynanthus (c 185; northern and southern India, southern China, Malesia to New Guinea and Solomon Islands; paraphyletic?), Agalmyla (>95; Malesia to New Guinea), Allocheilos (2; A. cortusiflorus, A. guangxiensis; southern China, Vietnam), Liebigia (12; Sumatra, Java, Bali), Didymocarpus (>75; northern India, Nepal and southern China to the Malay Peninsula and northern Sumatra), Oreocharis (100–140; the Himalayas, central and southern China, Burma, Thailand, Vietnam), Cathayanthe (1; C. biflora; Hainan), Petrocodon (19; southern China, northeastern Thailand, northern Vietnam), Primulina (115–120; southern China), Deinostigma (1–7; D. poilanei; southern China, Vietnam), Loxostigma (>7; L. brevipetiolatum, L. cavaleriei, L. fimbrisepalum, L. glabrifolium, L. griffithii, L. mekongense, L. musetorum; northeastern India, the Himalayas, Burma, southern China, Indochina), Pseudochirita (1; P. guangxiensis; southern China, Vietnam), Allostigma (1; A. guangxiense; Guangxi in southern China), Petrocosmea (c 27; northeastern India, Burma, southern China, Southeast Asia), Chayamaritia (2; C. banksiae, C. smitinandii; Thailand, Laos), Briggsiopsis (1; B. delavayi; southern China), Glabrella (1; G. leiophylla; southern China), Hemiboea (c 27; central and southern China, the Ryukyu Islands, Taiwan, northern Vietnam), Anna (3; A. mollifolia, A. ophiorrhizoides, A. submontana; China, northern Vietnam), Lysionotus (c 30; eastern Himalayas, southern China, northern Thailand, northern Vietnam, southern Japan), ‘Raphiocarpus’ (c 15; southern China, Vietnam; polyphyletic). – Southern Europe, India, Sri Lanka and southern China to Japan, Taiwan, Malesia and tropical Australia, Solomon Islands and islands in the Pacific to the Hawaiian Islands, few species in tropical Africa and Madagascar. Nodes sometimes 1:1 or 3:3 with split laterals, or 5:5. Stamens sometimes two (abaxial pair). Ovary usually gradually narrowing into style. Placentae lamelliform-recurved. Ovules restricted to distal end. Fruit elongated, twisted, sometimes both loculicidal and septicidal (sometimes a pyxidium or a berry). Testal cells little elongated (sometimes ornamented, with extremely long hairs). n = (4, 8) 9–11 (12, 13) 14–17 or more. Polyploidy often occurring. Dihydrocaffeoyl ester present in at least Haberlea and Ramonda. – Jerdonia may be sister to the remaining Didymocarpoideae. Jerdonia has flattened filaments, pollen grains shed in tetrads, four separate parietal placentae, alveolate endosperm, isocotylous seedlings, and n = 14.

Epithemateae C. B. Clarke, Commelyn. et cyrtandr. Bengal. 67. 1874 [‘Epithemeae’]

7/>80. Loxotidinae G. Don, Gen. Hist. 4: 645, 664. 1837–8 Apr 1838 [’Loxotieae’]. Rhynchoglossum (c 10; India and southern China to Southeast Asia, Malesia to New Guinea, one to three species, R. azureum, in southern Mexico, Central America, Colombia and Venezuela). – Monophyllaeinae A. Weber et Mich. Möller in Selbyana 31(2): 86. Dec 2013. Whytockia (8; southern China, Taiwan), Monophyllaea (>30; peninsular Thailand, Malesia to New Guinea). – Loxoniinae A. DC. in A. P. et A. L. P. P. de Candolle, Prodr. 9: 274. 1 Jan 1845 [‘Loxonieae’]. Stauranthera (3; S. coerulea, S. grandifolia, S. umbrosa; northeast India and southern China to Southeast Asia, Malesia to New Guinea), Loxonia (3; L. burttiana, L. discolor, L. hirsuta; the Malay Peninsula, Sumatra, Java, Borneo), Gyrogyne (1; G. subaequifolia; Guangxi in southern China; possibly extinct). – Epithematinae DC. ex Meisn. Plant. Vasc. Gen.: Tab. Diagn. 303, Comm. 212. 25-31 Oct 1840 [‘Epithemeae’] 1/>20. Epithema (22; northeastern India and Nepal to southern China and Southeast Asia, Taiwan, Malesia to New Guinea, one species, E. tenue, in tropical West and Central Africa). – India, Southeast Asia, Malesia, one species in tropical West Africa, one species in tropical America. Secretory canals present. Medullary bundles present in Rhynchoglossum. Inflorescence (cymes) without floral prophylls (bracteoles). Abaxial corolla lobe sometimes inside others in bud. Nectaries sometimes vascularized. Stamens sometimes two (adaxial pair). Ovary short, abruptly narrowed into style. Placentation sometimes axile. Endosperm absent? n = (8–)10(–12). Dihydroxyphenols (i.a. acteoside) not found.

Unplaced Gesneriaceae

Hygea (1; H. barbigera; Chile).

Cladogram of Gesnerieae based on DNA sequence data (Zimmer & al. 2002).

Phylogeny of Gesneriaceae, according to Weber & al. (2013).

LAMIACEAE Martinov

( Back to Plantaginales )

Martinov, Tekhno-Bot. Slovar: 355. 3 Aug 1820 [’Lamieae’], nom. cons.

Labiatae Juss., Gen. Plant.: 110. 4 Aug 1789, nom. cons. et nom. alt.; Viticaceae Juss., Gen. Plant.: 106. 4 Aug 1789 [’Vitices’]; Glechomaceae Martinov, Tekhno-Bot. Slovar: 288. 3 Aug 1820 [‘Glecomeae’]; Melissaceae Bercht. et J. Presl, Přir. Rostlin: 245. Jan-Apr 1820 [’Melisseae’]; Melittidaceae Martinov, Tekhno-Bot. Slovar: 390. 3 Aug 1820 [‘Meliteae, Melytteae’]; Nepetaceae Bercht. et J. Presl, Přir. Rostlin: 245. Jan-Apr 1820 [‘Nepeteae’]; Salviaceae Bercht. et J. Presl, Přir. Rostlin: 245. Jan-Apr 1820; Viticales Link, Hort. Berol. 1: 446. 4-11 Jul 1829 [‘Viticeae’]; Menthaceae Burnett, Outl. Bot.: 969, 1095, 1106. Feb 1835; Menthales Burnett, Outl. Bot.: 1106. Jun 1835 [‘Menthinae’], nom. illeg.; Aegiphilaceae Raf., Sylva Tellur.: 161. Oct-Dec 1838 [‘Aegiphilia’]; Siphonanthaceae Raf., Fl. Tellur. 4: 87. med 1838 [‘Siphonanthia’]; Ajugaceae Döll, Rhein. Fl.: 375. 24-27 Mai 1843 [‘Ajugoideae’]; Scutellariaceae Döll, Rhein. Fl.: 373. 24-27 Mai 1843 [‘Scutellarineae’]; Stachydaceae Döll, Rhein. Fl.: 366. 24-27 Mai 1843 [’Stachydeae’]; Ajugineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1162, 1195. 1846; Menthineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1161, 1164. 1846; Nepetineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1162, 1179. 1846; Scutellariineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1162, 1193. 1846 [‘Scutellarineae’]; Stachydineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1162, 1193. 1846 [‘Stachydeae’]; Symphoremataceae Wight, Icon. Plant. Ind. Orient. 4(3): 13. Apr 1849 [’Symphoremeae’]; Dicrastylidaceae J. Drumm. ex Harv. in Hooker’s J. Bot. Kew Gard. Misc. 7: 56. 1855 [‘Dicrastyleae’], nom. nud.; Monardaceae Döll, Fl. Baden 2: 661. med 1858 [’Monardeae’]; Saturejaceae Döll, Fl. Baden 2: 664. med 1858 [‘Satureineae’]; Lamiineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 464. 15 Nov 1895 [‘Lamiales’]; Chloanthaceae Hutch., Fam. Fl. Pl., ed. 2: 396. 4 Jun 1959; Salazariaceae F. A. Barkley in Phytologia 32: 304. 29 Oct 1975

Genera/species 215/6.860–7.450

Distribution Cosmopolitan except polar areas.

Fossils Various fossil leaves, flowers and fruits have been reported from the Eocene onwards of North America, India and Europe. Pollen grains of Lamiaceae have been found in Miocene layers of Central Europe and China.

Habit Usually bisexual (sometimes gynomonoecious or gynodioecious, rarely polygamomonoecious or dioecious), usually perennial, biennial or annual herbs (sometimes evergreen trees or shrubs, rarely lianas). Many species are xerophytic. Young stems and branches usually quadrangular in cross-section. Usually aromatic.

Vegetative anatomy Roots usually fibrous. Phellogen ab initio superficial or deeply seated. Primary medullary rays narrow or wide. Primary vascular tissue cylinder consisting of four bundles. Secondary lateral growth normal, anomalous (from concentric cambia) or absent. Endodermis often prominent. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements usually libriform fibres (sometimes very long; sometimes fibre tracheids) with simple pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (sometimes homocellular). Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric or banded, or absent (rarely apotracheal diffuse or diffuse-in-aggregates). Wood elements sometimes storied. Tyloses sometimes abundant. Sieve tube plastids S type. Nodes usually 1:1 (sometimes 1:2 or 2:2), unilacunar with one or two leaf traces (sometimes bilacunar with two traces); nodes often swollen. Pericycle usually with sclerenchymatous cells. Medulla often with crystal inclusions. Heartwood sometimes with gum-like substances. Silica bodies present in wood ray cells in some representatives. Calciumoxalate as styloids, crystal sand and acicular or prismatic crystals sometimes frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, sometimes stellate or dendritic (rarely lepidote); often multicellular glandular hairs (above all in Nepetoideae), sometimes containing ethereal oils.

Leaves Usually opposite (sometimes verticillate, rarely alternate), usually simple (sometimes pinnately or palmately compound), entire or lobed, often coriaceous, often ericoid, with various ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection usually arcuate (sometimes annular). Venation pinnate or palmate, usually eucamptodromous to semicraspedodromous (sometimes brochidodromous), or leaves one-veined. Stomata usually diacytic or anomocytic (sometimes diallelocytic with three subsidiary cells, anisocytic or paracytic, rarely other types). Cuticular wax crystalloids? Crystal inclusions of different kinds often present. Leaf margin usually serrate, crenate or lobate (rarely entire). Glandular hairs with ethereal oils, etc. present. Extrafloral nectaries present in some genera on abaxial side of lamina and petiole.

Inflorescence Terminal or axillary, usually panicle, raceme-, spike- or head-like thyrse, often with whorls of dichasial or circinate partial inflorescences (flowers rarely solitary axillary). Bracts and floral prophylls (bracteoles) often large and petaloid.

Flowers Usually zygomorphic (rarely resupinate; rarely actinomorphic). Hypogyny. Sepals (four or) five (to nine), usually with imbricate or open (rarely valvate) aestivation, often unequal in size, usually persistent and often accrescent, connate (sometimes bilabiate); calciumoxalate crystals often frequent. Petals (four or) five (to 16), with imbricate aestivation, connate into usually bilabiate (usually two upper and three lower petals; in Ajugoideae five lower petals; sometimes unilabiate) corolla. Nectariferous disc intrastaminal, entire or lobed, often developed on abaxial side only (rarely absent).

Androecium Stamens usually two longer and two shorter (sometimes two, rarely five, six or up to 16), antesepalous, alternipetalous. Filaments usually free from each other (rarely connate), adnate to corolla tube (epipetalous). Anthers often connivent, basifixed or dorsifixed, often versatile, usually tetrasporangiate (in ‘Salvia’ disporangiate; in Ocimeae synthecal), usually extrorse (sometimes introrse?), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores); connective often prolonged (in ‘Salvia’ modified into lever-shaped structure). Tapetum secretory, with multinucleate cells. Staminodia usually absent (sometimes one adaxial-median or two adaxial-lateral or abaxial-lateral staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (rarely tetra- or pentacolpate, tricolporate? or triporate; in Nepetoideae hexacolpate, rarely octacolpate), shed as monads, usually bicellular (in Nepetoideae tricellular) at dispersal. Exine tectate or semitectate, with usually columellate (in some Ajugoideae granular) infratectum, microperforate to reticulate, rugulate, psilate-punctate, granulate or spinulate, or smooth.

Gynoecium Pistil composed of two (to five) connate carpels. Gynophore present in some genera. Ovary superior, bilocular (to quinquelocular; sometimes pseudomonomerous), during maturation with locules usually bipartite by anterio-posterior secondary septa through ingrowth of ovary wall (ovary secondarily quadrilocular; in Symphorematoideae incompletely quadrilocular). Style single, usually bifid at apex (sometimes unequally bilobate), usually gynobasic (sometimes terminal). Stigmas usually punctate (rarely one capitate or quadrilobate stigma), papillate, usually Dry (sometimes Wet) type. Pistillodium absent.

Ovules Placentation usually basal or basal to axile (rarely apical; in Symphorematoideae free central), attached to sides of inrolled carpellary walls (cf. Verbenaceae). Ovules two per carpel, usually anatropous (sometimes hemianatropous, rarely orthotropous), usually ascending (in Symphorematoideae pendulous), apotropous or epitropous, unitegmic, tenuinucellar. Integument five to nine cell layers thick. Funicular obturator present or absent. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes persistent. Endosperm development cellular. Endosperm haustoria chalazal and micropylar (sometimes absent). Embryogenesis usually onagrad (sometimes asterad).

Fruit A drupe (Symphorematoideae, Viticoideae) or a schizocarp with usually four one-seeded (sometimes two two-seeded) usually nutlike (in Prasium drupaceous) mericarps (rarely a capsule), with persistent and sometimes accrescent calyx. Exocarp in Nepetoideae usually with mucilaginous idioblasts.

Seeds Aril absent. Exotestal cells often elongate, usually with radial and often inner walls thickened. Hypodermal cells sometimes sclerenchymatous. Endotesta unspecialized? Perisperm not developed. Endosperm usually sparse (sometimes copious) or absent. Embryo usually straight (sometimes curved), oily (rich in linolic acid or linolenic acid), well differentiated, without chlorophyll. Cotyledons two, flat. Radicula usually directed downwards towards fruit base. Germination phanerocotylar or cryptocotylar.

Cytology n = 5–13, 15–18, 23–26 (up to 120) – Polyploidy frequently occurring.

DNA Deletion in plastid gene matK. Mitochondrial coxI intron present.

Phytochemistry Flavonols, 6- and/or 8-hydroxylated flavone glycosides, 6- or 8-hydroxyflavones or 6-methoxyflavones, apigenin- and luteolin-derived flavonoids, Route II iridoids (also C4-decarboxylated iridoids: iridoid glycosides, glycosides of monoterpenoid lactones; especially in Lamioideae, Viticoideae, Ajugoideae, and Scutellarioideae; harpagide and harpagioside in, e.g., Caryopteris), Route I carbocyclic iridoids (aucubin in Vitex, catalpol, melittoside), Route II carbocyclic iridoids (galiridoside, lamiide, caryoptoside, ipolamiide, lamalbide, ajugol, ajugoside, lamiol, lamioside), Group X secoiridoids (desoxyloganin, nepeta lactones), ethereal oils consisting of monoterpenoids, sesquiterpenoids, phenylpropanoids etc. (above all in Nepetoideae), diterpenoids (labdanes, neoclerodanes, abietanes, primaranes, ent-kaurans etc.; in Viticoideae, Ajugoideae, Prostantheroideae, Scutellarioideae, Lamioideae, Nepetoideae), triterpenoids, ursolic acid, caffeic acid esters (phenylethyl caffeoylic glucosides, e.g. verbascosides, acteoside, cornoside; in Ajugoideae, Lamioideae, Prostantheroideae, Scutellarioideae, and Viticoideae; rosmarinic acid in Nepetoideae), betaines (e.g. glycine betaine, particularly in Lamioideae, Ajugoideae and Nepetoideae), alkaloids, triterpene saponins, cyanogenic compounds, and shikimic acid derived arthroquinones (in Tectona) present. Ellagic acid, tannins, and proanthocyanidins not found. Carbohydrates stored as stachyose and other oligosaccharides. Cell walls containing arabinoxyloglucans and often galactoxyloglucan hemicelluloses.

Use Ornamental plants, spices, perfumes (Lavandula, Marrubium, Mentha, Pogostemon, Rosmarinus, Salvia), medicinal plants, honey (Phacelia etc.), timber (Viticoideae, Ajugoideae, Tectona grandis, etc.), seed-oils (Perilla etc.).

Systematics Lamiaceae are sister-group to a clade with the following topology: [Mazaceae+[Phrymaceae+[Paulowniaceae+[Rehmanniaceae+Orobanchaceae]]]].

Gynobasic style and schizocarp with four nutlike mericarps have evolved several times in Lamiaceae.

A probable topology is (Li & al. 2016) [[Callicarpa+Prostantheroideae]+[[Symphorematoideae+Viticoideae]+Nepetoideae+[Tectona+[Premnoideae+Ajugoideae+[Peronematoideae+[Scutellarioideae+[Cymarioideae+Lamioideae]]]]]]]].

Calliprostantherina B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 14. Oct 2016

Calliprostantherina consist of Callicarpa and Prostantheroideae.

Callicarpa clade

1/c 140. Callicarpa (c 140; tropical and subtropical regions on both hemispheres). – Callicarpa has branched/stellate hairs, and actinomorphic 4–5(–7)-merous flowers. – Bendiksby & al. (2011) found Callicarpa to be sister of all other Lamiaceae investigated.

Prostantheroideae Luerss., Handb. Syst. Bot. 2:1014, 1016. Sep 1882 [’Prostanthereae’]

14/310–320. Australia. Sometimes aromatic. Flowers usually actinomorphic (sometimes zygomorphic), tetramerous to octamerous. Nectariferous disc sometimes absent. Staminodia sometimes two. Endosperm present. n = ?

Westringieae Bartl., Ord. Nat. Plant.: 182. Sep 1830 [’Westringiea’]

5/210–215. Prostanthera (>100; Australia, Tasmania), Westringia (25–30; Australia, Tasmania, Lord Howe), Hemiandra (14; southwestern Western Australia), Microcorys (c 20; southwestern Western Australia), Hemigenia (c 50; Australia). – Australia, Tasmania, Lord Howe.

Chloantheae Benth. et Hook. f., Gen. Plant. 2: 1132. 1-16 Mai 1876

9/100–105. Brachysola (2; B. coerulea, B. halganiacea; southwestern Western Australia), Physopsis (5; P. chrysophylla, P. chrysotricha, P. lachnostachya, P. spicata, P. viscida; southwestern Western Australia), Newcastelia (10; Australia), Lachnostachys (6; L. albicans, L. bracteosa, L. coolgardiensis, L. eriobotrya, L. ferruginea, L. verbascifolia; Western Australia), ’Pityrodia’ (c 40; Western Australia, Northern Territory, eastern Queensland; polyphyletic), Dicrastylis (c 25; Australia), Cyanostegia (5; C. angustifolia, C. corifolia, C. cyanocalyx, C. lanceolata, C. microphylla; Western Australia, southern Northern Territory), Hemiphora (5; H. bartlingii, H. elderi, H. exserta, H. lanata, H. uncinata; southwestern Western Australia), Chloanthes (4; C. coccinea, C. glandulosa, C. parviflora, C. stoechadis; southwestern Western Australia, eastern Queensland, eastern New South Wales). – Australia. – Chloantheae have occasionally been identified as sister to Wenchengia (here in Scutellarioideae), and these as sister-group to Ajugoideae.

[[Symphorematoideae+Viticoideae]+Nepetoideae+[Tectona+[Premnoideae+Ajugoideae+[Peronematoideae+[Scutellarioideae+[Cymarioideae+Lamioideae]]]]]]

Viticisymphorina B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 13. Oct 2016

Fruit a drupe. – Viticisymphorina comprise Symphorematoideae and Viticoideae.

Symphorematoideae Briq. in Engler et Prantl, Nat. Pflanzenfam. IV, 3a: 144. 26 Feb 1895 [‘Symphoremoideae’]

3/c 30. Congea (11; northeastern India to Yunnan and Southeast Asia, West Malesia), Sphenodesme (c 15; northeastern India to southern China and Southeast Asia, West Malesia), Symphorema (3; S. involucratum, S. luzonicum, S. polyandrum; India, Sri Lanka, Yunnan, Burma, the Andaman Islands, Thailand, the Philippines, the Moluccas). – India to West Malesia. Lianas. Inflorescence three- to seven-flowered capitate cymose, with involucre consisting of more or less petaloid showy bracts. Flowers actinomorphic. Calyx 5–8-lobate. Corolla 5–16-lobate. Nectariferous disc absent. Stamens four to 18. Ovary incompletely bilocular. Placentation free central to apical. Ovules orthotropous, pendulous. Funicle absent. Megagametophyte present on surface of ovule. Endosperm absent? n = 12, 14, 17, 18. Diterpenoids absent? – Congea has occasionally been recovered as sister to the remaining Lamiaceae.

Viticoideae Briq. in Engler et Prantl, Nat. Pflanzenfam. IV, 3a: 144. 26 Feb 1895

4/c 285. ’Vitex’ (c 250; temperate to tropical regions on both hemispheres; non-monophyletic), Petitia (2; P. domingensis, P. urbanii; the West Indies), Pseudocarpidium (9; the West Indies, with their largest diversity on Cuba), Teijsmanniodendron (23; Southeast Asia, the Nicobar Islands, Malesia to New Guinea, the Bismarck Archipelago and Solomon Islands). – Temperate to tropical regions on both hemispheres.

Nepetoideae Burnett, Outlines Bot.: 971. Feb 1835 [’Nepetidae’]

105/3.540–3.840. Elsholtzieae Burnett, Outlines Bot.: 971. Feb 1835 [’Elscholzieae’]. Ombrocharis (1; O. dulcis; Hunan), Perillula (1; P. reptans; Japan); Elsholtzia (c 40; temperate regions and tropical mountains in the Old World), Collinsonia (11; China, Japan, Taiwan, southeastern Canada, eastern and southeastern United States), Mosla (c 15; the Caucasus, eastern India and the Himalayas to Burma, China, the Korean Peninsula Japan, the Ryukyu Islands, the Kuril Islands and the Russian Far East, Southeast Asia, Malesia to the Philippines), Perilla (1–4; P. frutescens; India and Bhutan to China, the Korean Peninaula and Japan, Taiwan, Southeast Asia, Java). – Mentheae Dumort., Fl. Belg.: 48. 1827. Salvia (900–1.000; temperate to tropical regions on both hemispheres, with their highest diversity in western North America, southwestern and Central Asia, the Himalayas and western China), Lepechinia (40–45; tropical and subtropical America); Lycopus (19; Europe, northwestern Asia, temperate North America, one species, L. australis, in southeastern South Australia to southeastern Queensland, Tasmania), Hyssopus (6; H. ambiguus, H. cuspidatus, H. latilabiatus, H. macranthus, H. officinalis, H. seravschanicus; southern Europe, the Mediterranean to Pakistan, Central Asia and Mongolia), Mentha (18–19; nearly cosmopolitan except South America), Thymbra (4; T. calostachya, T. capitata, T. sintenisii, T. spicata; the Mediterranean to Iran), Thymus (c 350; Europe, temperate Asia), Origanum (40–45; Europe, the Mediterranean, temperate Asia), Zataria (1; Z. multiflora; Iran, Afghanistan, Pakistan, Kashmir), Pentapleura (1; P. subulifera; Kurdistan), Satureja (c 45 or 200–210; nearly cosmopolitan, with their highest diversity in temperate regions in the Old World), Killickia (4; K. compacta, K. grandiflora, K. lutea, K. pilosa; Drakensberg, KwaZulu-Natal), Gontscharovia (1; G. popovii; Central Asia), Saccocalyx (1; S. saturejoides; Morocco, Algeria), Bystropogon (7; B. canariensis, B. maderensis, B. odoratissimus, B. origanifolius, B. plumosus, B. punctatus, B. wildpretii; Madeira, the Canary Islands), Cuminia (1; C. eriantha; Robinson Crusoe Island in the Juan Fernandez Islands), Minthostachys (17; the Andes from Venezuela to central Argentina), Cyclotrichium (9; southwestern Asia to Iran), Obtegomeria (1; O. caerulescens; Sierra Nevada de Santa Marta in northeastern Colombia), Kurzamra (1; K. pulchella; the Andes in Chile and northwestern Argentina), Ziziphora (17; the Mediterranean and the Caucasus to Russia, Central Asia, Afghanistan, Mongolia, Siberia and the Himalayas), Stachydeoma (1; S. graveolens; northwestern Florida), Hedeoma (c 40; southwestern Canada, United States, Mexico, Guatemala, subtropical South America), Rhododon (1; R. ciliatus; Texas), Pogogyne (7; P. abramsii, P. clareana, P. douglasii, P. floribunda, P. nudiuscula, P. serpylloides, P. ziziphoroides; southern Oregon to Baja California), Poliomintha (8; southwestern United States, northwestern Mexico), Hesperozygis (7; H. dimidiata, H. kleinii, H. myrtoides, H. nitida, H. rhododon, H. ringens, H. spathulata; southern Brazil), Glechon (7; G. ciliata, G. discolor, G. elliptica, G. hoehneana, G. marifolia, G. spathulata, G. thymoides; southern Brazil, Uruguay, Paraguay, northern Argentina), Hoehnea (4; H. epilobioides, H. minima, H. parvula, H. scutellarioides; southern Brazil, Paraguay), Rhabdocaulon (7; R. coccineum, R. denudatum, R. erythrostachys, R. gracile, R. lavanduloides, R. stenodontum, R. strictum; southern Brazil, Uruguay, Paraguay, northeastern Argentina), Eriothymus (1; E. rubiaceus; Minas Gerais in Brazil), Cunila (18; eastern and central United States, Mexico, Central America, tropical South America to Uruguay and Argentina), Piloblephis (1; P. rigida; Georgia, Florida, the Bahamas), Conradina (7; C. brevifolia, C. canescens, C. cygniflora, C. etonia, C. glabra, C. grandiflora, C. verticillata; southeastern United States), Dicerandra (6–11; D. densiflora, D. frutescens, D. fumella, D. linearifolia, D. odoratissima, D. radfordiana; southeastern United States), Monardella (c 35; southwestern Canada, western United States), Pycnanthemum (c 20; southeastern Canada, eastern and southeastern United States), Acanthomintha (4; A. duttonii, A. ilicifolia, A. lanceolata, A. obovata; California, northern Baja California), Monarda (c 20; southern Canada, United States, northern Mexico), Blephilia (3; B. ciliata, B. hirsuta, B. subnuda; southern Canada, eastern and central United States), Prunella (8; Europe, the Mediterranean and North Africa to the Caucasus, Russia and temperate Asia, one species, P. vulgaris, also in North America), Cleonia (1; C. lusitanica; western Mediterranean to Algeria), Horminum (1; H. pyrenaicum; the Alps, the Pyrenees); Nepeta (200–250; Europe, the Canary Islands, the Mediterranean, North Africa, mountains in tropical Africa, temperate Asia), Drepanocaryum (1; D. sewerzowii; Central Asia), Lophanthus (20–25; mountains in Iran, western Himalayas, Central Asia to Siberia and China), Hymenocrater (10–12; the Caucasus, Iran, Afghanistan), Marmoritis (5; M. complanata, M. decolorans, M. nivalis, M. pharica, M. rotundifolia; Pakistan and the Himalayas to Tibet and China), Agastache (22; mountains and arid regions in temperate Asia, Taiwan, southern Canada, United States and Mexico), Meehania (7; M. cordata, M. faberi, M. fargesii, M. henryi, M. montis-koyae, M. pinfaensis, M. urticifolia; China to Japan and the Russian Far East, eastern United States), Glechoma (7; G. biondiana, G. grandis, G. hederacea, G. hirsuta, G. longituba, G. sardoa, G. sinograndis; Europe, the Mediterranean, temperate Asia to the Russian Far East, Japan,Taiwan and Vietnam), Dracocephalum (60–70; Europe, the Mediterranean, North Africa, temperate Asia, North America), Lallemantia (5; L. baldshuanica, L. canescens, L. iberica, L. peltata, L. royleana; Turkey and the Caucasus to the Arabian Peninsula, Iran, Central Asia, the Himalayas and western Siberia), Cedronella (1; C. canariensis; Macaronesia except Cape Verde Islands), Schizonepeta (2; S. multifida, S. tenuifolia; southern Siberia, Mongolia, northern China), Melissa (4; M. axillaris, M. flava, M. officinalis, M. yunnanensis; Europe, the Mediterranean, North Africa, southwestern Asia to Iran, Central Asia, the Himalayas, Tibet, Yunnan, Southeast Asia and West Malesia, Taiwan). – Ocimeae Dumort., Anal. Fam. Plant.: 22. 1829 [’Ocymeae’]. Lavandula (45–50; Macaronesia, the Mediterranean, North Africa to Somalia, southwestern Asia to southeastern India); Isodon (c 105; tropical and subtropical regions in Africa and Asia)?, Siphocranion (2; S. macranthum, S. nudipes; China, Tibet, Burma, Southeast Asia; in Hanceola?)?, Hanceola (8; China; incl. Siphocranion?)?, Hyptidendron (17–18; the Guayana Highlands and northeastern Brazil to Peru and Bolivia), Eriope (>30; tropical South America, with their highest diversity in central and eastern Brazil), Hypenia (c 25; northeastern South America to Venezuela and Bolivia), Marsypianthes (5; M. burchellii, M. chamaedrys, M. foliolosa, M. hassleri, M. montana; southern Mexico, Central America, the West Indies, tropical South America to Paraguay and Argentina), Hyptis (300–400; southern United States, Mexico, Central America, the West Indies, tropical and subtropical South America to Peru and Argentina), Physominthe (2; P. longicaulis, P. vitifolia; eastern and northeastern Brazil), Mesosphaerum (c 25; southern Mexico, Central America, western tropical South America), Cantinoa (23; southeastern United States, Central America, the West Indies, tropical South America to Argentina), Oocephalus (14; Brazil, eastern Bolivia), Condea (26; western United States, Central America, the West Indies, tropical South America), Cyanocephalus (25; Cuba, tropical South America to eastern Paraguay and Bolivia), Eplingiella (3; E. brightoniae, E. cuniloides, E. fruticosa; northeastern Brazil), Eriopidion (1; E. strictum; southern Venezuela, northeastern Brazil), Gymneia (6; G. chapadensis, G. interrupta, G. malacophylla, G. moniliformis, G. platanifolia, G. virgata; northeastern and central Brazil), Leptohyptis (5; L. calida, L. leptostachys, L. macrostachys, L. pinheiroi, L. siphonantha; Brazil, especially northeastern parts), Martianthus (4; M. elongatus, M. leucocephalus, M. sancti-gabrielii, M. stachydifolius; northeastern Brazil, Huarochi in Peru), Medusantha (8; northeastern and central Brazil), Rhaphiodon (1; R. echinus; northeastern Brazil), Asterohyptis (4; A. mocinoana, A. nayarana, A. seemannii, A. stellulata; Mexico to Costa Rica), Ocimum (c 65; tropical and subtropical regions on both hemispheres, with their largest diversity in Africa), Syncolostemon (c 45; southern and southeastern Africa), Orthosiphon (40–45; tropical regions in the Old World; incl. Heterolamium?), Heterolamium (2; H. debile, H. flaviflorum; China; in Orthosiphon?), Fuerstia (9; tropical East and southern Africa), Benguellia (1; B. lanceolata; Angola), Catoferia (4; C. capitata, C. chiapensis, C. martinezii, C. spicata; southern Mexico, Central America, Colombia to Peru), Endostemon (20; tropical and southern Africa and Madagascar to the Arabian Peninsula and India), Haumaniastrum (c 35; tropical Africa, one species, H. villosum, on Madagascar), Platostoma (c 45; tropical Africa, Madagascar, tropical Asia to New Guinea), Dauphinea (1; D. brevilabra; Madagascar), Capitanopsis (3; C. albida, C. angustifolia, C. cloiselii; Madagascar), Madlabium (1; M. magenteum; northern Madagascar), Plectranthus (c 350; tropical and subtropical Africa, Madagascar, tropical Asia to tropical Australia and islands in the Pacific), Thorncroftia (6; T. greenii, T. longiflora, T. lotteri, T. media, T. succulenta, T. thorncroftii; Northern Province, Mpumalanga), Tetradenia (c 20; tropical and southern Africa, Madagascar), Anisochilus (17; tropical Asia from India and Sri Lanka to Indochina), Leocus (5; L. africanus, L. caillei, L. lyratus, L. membranaceus, L. pobeguinii; tropical Africa), Aeollanthus (40–45; tropical and subtropical Africa), Alvesia (3; A. clerodendroides, A. cylindricalyx, A. rosmarinifolia; Central Africa), Pycnostachys (c 35; tropical and southern Africa, one species, P. coerulea, on Madagascar). – Cosmopolitan, with their highest diversity in warm-temperate regions. Usually aromatic herbs. Prismatic calcium oxalate crystals present in calyx epidermis. Pollen grains hexacolpate, tricellular at dispersal. Style gynobasic. Endosperm development highly asymmetrical, two haustoria present adjacent to each other. Exocarp with mucilaginous cells (myxocarpy) producing hygroscopic spiral fibrils. Endosperm single-layered. Cotyledons enclosing embryo. Seeds with special fatty acids. n = 6 or more. Volatile terpenoids, rosmarinic acid, nepetoidin A and B (caffeic acid esters), and betaine (sparse) present. Iridoid glucosides and acteosides usually absent. – The polyphyletic ’Salvia’ is represented by at least three different clades in Mentheae. Lavandula is sister to the remaining Ocimeae (Li & al. 2016). The clade [Ombrocharis+Perillula] is sister-group to the rest of Elsholtzieae (Chen & al. 2016). Ombrocharis dulcis consists of aromatic tuberous perennial herbs with 11-veined two-lipped calyx, two lipped corolla with deeply bilobate upper lip and trilobate lower lip, and schizocarp with nutlike mericarps.

[Tectona+[Premnoideae+Ajugoideae+[Peronematoideae+[Scutellarioideae+[Cymarioideae+Lamioideae]]]]]

Tectona clade

1/3. Tectona (3; T. grandis, T. hamiltoniana, T. philippinensis; Pakistan, India, Burma, southern China, Southeast Asia, the Philippines). – Tropical Asia to southern China and the Philippines. Trees, shrubs or herbs. Hairs sometimes branched. Leaves sometimes compound. Nectariferous disc poorly developed or absent. Fruit a drupe. – Tectona has shikimic acid derived arthroquinones.

[Premnoideae+Ajugoideae+[Peronematoideae+[Scutellarioideae+[Cymarioideae+Lamioideae]]]]

Premnoideae B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 10. Oct 2016

3/170–180. Cornutia (8; southern Mexico, Central America, the West Indies, tropical South America); Premna (130–140; tropical Africa, Madagascar, Indian Ocean islands, tropical Asia to tropical Australia and islands in the Pacific), Gmelina (c 35; Pakistan, India, Sri Lanka, southern China, Southeast Asia, Malesia to New Guinea, tropical Australia, Solomon Islands, New Caledonia, Fiji). – Pantropical. Premnoideae have a drupe with a single four-seeded pyrene. – Cornutia is sister to [Premna+Gmelina].

Ajugoideae Luerss., Handb. Syst. Bot. 2: 1016. Sep 1882

23/740–770. Monochilus (2; M. gloxinifolius, M. obovatus; Brazil)?, Karomia (9; eastern and southern Africa, Madagascar, one species, K. fragrans, in Vietnam), Discretitheca (1; D. nepalensis; Nepal), Rotheca (35–60; tropical and southern Africa, tropical Asia), Glossocarya (10; Sri Lanka, Burma, Southeast Asia, New Guinea, eastern Queensland); Ajuga (35–40; temperate to tropical regions in the Old World to eastern Siberia, Japan, Australia and islands in western Pacific), Pseudocaryopteris (3; P. bicolor, P. foetida, P. paniculata; India, the Himalayas, Burma, southern China, Southeast Asia), Tripora (1; T. divaricata; China, the Korean Peninsula, Japan), Amethystea (1; A. caerulea; Central Asia, southern Siberia, Mongolia, China, the Korean Peninsula, Japan), Trichostema (17–19; southern Canada, United States, Mexico), Caryopteris (7; C. forrestii, C. glutinosa, C. incana, C. jinshajiangensis, C. mongholica, C. tangutica, C. trichosphaera; Central Asia, Tibet, Mongolia, China, the Korean Peninsula, Japan, Taiwan); Hosea (1; H. lobbii; northwestern Borneo), Oxera (24; Borneo to New Guinea, the Bismarck Archipelago, Solomon Islands, Queensland, New Caledonia, Vanuatu, Fiji, Samoa, Tonga), Ovieda (1; O. spinosa; Hispaniola), Aegiphila (c 150; Florida, southern Mexico, Central America, the West Indies, tropical South America), Clerodendrum (c 150; tropical and subtropical regions on both hemispheres), Tetraclea (1; T. coulteri; southern Arizona, western Texas, northern Mexico), Amasonia (6; A. angustifolia, A. arborea, A. calycina, A. campestris, A. hirta, A. obovata; Trinidad, tropical South America), Kalaharia (1; K. uncinata; tropical and southern Africa), Volkameria (c 30; tropical regions on both hemispheres); Schnabelia (5; S. aureoglandulosa, S. nepetifolia, S. oligophylla, S. terniflora, S. tetradonta; southwestern and southern China), Rubiteucris (2; R. palmata, R. siccanea; northeastern India, the Himalayas, Tibet, Burma, China, Taiwan), Teucrium (c 250; nearly cosmopolitan, with their highest diversity in the Mediterranean). – Subcosmopolitan (abundant in temperate areas), with their highest diversity in Southeast Asia to Australia. Sometimes aromatic. Flowers actinomorphic or zygomorphic (unilabiate, in Teucrium 0:5), in Aegiphila tetramerous. Nectariferous disc poorly developed or absent. Exine spinulate or verrucate, usually with branched (sometimes simple, granulate, etc.) columellae. Style often terminal. Antipodal cells sometimes numerous. Fruit a schizocarp with achene-like mericarps. Endosperm multilayered or absent. Cotyledons enclosing rest of embryo. n = 7, 10, 13, 14, 16 or more. Terpenoids sometimes absent. – The clade [Karomia+Rotheca] was recovered as sister-group to the remaining Ajugoideae (Li & al. 2016). However, Glossocarya and Discretitheca were not included in their analyses.

Perolamiina B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 13. Oct 2016

[Peronematoideae+[Scutellarioideae+[Cymarioideae+Lamioideae]]]

Peronematoideae B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 11. Oct 2016

4/17. Peronema (1; P. canescens; southern Burma to Sumatra and Borneo), Garrettia (1; G. siamensis; Yunnan, Thailand, Java), Hymenopyramis (7; H. acuminata, H. brachiata, H. cana, H. parvifolia, H. pubescens, H. siamensis, H. vesiculosa; Burma, Hainan, Thailand, Indochina), Petraeovitex (8; southern Burma and Thailand, the Malay Peninsula, Malesia to New Guinea, the Bismarck Archipelago, Solomon Islands and northeastern Queensland). – Shrubs, lianas or trees. Leaves simple or trilobate. Inflorescence in Garrettia axillary cyme with secondary monochasial branches. Flowers minute. Calyx quadridentate or quinquedentate. Corolla infundibuliform, unequally quadrilobate, or two-lipped, with lower lip trilobate and upper lip usually bilobate. Nectariferous disc absent or almost so. Stamens didynamous. Ovary bilocular when young, later secondarily quadrilocular, unlobed. Ovules one or two per locule. Stigma bilobate. Fruit a schizocarp with four single-seeded mericarps and inflated persistent calyx.

Scutelamiina B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 13. Oct 2016

[Scutellarioideae+[Cymarioideae+Lamioideae]]

Scutellarioideae Prantl, Handb. Bot.: 293. 1 Mar-15 Apr 1880 [’Scutellariae’]

5/325–385. Wenchengia (1; W. alternifolia; Hainan); Holmskioldia (3; H. gigas, H. speciosa, H. tettensis; the Himalayas), Renschia (1; R. heterotypica; northern Somalia), Tinnea (c 20; tropical and southern Africa), Scutellaria (300–360?; nearly cosmopolitan). – Subcosmopolitan. Inflorescence racemose. Calyx usually bilabiate (not in Holmskioldia), with rounded lobes and with numerous xylem fibres. Pericarps with tuberculate or elongate processes. Testa tuberculate. Endosperm various. n = 12 or more. Terpenoids sometimes absent. – Wenchengia is sister to the remaining Scutellarioideae. In Wenchengia the leaves are spirally arranged, the calyx bilobed and quinquedentate, and the style terminal.

Cymalamiina B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 12. Oct 2016

[Cymarioideae+Lamioideae]

Cymarioideae B. Li, R. G. Olmstead et P. D. Cantino in Sci. Rep. 6:34343: 12. Oct 2016

2/3. Cymaria (2; C. dichotoma, C. elongata; Hainan, Indochina, the Malay Peninsula to New Guinea), Acrymia (1; A. ajugiflora; Perak and Selangor on the Malay Peninsula). – Southeast Asia. Cymes axillary, with long peduncle and secondary monochasial branches. Ovary shallowly quadrilobate. Nutlets reticulately patterned.

Lamioideae Harley in Kew Bull. 58: 765. 26 Nov 2003

50/1.295–1.470. Pogostemoneae Briq. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. IV, 3a: 208. Dec 1895. Holocheila (1; H. longipedunculata; Yunnan), Colebrookea (1; C. oppositifolia; the Himalayas to Burma, western China and Thailand); Craniotome (1; C. furcata; the Himalayas, Tibet and southern China to Laos and Vietnam), ’Microtoena’ (c 25; the Himalayas, Tibet, China, the Korean Peninsula, Southeast Asia to Java; paraphyletic), Anisomeles (5; A. candicans, A. heyneana, A. indica, A. malabarica, A. salviifolia; Madagascar, islands in the Indian Ocean, tropical Asia to Tibet, China and New Guinea, the Bismarck Archipelago, tropical Australia), Pogostemon (80–85; southern tropical Africa, East Asia to Japan, tropical Asia and Australia); Eurysolen (1; E. gracilis; northeastern India to Yunnan, Burma, Thailand, West Malesia), Leucosceptrum (1; L. canum; the Himalayas, Tibet, Burma and southern China to northern Thailand and northern Indochina), Comanthosphace (5; C. formosana, C. japonica, C. nanchuanensis, C. ningpoensis, C. stellipila; China, Japan, Taiwan), Achyrospermum (c 25; tropical regions in the Old World). Stamens four, of about equal length. Style gynobasic. Ovule in, e.g., with glandular hairs. Megagametophyte with micropylar lobe longer and wider than chalazal lobe. Endosperm multilayered. Embryo spatulate. n = 6 or more. Seed oils with laballenic fatty acids. – Colquhounia clade Colquhounia (5; C. coccinea, C. compta, C. elegans, C. seguinii, C. vestita; southern Himalayas, Tibet, Burma, southwestern China, Thailand, Indochina). – Gomphostemmateae Scheen et Lindqvist in A.-C. Scheen, M. Bendiksby, O. Ryding, C. Mathiesen, V. A. Albert et C. Lindquist, Ann. Missouri Bot. Gard. 97: 216. 9 Jul 2010. Gomphostemma (c 40; northeastern India, Burma, southern China, Southeast Asia, the Andaman Islands, the Malay Peninsula, Malesia to the Lesser Sunda Islands, Taiwan), Chelonopsis (17; Pakistan, Tibet, China, the Korean Peninsula, Japan, Taiwan). – Synandreae Raf., Fl. Tellur. 3: 84. Nov-Dec 1837 [’Synandrines’]. Synandra (1; S. hispidula; eastern United States), Macbridea (2; M. alba, M. caroliniana; southeastern United States), Physostegia (12; southern and southwestern Canada, United States, Mexico), Warnockia (1; W. scutellarioides; Oklahoma, Texas, northwestern Mexico), Brazoria (3; B. arenaria, B. enquistii, B. truncata; southern and central Texas). – Betonica-Galeopsis clade Betonica (6; B. alopecuros, B. graeca, B. haussknechtii, B. macrantha, B. monieri, B. officinalis; Europe, the Mediterranean, North Africa, western and southwestern Asia to the Caucasus and northern Iran), Galeopsis (9; Europe to the Caucasus and Siberia). – Stachydeae Dumort., Fl. Belg.: 44. 1827. Melittis (1; M. melissophyllum; Europe, Turkey), Stachys (300–450 or 450–600; temperate and subtropical regions on both hemispheres, tropical mountains, absent from Australia; incl. Sideritis: c 140; temperate regions in the Old World south to Macaronesia and North Africa). – Paraphlomideae Bendiksby in Taxon 60: 481. 8 Apr 2011. Paraphlomis (c 25; China, Southeast Asia to Central Malesia, Taiwan), Ajugoides (1; A. humilis; Japan), Matsumurella (5; M. chinensis, M. kwangtungensis, M. szechuanensis, M. tuberifera, M. yangsoensis; China, Japan, the Ryukyu Islands, Taiwan). – Phlomideae Mathiesen in A.-C. Scheen, M. Bendiksby, O. Ryding, C. Mathiesen, V. A. Albert et C. Lindquist, Ann. Missouri Bot. Gard. 97: 216. 9 Jul 2010. Phlomis (80–85; the Mediterranean to Central Asia and western China), Eremostachys (c 130; southeastern Russia and the Caucasus to West and Central Asia, India, the Himalayas, Mongolia, western and central China). – Leonureae Dumort., Fl. Belg.: 46. 1827. Lagochilus (c 45; the Caucasus, Iran, Afghanistan, Central Asia to northwestern China and Mongolia), Chaiturus (1; C. marrubiastrum; Europe, Russia, Turkey and the Caucasus to Central Asia), Loxocalyx (3; L. ambiguus, L. quinquenervius, L. urticifolius; China, Japan), Leonurus (c 25; Europe, the Mediterranean, the Caucasus, temperate Asia to the Himalayas, Japan and Taiwan), Lagopsis’ (5; L. darwiniana, L. eriostachya, L. flava, L. marrubiastrum, L. supina; Kashmir, Central Asia, Tibet, China and Mongolia to Siberia and Japan; polyphyletic?), Panzerina (2; P. canescens, P. lanata; western Siberia, Mongolia, northern China). – Marrubieae Vis., Fl. Dalmat. 2: 214. 1847. Roylea (1; R. cinerea; western Himalayas); Acanthoprasium (2; A. frutescens: the Alps; A. integrifolium: Cyprus); Moluccella (8; the Mediterranean, the Caucasus, the Middle East to northwestern India); ’Ballota’ (c 30; Europe, the Mediterranean, North Africa, the Arabian Peninsula, western Asia, one species, B. africana, in southern Namibia and South Africa; polyphyletic), Marrubium (c 45; southeastern Europe, the Mediterranean, the Caucasus, the Middle East to Iran and Kashmir). – Lamieae Coss. et Germ., Fl. Desc. Anal. Paris. 311, 324. 22 Feb 1845 [’Lamioideae’]. ’Lamium’ (40–50; Europe, the Mediterranean, North Africa, temperate Asia to Kamchatka, Japan and Taiwan; polyphyletic), Stachyopsis (4; S. lamiiflora, S. maleolens, S. marrubioides, S. oblongata; Central Asia, western China), Eriophyton (6; E. nepalense, E. rhomboideum, E. staintonii, E. sunhangii, E. tuberosum, E. wallichii; the Himalayas, Central Asia, Tibet, Yunnan). – Leucadeae Scheen et Ryding in A.-C. Scheen, M. Bendiksby, O. Ryding, C. Mathiesen, V. A. Albert et C. Lindquist, Ann. Missouri Bot. Gard. 97: 217. 9 Jul 2010. Rydingia (4; R. integrifolia, R. limbata, R. michauxii, R. persica; Ethiopia, southern Arabian Peninsula, Iran, Pakistan), ’Leucas’ (130–135; tropical and southern Africa, the Arabian Peninsula, southern and tropical Asia to Japan and Malesia to Queensland; polyphyletic), Isoleucas (2; I. somala: Somalia; I. arabica: southeastern Yemen), Otostegia (9; Cameroon, northeastern Africa, the Arabian Peninsula and Israel to Jordan, Iran to Pakistan),’Leonotis’ (1[–9]; L. leonurus; southern Africa, Angola [tropical Africa]; polyphyletic), ’Acrotome’ (8; southern and southeastern Africa; paraphyletic). – Unplaced Lamioideae Paralamium (1; P. griffithii; northeastern India and Yunnan to northern Burma and northern Vietnam; in Pogostemoneae?), Pseudomarrubium (1; P. eremostachydioides; Karatau Mountains in Kazakhstan), Metastachydium (1; M. sagittatum; Central Asia to western China). – Subcosmopolitan, with their highest diversity in temperate and subtropical regions in Europe and Asia. – Holocheila and Colebrookea form a sister-group to the rest of Pogostemoneae (Li & al. 2016). They have a two-lipped calyx with ten veins and the corolla lips are entire.

50% majority rule consensus tree (simplified) partitioned Bayesian analysis of Lamiaceae based on DNA sequence data (Bendiksby & al. 2011).

Phylogeny of Lamiaceae based on DNA sequence data (Li & al. 2016).

LENTIBULARIACEAE A. Rich.

( Back to Plantaginales )

Richard in P. A. Poiteau et P. J. F. Turpin, Fl. Paris. 1, ed. 4o: 26; ed. fol.: 23. 18 Jan 1808 [’Lentibulariae’], nom. cons.

Utriculariaceae Hoffmanns. et Link, Fl. Portug. 1: 62. 1 Sep 1809 [’Utriculinae’]; Lentibulariales Rich. ex Bercht. et J. Presl, Přir. Rostlin: 242. Jan-Apr 1820 [‘Lentibulariae’]; Lentibulariineae Link, Handbuch 1: 511. 4-11 Jul 1829 [‘Lentibulariae’]; Pinguiculaceae Dumort., Anal. Fam. Plant.: 19, 23. 1829; Pinguiculales Dumort., Anal. Fam. Plant.: 19. 1829 [’Pinguicularieae’]; Utriculariales Döll, Rhein. Fl.: 283. 24-27 Mai 1843

Genera/species 3/320–330

Distribution Cosmopolitan except polar and arid regions.

Fossils Unknown.

Habit Bisexual, perennial or annual herbs. Roots present (Pinguicula) or absent (Genlisea, Utricularia). Mycorrhiza absent. Aquatic or helophytic; some species are epiphytic. Carnivorous. Stem in Genlisea and Utricularia photosynthesizing. Leaves in Utricularia with bladder-shaped suction traps (in terrestrial species with chemical attractants), in Genlisea modified into ‘eel trap’-like structures. Tubers or rhizomes present in many terrestrial species of Utricularia.

Vegetative anatomy Mycorrhiza absent. Radicula degenerating very soon after germination. Lateral roots at least usually without root cap. Phellogen absent. Stem vascular tissue in Genlisea and Utricularia not organized into vascular bundles (phloem and xylem independently distributed). Vascular bundles in Pinguicula cylindrical. Secondary lateral growth absent. Xylem weakly developed. Vessel elements with ? perforation plates; lateral pits? Imperforate tracheary xylem elements? Wood rays absent. Axial parenchyma? Sieve tube plastids S type. Nodes 1:?, unilacunar with ? leaf traces. Calciumoxalate crystals and crystalloids present in Pinguicula and Utricularia.

Trichomes Hairs unicellular or multicellular, uniseriate; glandular hairs stalked or sessile; each secretory gland attached to a single epidermal cell without contact with vessel elements.

Leaves Usually alternate (spiral, often arranged in rosette; in Utricularia often opposite or verticillate), simple or compound (in Utricularia extremely variable), entire or lobed (in Utricularia sometimes peltate leaf-like organs), with adaxial glands or with special traps secreting proteolytic enzymes, with ? ptyxis; leaf dimorphism present in Genlisea and Utricularia (leaves sometimes absent). Genlisea with tubular leaves arising from basal rhizophylls and modified into bipartite ‘eel traps’, spirally twisted structures trapping (mostly unicellular) organisms; inner sides of ‘eel traps’ beset with rows of inwardly directed stiff hairs. Utricularia with small bladder-shaped ‘suction traps’ with lid at orifice; stimulation of one of four tactile-sensitive hairs at corolla mouth causing lid rapidly spring open (change in concentration of cytochrome c oxidase may increase respiration rate); Pinguicula with ‘fly paper traps’ with stalked and sessile glandular hairs on adaxial foliar surface; glandular hairs secreting mucilage and proteolytic enzymes. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually diacytic (sometimes anomocytic or anisocytic) or absent. Cuticular wax crystalloids absent? Epidermis with calciumoxalate crystals and crystalloids. Leaf margin serrate (numerous species of Utricularia) or entire.

Inflorescence Terminal or lateral, raceme or spike, or flowers solitary terminal.

Flowers Zygomorphic. Hypogyny. Sepals in Genlisea and Pinguicula five, equal or unequal in size, with imbricate or open aestivation, persistent, connate; in Utricularia usually connate into bilobate calyx (in subgenus Polypompholyx quadrilobate), often bilabiate, with open aestivation; median sepal adaxial. Petals five, with descending cochlear aestivation (abaxial lobe outside remaining lobes), connate into usually bilabiate (upper lip bilobate, lower lip trilobate) corolla, with abaxial nectar-secreting spur or bent inwards at base. Disc absent.

Androecium Stamens two (abaxial-lateral), antesepalous, alternipetalous. Filaments usually stout, free from each other, adnate to corolla tube (epipetalous). Anthers connivent (with thecae often confluent; sometimes superposed), dorsifixed, with ephemeral epidermal cells, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia usually absent (sometimes two adaxial-lateral).

Pollen grains Microsporogenesis simultaneous. Pollen grains (3–)4–8-colporate (in Genlisea violacea spiraperturate; in some species of Utricularia stephanocolporate), usually shed as monads (rarely tetrads), tricellular at dispersal. Exine tectate to semitectate, with columellate infratectum, perforate, microreticulate or reticulate, often regulate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, unilocular. Style, single, simple, hollow, short, or absent. Stigma non-uniformly broadly bilobate (bilabiate, with adaxial lobe reduced; sometimes sensitive), papillate, Wet type. Pistillodium absent.

Ovules Placentation free central or basal. Ovules usually numerous (in some species of Utricularia two) per carpel, anatropous, unitegmic, tenuinucellar (reduced, with meiocyte semi-inferior). Integument two to six cell layers thick. Megagametophyte monosporous, Polygonum type (sometimes protruding into micropyle). Antipodal cells sometimes persistent (uppermost cell sometimes strongly enlarged). Endosperm development cellular. Endosperm haustoria micropylar and (reduced) chalazal. Embryogenesis onagrad, asterad, or chenopodiad.

Fruit Usually a capsule, regularly dehiscing (with valves or pores, sometimes a pyxidium) or irregularly dehiscing (in some species of Utricularia a one-seeded nut).

Seeds Aril absent. Testa in some species of Utricularia winged or provided with hooks or hairs (in Genlisea possibly multiplicative). Exotestal cell walls thickened in different ways. Endotesta? Perisperm not developed. Endosperm absent (ab initio starchy). Embryo globular or ovoid, in Utricularia poorly differentiated when mature (two to 13 little differentiated cotyledon-like organs present), with chlorophyll. Cotyledons one or two (Pinguicula) or two often minute, undifferentiated. Germination phanerocotylar. Radicula ephemeral.

Cytology n = 7–12, 14, 15, 16, 21, 22, 24, 32. – Some species of Genlisea (e.g. G. margaretae) have the smallest known genome among angiosperms.

DNA Deletion in plastid gene matK. Mitochondrial coxI intron present. Mutation frequency in plastid gene matK in Utricularia and, above all, Genlisea among highest known in angiosperms; other genes in Lentibulariaceae also with high mutation frequency.

Phytochemistry Flavones (8-hydroxyapigenin, apigenin, luteolin, 6- and 8-hydroxyluteolin etc.), Route II decarboxylated iridoids, Group I carbocyclic iridoids (aucubin, catalpol), caffeic acid, iridoid glycosides, cyanogenic glycosides, and p-coumaride present. Alkaloids sometimes present. Flavonols, verbascosides, ellagic acid, proanthocyanidins, and saponins not found. Carbohydrates stored as stachyose and other oligosaccharides. Aluminium accumulated in some species of Utricularia.

Use Ornamental plants, nutrients (fermented viscous mild, using enzymes of Pinguicula vulgaris).

Systematics Pinguicula (c 80; temperate regions on the Northern Hemisphere, the Mediterranean, Central and South America), Genlisea (21; tropical and southern Africa, Madagascar, tropical America), Utricularia (220–230; cosmopolitan, with their highest diversity in Western Australia and tropical America).

The sister-group relationship of Lentibulariaceae is unresolved.

Pinguicula is sister to [Genlisea+Utricularia] (Jobson & al. 2003).

Insect-trapping in Byblidaceae, Lentibulariaceae and Plantaginaceae (Philcoxia) have evolved in parallel.

Cladogram of Lentibulariaceae based on DNA sequence data (Jobson & al. 2003).

LINDERNIACEAE (Reichb.) Borsch, K. Müller et E. Fischer

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Borsch, Müller et Fischer, Pl. Biol. (Stuttgart) 7: 76. 28 Jan 2005

Genera/species 21/155–165

Ditribution Tropical to warm-temperate regions in the Northern and Southern Hemispheres, with their largest diversity in tropical Africa and Southeast Asia.

Fossils Unknown.

Habit Bisexual, annual or perennial herbs (sometimes lignified at base, rarely shrubs). Chamaegigas intrepidus is poikilohydric aquatic resurrection plant.

Vegetative anatomy Phellogen? Stem often tetragonal in cross-section, with four longitudinal ridges with thin strands of lignified tissue inside. Vessel elements with simple? perforation plates; lateral pits? Vestured pits? Imperforate tracheary xylem elements libriform fibres? Wood rays? Axial parenchyma? Sieve tube plastids S type. Nodes 1:3, unilacunar with three leaf traces. Crystals?

Trichomes Hairs simple, unicellular or multicellular, or absent; glandular hairs often present, heads of glandular hairs with vertically divided cells.

Leaves Opposite, usually simple (in Scolophyllum sometimes pinnately compound), usually entire (rarely lobed), sometimes pairwise connate at base, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles?; petiole often with wing bundles. Venation pinnate or palmate. Stomata? Cuticular wax crystalloids? Leaf margin serrate, crenate or entire.

Inflorescence Terminal or axillary, raceme or head, or flowers solitary axillary. Floral prophylls (bracteoles) absent.

Flowers Zygomorphic (in Lindernia hypandra resupinate). Hypogyny. Sepals (four or) five, sometimes unequal in size, often winged, more or less connate (sometimes at base only). Petals four or five, connate into bilabiate to infundibuliform corolla, with glandular hairs on adaxial side (in ‘Lindernia’ with lobes covering anthers). Nectary? Disc?

Androecium Stamens usually four fertile or two fertile adaxial stamens and two abaxial staminodia (in Micranthemum two abaxial stamens). Filaments free from each other, adnate to corolla tube (epipetalous). Anthers parallel or head-to-head, dorsifixed, sometimes (in, e.g., Torenia) connate, non-versatile?, tetrasporangiate (in, e.g., Hemiarrhena and Torenia disporangiate), extrorse or introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia geniculate and Z-shaped or long and curved with clavate or spur-like appendage and blue and yellow glandular hairs, sometimes strongly reduced or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–5)-colpate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, smooth (‘Lindernia’).

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular. Style single, simple. Stigma bilobate, papillate, Wet type, usually sensitive. Pistillodium absent.

Ovules Placentation axile to basal. Ovules few to numerous per carpel, orthotropous, anatropous or hemitropous, unitegmic, tenuinucellar. Integument up to four cell layers thick. Megagametophyte monosporous, Polygonum type; often spathulate, in some species of ’Lindernia’ and Torenia bulging out of ovule. Endosperm development cellular? Endosperm haustoria? All endothelial cells curved inwards into endosperm; these may fuse making endosperm surface alveolated. Embryogenesis?

Fruit Usually a septicidal and septifragal capsule (in Torenia a septicidal and poricidal capsule with persistent and accrescent calyx).

Seeds Testa smooth or pitted. Aril absent? Exotesta? Endotesta? Perisperm not developed. Endosperm often ruminate (due to inpushings of endothelial cells), often alveolated, usually furrowed. Embryo?, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = 7–9, 12, 14, 17, 20, 21 (28) – Protein bodies absent from cell nuclei.

DNA

Phytochemistry Virtually unknown. Iridoids not found.

Use Ornamental plants, aquarium plants (Micranthemum).

Systematics Stemodiopsis (5; S. buchananii, S. eylesii, S. glandulosa, S. humilis, S. rivae; tropical Africa), Micranthemum (17–18; eastern United States, Mexico, Central America, Cuba, tropical South America), Bryodes (1; B. micrantha; Madagascar, Mauritius, the Seychelles), Psammetes (1; P. madagascariensis; tropical Africa, Madagascar), Torenia (40–50; tropical and southern Africa, Indian Ocean islands, Southeast Asia; incl. Legazpia?), Legazpia (1; L. polygonoides; East Asia to islands in western Pacific; in Torenia?), Encopella (1; E. tenuifolia; Cuba), Dintera (1; D. pterocaulis; Waterberg Plateau in Namibia), Bythophyton (1; B. indicum; Southeast Asia, Malesia), Artanema (5; A. bantamense, A. evrardii, A. fimbriatum, A. finetianum, A. longifolium; tropical Africa, Southeast Asia, Malesia), Picria (1; P. felterrae; southern China, Southeast Asia, Malesia), Pierranthus (1; P. capitatus; Indochina), Chamaegigas (1; C. intrepidus; Namibia), ’Lindernia’ (c 30; warm-temperate to tropical regions on both hemispheres; non-monophyletic; incl. Hemiarrhenia, Schizotorenia, Scolophyllum?), Hemiarrhena (1; H. plantaginea; northernmost Western Australia; in Lindernia?), Schizotorenia (2; S. atropurpurea, S. finetiana; Southeast Asia; in Lindernia?), Scolophyllum (2; S. spinifidium, S. ubonensis; Southeast Asia; in Lindernia?), Craterostigma (9; tropical and southern Africa, Madagascar, Yemen, Socotra, India), Crepidorhopalon (28; tropical and southern Africa, Madagascar), Hartliella (4; H. bampsii, H. capitata, H. cupricola, H. suffruticosa; Katanga Province in Congo), Bampsia (2; B. lawalreana, B. symoensiana; Katanga Province in Congo).

The sister-group relationship of Linderniaceae is unresolved. In Schäferhoff & al. (2010) it is part of trichotomy also including Byblis (Byblidaceae) and the remaining Plantaginales “beyond” Scrophulariaceae.

Cladogram of Linderniaceae based on DNA sequence data (Rahmanzadeh & al. 2005).

Phylogeny of Linderniaceae based on DNA sequence data (Schäferhoff & al. 2010).

MARTYNIACEAE Horan.

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Horaninov, Char. Ess. Fam.: 130. 30 Jun 1847 [’Bignoniaceae s. Martyniaceae’], nom. cons.

Genera/species 5/13

Distribution Warm and arid or semiarid regions from southern United States and Mexico to Argentina.

Fossils Unknown.

Habit Bisexual, usually annual (rarely perennial) herbs (rarely shrubs, in Holoregmia with fleshy young stems). Craniolaria annua has a large root tuber. With densely spaced glandular hairs and evil-smelling. Mycorrhiza probably absent. Some species (Proboscidea, Ibicella) are possibly carnivorous; the glands are similar to those in Byblis and Lentibulariaceae, although this may be a precursor to insect-trapping.

Vegetative anatomy Phellogen superficial. Vessel elements with simple perforation plates; lateral pits alternate, simple pits? Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal scanty vasicentric. Wood non-storied. Sieve tube plastids Ss type. Nodes unilacunar? with ? leaf traces. Crystals absent in parenchyma.

Trichomes Hairs unicellular or multicellular, uniseriate; mucilage glandular hairs consisting of unicellular or multicellular uniseriate stalk and multicellular apical head with flattened apex.

Leaves Usually opposite (rarely alternate, spiral), simple, entire or palmately lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection strongly arcuate; petiole also with adaxial cortical and medullary bundles. Venation pinnate? or palmate. Stomata usually anomocytic or anisocytic (rarely paracytic or diacytic). Cuticular wax crystalloids? Leaf margin serrate or entire. Extrafloral nectaries?

Inflorescence Terminal raceme.

Flowers Zygomorphic, often large. Hypogyny. Sepals five, with imbricate quincuncial aestivation, free or connate; median sepal adaxial. Petals five, with descending imbricate aestivation, connate into quinquelobate, tubular and bilabiate (upper lip bilobate, lower lip trilobate) or somewhat non-uniform corolla. Nectariferous disc intrastaminal, annular.

Androecium Stamens usually two longer and two shorter (didynamous; sometimes two, equal in length). Median adaxial stamen staminodial (when four fertile stamens present); in Martynia one median adaxial and two adaxial lateral stamens staminodial (when two fertile stamens present), antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, connivent, with thecae separated 180°, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective with gland at apex. Tapetum secretory, with binucleate cells. Staminodia one or three.

Pollen grains Microsporogenesis simultaneous. Pollen grains inaperturate (or polyporate?), shed as monads, bicellular or tricellular at dispersal. Exine semitectate (sometimes intectate), subdivided into c. 20 to c. 40 platelets evenly distributed on tectal surface, with columellate infratectum, areolate (Ibicella, Proboscidea), annular (Craniolaria, sexine consisting of smooth mural rings supported by columellae encircling smooth to granulate aperturoid areas, sexine between rings consisting of rodlets, clavulae, pilulae, or cone-shaped elements) or reticulate (Martynia).

Gynoecium Pistil composed of two paracarp-connate carpels. Ovary superior, primarily unilocular, yet quadrilocular to multilocular due to secondary septa (often divided by gradually connate placentae). Style single, simple, filiform. Stigma bilobate, type? Pistillodium absent.

Ovules Placentation parietal with two T-shaped placentae. Ovules two to numerous per carpel, anatropous, pendulous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent. Endosperm development cellular. Endosperm haustoria micropylar and chalazal (Martynia). Embryogenesis onagrad (Catalpa variant).

Fruit An incompletely loculicidal capsule, usually with well developed beak-shaped apical part formed from sterile upper part of ovary. Exocarp and mesocarp fleshy, caducous when ripe. Endocarp lignified, with bristles at apex, sharp edges, and prickle-, comb-, horn- or spur-like outgrowths.

Seeds Aril? Testa subgelatinous or inner and radial testal cell walls with band of cellulose. Inner testal layers in Proboscidea lignified and tanniniferous; in Martynia radial and inner tangential cell walls of outer epidermis lignified, remaining layers crushed. Perisperm not developed. Endosperm thin or almost absent, oily and aleuroniferous. Embryo straight, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology n = 15, 16

DNA Deletion in plastid gene matK?

Phytochemistry Flavonols (kaempferol), flavone glucosides, Group I carbocyclic iridoids (catalpol), Group II iridoids (harpagide, harpagioside and other 8β-8α-methylsubstituted iridoids, 10-hydroxylated carboxylic iridoids), and caffeic acid esters (verbascosides, cornoside, martynoside) present. Ellagic acid, proanthocyanidins, saponins, and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Ibicella (3; I. lutea, I. nelsoniana, I. parodii; Brazil, Bolivia, Argentina), Martynia (1; M. annua; Mexico, Central America, the Greater Antilles), Craniolaria (3; C. annua, C. argentina, C. integrifolia; the Greater Antilles, South America), Holoregmia (1; H. viscida; Bahia in northeastern Brazil), Proboscidea (5; P. althaeifolia, P. louisianica, P. parviflora, P. sabulosa, P. spicata; southern United States, Mexico, Peru).

Martyniaceae are probably sister to Schlegeliaceae. Gutierrez 2008 suggested a sister-group relationship with Verbenaceae.

The heavily glandular Ibicella and Proboscidea have been reported to capture insects and other arthropods (Schäferhoff & al. 2010). However, protease activity has not been detected in the glands (Plachno & al. 2009). Yet there may be mutualistic relationships between predatory insects and the plants in a way similar to that in Byblis (Byblidaceae) and Roridula (Roridulaceae) (Rice 2008). In other words, the leaves may absorb exudates/faeces from carnivorous insects feeding on the trapped animals.

Cladogram of Martyniaceae based on DNA sequence data (Gutierrez 2008).

Cladogram of Martyniaceae based on DNA sequence data (Gormley & al. 2015).

MAZACEAE Reveal

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Reveal in Kew Bull. 66: 47. Mar 2011

Genera/species 2/c 22

Distribution Tibet, East and Southeast Asia, Malesia to New Guinea, southeastern Australia, Tasmania, New Zealand.

Fossils Unknown.

Habit Bisexual, usually perennial or annual herbs, often with rhizome. Young stem quadrangular in cross-section.

Vegetative anatomy Roots fibrous. Phellogen absent? Pericyclic envelope absent in Mazus. Primary vascular tissue cylinder of bundles. Vessel elements with simple? perforation plates; lateral pits? Imperforate tracheary xylem elements ? with simple pits, septate? Wood rays absent? Axial parenchyma paratracheal? Sieve tube plastids S type. Nodes 1:1 (Mazus), unilacunar with one leaf trace. Crystals?

Trichomes Hairs multicellular, uniseriate; glandular hairs often present.

Leaves Usually opposite (sometimes spiral), simple, entire or lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Leaf margin serrate.

Inflorescence Terminal or axillary cyme or flowers solitary axillary.

Flowers Usually zygomorphic. Hypogyny. Sepals five, with valvate aestivation, persistent, connate into campanulate or bilabiate (3:2, three upper and two lower lobes) calyx. Petals (two to) four or five, connate into tubular or bilabiate (2:3, two upper and three lower lobes) corolla, sometimes with spur. Nectaries present. Disc absent.

Androecium Stamens usually two longer and two shorter (rarely two), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers usually with divergent (sometimes confluent) thecae, dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate?, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, often microreticulate, granulate or spinulate?

Gynoecium Pistil composed of two connate carpels (abaxial carpel often degenerated). Ovary superior, usually unilocular (pseudomonomerous or bilocular). Style single, simple. Stigma bilobate to fan-shaped, sensitive, without asymmetrically swollen stigmatoid tissue, type? Pistillodium absent.

Ovules Placentation usually subbasal or axile. Ovules one or several per carpel, orthotropous or hemianatropous?, epitropous?, unitegmic, tenuinucellar. Integument five or six cell layers thick. Hypostase present? Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus? Antipodal cells persistent? Endosperm development cellular. Endosperm haustorium chalazal. Embryogenesis solanad?

Fruit A berry-like more or less fleshy indehiscent fruit, enclosed by persistent and sometimes accrescent calyx.

Seeds Aril absent? Testa often thin. Exotesta? Endotesta? Perisperm not developed. Endosperm sparse or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = 19 (Mazus)

DNA Deletion in plastid gene matK?

Phytochemistry Very insufficiently known. Group I carbocyclic iridoids (aucubin, catalpol) present.

Use Ornamental plants.

Systematics Lancea (2; L. hirsuta, L. tibetica; Tibet, China), Mazus (c 20; East and Southeast Asia, Malesia to New Guinea, southeastern Queensland to southeastern South Australia, Tasmania, New Zealand).

Mazaceae are sister-group to a clade with the following topology: [Phrymaceae+[Paulowniaceae+[Rehmanniaceae+Orobanchaceae]]]. Other analyses have placed the two genera very close to Mimulus (Phrymaceae).

OLEACEAE Hoffmans. et Link

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Hoffmannsegg et Link, Fl. Portug. 1: 62. 1 Sep 1809 [’Oleinae’], nom. cons.

Jasminaceae Juss., Gen. Plant.: 104. 4 Aug 1789 [’Jasmineae’]; Lilacaceae Vent., Tabl. Règne Vég. 2: 307. 5 Mai 1799 [‘Lilaceae’], nom. illeg.; Fraxinaceae Vest, Anleit. Stud. Bot.: 269, 288. 1818 [’Fraxinoideae’]; Fraxinales Bercht. et J. Presl, Přir. Rostlin: 224. Jan-Apr 1820 [’Fraxineae’]; Jasminales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 248. Jan-Apr 1820 [‘Jasmineae’]; Oleales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 248. Jan-Apr 1820 [‘Oleaceae’]; Ligustraceae G. Mey., Chloris Han.: 245, 254. Jul-Aug 1836 [’Ligustrinae’]; Bolivariaceae Griseb., Gen. Sp. Gent.: 20. Oct 1838; Ligustrales Bartl. ex Bisch., Lehrb. Bot.: 3(2): 529. 1840 [‘Ligustrinae’]; Ligustropsida Bartl. ex Meisn., Plant. Vasc. Gen.: Comm.: 164. 5-11 Apr 1840 [’Ligustrinae’]; Forestieraceae (Endl.) Meisn., Plant. Vasc. Gen.: Tab. Diagn.: 345, Comm. 257. 13-15 Feb 1842 [’Forestiereae’]; Oleineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1031. 1846 [‘Oleineae’]; Syringaceae Horan., Char. Ess. Fam.: 115. 30 Jun 1847 [’Syringaceae nob (s. Oleaceae)’]; Nyctanthaceae J. Agardh, Theoria Syst. Plant.: 284. Apr-Sep 1858 [’Nyctantheae’]; Schreberaceae (Wight) Schnizlein, Iconogr. Fam. Regni Veg. 2: ad t. 151*. 1857-1870; Oleanae Takht., Divers. Classif. Fl. Pl.: 449. 24 Apr 1997

Genera/species 23/560–570

Distribution Cosmopolitan except polar areas, with their largest diversity in Southeast Asia and Australia.

Fossils Fossil wood of Oleaceae has been found in the Late Cretaceous of India. Winged seeds assigned to Oleaceae (possibly Fraxinus) have been found in Eocene and younger layers in North America, Europe and Asia. Fossil pollen of Oleaceae have been recovered from Late Cretaceous (Early Campanian) and Eocene layers in North America, but also from younger strata in Europe.

Habit Usually bisexual (rarely polygamomonoecious or dioecious), evergreen or deciduous trees, shrubs or lianas (rarely suffrutices).

Vegetative anatomy Phellogen ab initio usually superficial (rarely deeply seated). Medulla in Forsythia and Jasminum sometimes septated by diaphragms? Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids and fibre tracheids (sometimes libriform fibres) with simple and/or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually paratracheal scanty vasicentric, aliform, confluent, or banded (sometimes apotracheal diffuse), or absent. Sieve tube plastids S type. Nodes 1:1?, unilacunar with one? leaf trace. Stem and leaves with calciumoxalate as druses, sphaerites, styloids, raphides, or acicular to prismatic (often very small) crystals and/or crystal sand; crystals often frequent in epidermal cells of trichome bases.

Trichomes Hairs unicellular or multicellular, often lepidote or peltate; glandular hairs present, also peltate-lepidote; cells in head with exclusively vertical walls; groups of multicellular secretory hairs sometimes forming extrafloral nectaries.

Leaves Usually opposite (in some species of Jasminum alternate), simple entire or imparipinnate or trifoliolate (sometimes unifoliolate), with conduplicate ptyxis (‘Chionanthus’). Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation usually pinnate (rarely palmate). Stomata usually anomocytic (sometimes paracytic). Cuticle deeply furrowed. Cuticular wax crystalloids as rodlets or platelets. Domatia as pockets, pits or hair tufts, or absent. Mesophyll often with sclerenchymatous idioblasts with sclereids; calciumoxalte crystals frequent; nuclei of mesophyll parenchyma cells often with special crystalline enclosings. Leaf margin or leaflet margins serrate or entire. Hairs often peltate or as secretory glandular hairs – often as extrafloral nectaries – or sometimes sunken translucent points on abaxial side of lamina.

Inflorescence Terminal or axillary, usually thyrsoid, panicle, cymose or raceme (flowers sometimes solitary axillary).

Flowers Actinomorphic. Hypogyny. Sepals usually four (in Jasminum five to nine [to 15]), with usually valvate (sometimes open) aestivation, connate (rarely absent), orthogonally arranged/initiated. Petals usually four (in Jasminum five to nine [to twelve]), with imbricate, valvate, induplicate-valvate or convolute aestivation, usually more or less connate (rarely free or absent), diagonally arranged/initiated. Nectariferous disc intrastaminal, annular, or absent.

Androecium Stamens two (to five), antesepalous. Filaments free, usually adnate to corolla tube (epipetalous). Anthers basifixed or dorsifixed, with thecae arranged back-to-back, often with osmophores, usually non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Placentoid? Tapetum secretory. Female flowers sometimes with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colpate or (2–)3(–4)-colpor(oid)ate, shed as monads, usually bicellular (rarely tricellular) at dispersal. Exine semitectate, with columellate infratectum, reticulate, beset with rounded or elongate supratectal knobs (piloid excrescences), or psilate.

Gynoecium Pistil composed of two connate carpels, usually median (sometimes transverse or oblique). Ovary superior, bilocular (in Ligustrum nectar-secreting). Style single, simple, often short or absent. Stigma elongate-clavate, bilobate, papillate or non-papillate, Dry type. Male flowers sometimes with pistillodium.

Ovules Placentation axile, apical or basal. Ovules (one or) two (to four; in Forsythia numerous) per carpel, usually anatropous (rarely hemianatropous or amphitropous), pendulous or ascending, apotropous or epitropous, unitegmic, tenuinucellar. Integument usually approx. seven (rarely up to c. 20) cell layers thick, usually massive. Endothelium present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Endosperm development usually cellular (rarely nuclear). Endosperm haustoria absent. Embryogenesis caryophyllad or solanad.

Fruit A loculicidal capsule (in ’Menodora’ clade of Jasminum a pyxidium), a samara (Fraxinus), a berry, a drupe (in Dimetra and Nyctanthes a bipartite schizocarp with samaroid mericarps).

Seeds Aril absent. Testa often vascularized, sometimes winged. Exotesta often palisade, moderately and evenly thickened. Endotesta sometimes fibrous. Endothelium usually persistent. Perisperm not developed. Endosperm copious or sparse, oily, or absent. Embryo straight, well differentiated to little differentiated or rudimentary, without chlorophyll. Cotyledons two, flattened, sometimes nutrient-storing. Germination phanerocotylar or cryptocotylar.

Cytology n = 14 (Forsythieae); n = 13 (Fontanesia); n = 11, 12 (Myxopyreae; in Nyctanthes 11, 18, 22, 23); n = 11–13 (Jasminum); n = 23 (Oleeae) – Polyploidy occurring. Globular crystalline protein bodies present in nucleus.

DNA Deletion of 9 bp in plastid gene ndhF. Plastid genome in Jasminum with two inversions (i.a. one 21 kb inversion). Plastid gene clpP lost from at least two species of Jasminum; plastid gene accD (ORF512, zpfA) lost in some species. Mitochondrial coxI intron present in Jasminum.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, flavone glycosides (in Oleeae), Route I iridoids (carbocyclic iridoids and secoiridoids), Group IV carbocyclic iridoids (forsythiide, kingiside), Group VII secoiridoids (swertiamarin, gentiopicroside), Group VIII secoiridoids (oleuropein), Group IX secoiridoids (indole alkaloids of corynantheane type), Group X secoiridoids (loganin, ketologanin), triterpenes, ursolic acid and caffeic acid esters (cornosides in Forsythieae; oleoside in Jasminum and Oleeae; verbascosides, orobanchin, etc.), saponins, syringin, coumarins, coniferin, lignans, and polyols present. Ellagic acid, tannins, proanthocyanidins and cyanogenic compounds not found. Carbohydrates usually stored as oligosaccharides (mannitol). Cell walls with arabinoxyloglucans and often galactoxyloglucan hemicelluloses.

Use Ornamental plants, fruits and fruit oils (Olea), perfumes (Jasminum, ‘Osmanthus’), timber.

Systematics Oleaceae are sister to Carlemanniaceae.

The basal branching is unresolved. Jasminum is sister to Oleeae.

Forsythieae H. Taylor ex L. A. S. Johnson in Contr. N. S. W. Natl. Herb. 2: 397. 18 Nov 1957

2/10. Forsythia (9; western Balkan Peninsula, East Asia), Abeliophyllum (1; A. distichum; the Korean Peninsula). – The Balkan Peninsula, East Asia. Medulla septated. Petals with valvate aestivation. Ovules one or several per carpel. Fruit a samara or a capsule. n = 14. Cornosides present. Iridoids sometimes absent. – In some analyses, Forsythia and Abeliophyllum form a sister-group to the remaining Oleaceae.

Fontanesieae H. Taylor ex L. A. S. Johnson in Contr. N. S. W. Natl. Herb. 2: 397. 18 Nov 1957

1/2. Fontanesia (2; F. phillyreoides: Sicily, Turkey, Syria, Lebanon; F. fortunei: east central China). – Vestured pits present. Petals free, with imbricate aestivation. Ovule one per carpel. Fruit a samara. n = 13. Secologanoside, loganic acid, and 5-hydroxylated derivatives (e.g. swertiamarin) present.

Myxopyreae Boerl., Handl. Fl. Nederl. Ind. 2: 324. 1 Jan 1899

3/7. Myxopyrum (4; M. nervosum, M. ovatum, M. pierrei, M. smilacifolium; India, Burma, Hainan, Southeast Asia, the Andaman and Nicobar Islands, Malesia to New Guinea and the Bismarck Archipelago), Dimetra (1; D. craibeana; northeastern Thailand), Nyctanthes (2; N. aculeata, N. arbo-tristis; the Himalayas, Thailand, Java, Sumatra). – Tropical Asia. Cortical vascular bundles present in corners of angled stem in Myxopyrum and Nyctanthes. Sepals diagonally initiated (Nyctanthes). Petals in Nyctanthes with contorted aestivation. Gynoecium in Nyctanthes transversely orientated. Placentation basal or subbasal. Ovule one (to three) per carpel, ascending (Nyctanthes). Integument in Nyctanthes c. 20 cell layers thick. Megasporocytes occasionally several and megagametophyte disporous 8-nucleate (Allium type). Fruit a berry or a schizocarp. Exotesta in Nyctanthes poorly developed. Mesotesta in Nyctanthes persistent. Endotestal cells in Nyctanthes sclerotic, tangentially elongate. n = 11, 12 (18, 22, 23). Myxopyroside iridoid pathway present. – Myxopyreae are sister (with very high bootstrap support) to the remaining Oleaceae in analyses by Lee & al. (2007).

[Jasmineae+Oleeae]

Ovules two (to four) per carpel. Fruit drupaceous or baccate. Oleoside present.

Jasmineae Lam. et DC., Syn. Plant. Fl. Gall.: 216. 30 Jun 1806

1/c 200. Jasminum (c 200; warm-temperate to tropical regions in the Old World). – Sepals and petals up to 14 or more. First four sepals diagonally initiated. Petals with imbricate-quincuncial aestivation. Ovules usually two (rarely four) per carpel. Endothelium absent. Megasporocytes and megagametophytes occasionally several. Fruit bilobate, a berry or a pyxidium. Seed coat multilayered. Mesotestal cells with completely thickened or band-thickened anticlinal walls. x = 11–13. Inversion of 21 kb present in plastid genome. Secoiridoids present.

Oleeae Hoffmanns. et Link ex Dumort., Fl. Belg.: 52. 1827 [’Oleaceae’]

16/340–345. Ligustrum (c 60; Europe, the Mediterranean, North Africa, East and Southeast Asia, Malesia to eastern Australia and Tasmania, with their highest diversity in East Asia); Comoranthus (3; C. madagascariensis: western Madagascar; C. minor: southwestern Madagascar; C. obconicus: Mayotte Island), Schrebera (8; S. alata, S. arborea, S. trichoclada: tropical to southern Africa; S. capuronii, S. orientalis: Madagascar; S. swietenioides: India, the Himalayas, Southeast Asia; S. kusnotoi: Borneo; S. americana: Peru); Fraxinus (40–45; temperate regions on the Northern Hemisphere, few species in tropical regions); ‘Olea’ (32–33; warm-temperate to tropical regions in the Old World; polyphyletic), Haenianthus (3; H. incrassatus: Jamaica; H. salicifolius: the Greater Antilles; H. variifolius: Cuba),‘Chionanthus’ (c 60; warm-temperate to tropical regions in Africa, Madagascar, East Asia and eastern United States; polyphyletic), Notelaea (12; tropical and eastern Australia, Tasmania), Picconia (1; P. excelsa; Madeira, the Canary Islands), Noronhia (c 45; Madagascar, the Comoros), Phillyrea (2; P. angustifolia, P. latifolia; Madeira, the Mediterranean to Syria and northern Iran),‘Osmanthus’ (32; southwestern Asia, China to West Malesia, southeastern United States, Mexico; polyphyletic), Nestegis (5; N. apetala: New Zealand, Norfolk Island; N. cunninghamii, N. lanceolata, N. montana: New Zealand; N. sandwicensis: the Hawaiian Islands), Forestiera (c 20; southern United States, Mexico, Central America, the West Indies, Ecuador), Hesperelaea (1; H. palmeri; Guadalupe Island off Baja California; probably extinct), Priogymnanthus (2; P. apertus, P. hasslerianus; tropical South America). – Warm-temperate to tropical regions on both hemispheres. Vessels in multiples. Vestured pits sometimes present. Libriform fibres usually present. Fibre tracheids usually absent. Marginal parenchyma often present. Peltate scales sometimes present. Sepals sometimes diagonally initiated. Petals with usually valvate (sometimes imbricate) aestivation, sometimes free or absent. Stamens sometimes four. Ovules usually two (rarely four) per carpel. x = (20) 23. Flavone glycosides and secoiridoids present. – Ligustrum (including Syringa) may be sister to the remaining Oleeae.

Cladogram of Oleaceae based on DNA sequence data (Wallander & Albert 2000).

OROBANCHACEAE Vent.

( Back to Plantaginales )

Ventenat, Tabl. Règne Vég. 2: 292. 5 Mai 1799 [’Orobanchoideae’], nom. cons.

Pedicularidaceae Juss., Gen. Plant.: 99. 4 Aug 1789 [’Pediculares’]; Rhinanthaceae Vent., Tabl. Règne Vég. 2: 295. 5 Mai 1799 [’Rhinanthoideae’]; Euphrasiaceae Martinov, Tekhno-Bot. Slovar: 239. 3 Aug 1820 [‘Euphrasiae’]; Orobanchales Vent. ex Bercht. et J. Presl, Přir. Rostlin: 242. Jan-Apr 1820 [‘Orobanchoideae’]; Melampyraceae Rich. ex Hook. et Lindl. in W. J. Hooker, Fl. Scot. 2: 213. 11 Mar 1821; Rhinanthales Dumort., Anal. Fam. Plant.: 20. 1829 [’Rhinantharieae’]; Orobanchineae Link, Handbuch 1: 506. 4-11 Jul 1829 [‘Orobanchinae’]; Rhinanthineae Link, Handbuch 1: 513. 4-11 Jul 1829 [‘Rhinanthaceae’]; Phelypaeaceae Horan., Prim. Lin. Syst. Nat.: 73. 2 Nov 1834 [’Phelipaeaceae’]; Buchneraceae (Benth.) Lilja, Skånes Fl., ed. 2: 979. Apr-Dec 1870; Aeginetiaceae Livera in Ann. Roy. Bot. Gard. (Peradeniya) 10: 153. 31 Mai 1927; Cyclocheilaceae Marais in Kew Bull. 35: 805. 28 Apr 1981; Nesogenaceae Marais in Kew Bull. 35: 798. 28 Apr 1981; Lindenbergiaceae Doweld, Tent. Syst. Plant. Vasc.: xlix. 23 Dec 2001

Genera/species 98/1.850–2.150

Distribution Mainly temperate and alpine regions, with their highest diversity in temperate regions on the Northern Hemisphere and in Africa and Madagascar; some representatives in southern South America, South Asia, southern Australia, and New Zealand.

Fossils Unknown.

Habit Bisexual, usually perennial, biennial or annual herbs (rarely shrubs or suffrutices, e.g. Brandisia, Cyclocheilon, Asepalum). Nearly all species are either root hemiparasites having green assimilating leaves, or more or less succulent achlorophyllous root holoparasites with scale-like leaves. Often blackening when dry. Roots in Asepalum and Cyclocheilon at least usually red. Lindenbergia comprises autotrophic, photosynthesizing plants.

Vegetative anatomy Mycorrhiza absent. Roots with haustoria. Phellogen ab initio superficial? Medullary vascular bundles present or absent. Primary vascular tissue one or several cylinders of bundles. Secondary lateral growth normal or absent. Endodermis sometimes prominent (Euphrasia). Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements libriform fibres? with simple pits, septate? Wood rays ?-seriate, usually heterocellular? Axial parenchyma paratracheal. Sieve tube plastids S type. Nodes usually 1:1?, unilacunar with one? leaf trace. Crystals?

Trichomes Hairs unicellular or multicellular, usually uniseriate (sometimes branched; in Brandisia also stellate); glandular hairs sometimes dendritic or lepidote; head of glandular hairs without vertical septa (not vertically divided).

Leaves Alternate (spiral) or opposite, simple, entire or pinnately lobed in hemiparasites, scale-like and membranous or absent in holoparasites, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually anomocytic?, not closing (not even in Lindenbergia). Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts. Leaf margin lobate, serrate or entire. Extrafloral nectaries present as glandular hairs on lamina in some species of Melampyrum.

Inflorescence Terminal or axillary, usually raceme (in Lindenbergia branched raceme) or spike (flowers occasionally solitary axillary). Floral prophylls (bracteoles) lateral (in Asepalum and Cyclocheilon large, free or connate, enclosing bud, persistent and accrescent in fruit), at pedicel base, shortly above foliaceous bract, or immediately below flower.

Flowers Zygomorphic. Hypogyny. Sepals four or five, with valvate or open aestivation, persistent, connate (in Asepalum and Cyclocheilon almost absent); median sepal adaxial? Petals five, with imbricate quincuncial or descending-cochlear aestivation (adaxial-lateral, posterior, corolla lobes usually covered in bud by one or two abaxial-lateral lobes; sometimes also abaxial lobe covering adaxial-lateral lobes; often ascending-cochlear aestivation in which abaxial lobe covers all other lobes; abaxial lobe often inserted outside of adaxial lobes), caducous or persistent (collar-like base of corolla tube persistent), connate into tubular or infundibuliform (often obliquely) usually bilabiate corolla. Nectariferous disc intrastaminal, annular, often fleshy (nectaries and/or disc sometimes absent).

Androecium Stamens usually two longer and two shorter (didynamous), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, usually adnate to corolla tube (epipetalous). Anthers often connivent, with thecae parallel or confluent, often with one theca reduced, often hairy, sagittate to invertedly U-shaped (sometimes separate), dorsifixed, versatile?, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; in Bartsia and some species of Euphrasia poricidal, dehiscing by apical pores); connective sometimes prolonged at apex. Tapetum usually secretory (sometime amoeboid-periplasmodial). Staminodia usually absent (rarely one, adaxial-median).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–4)-colpate or (2–)3(–4)-porate (rarely inaperturate; in, e.g., Lindenbergia tricolporate; in Pedicularis often syncolpate), shed as monads, bicellular at dispersal. Exine usually tectate (rarely semitectate), with columellate infratectum, usually perforate, microreticulate or finely rugulate, often with verrucoid or globular supratectal elements (retipilate; rarely reticulate).

Gynoecium Pistil composed of two (or three or five) connate carpels. Ovary superior, entirely or partially unilocular or bilocular. Style single, simple. Stigma clavate (e.g. Brandisia) to capitate or bilobate to quadrilobate (rarely lingulate or spatulate), papillate, Dry type. Pistillodium absent.

Ovules Placentation axile (when ovary bilocular) or parietal (when ovary unilocular; placentae sometimes two bilobate, four or six). Ovules usually numerous (rarely, e.g., in Nesogenes one) per carpel, usually anatropous, usually epitropous (rarely apotropous), unitegmic, tenuinucellar. Integument (two to) four to twelve cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells often persistent. Endosperm development cellular. Endosperm haustoria chalazal and often micropylar. Embryogenesis onagrad.

Fruit Usually a loculicidal or septicidal capsule (in Radamaea a berry; in Leucosalpa and Tozzia a drupe; in Nesogenes a two-seeded drupe with persistent calyx; in Asepalum a schizocarp with two mericarps).

Seeds Aril absent. Seed pedestals often present. Testa often winged; radial cell walls of wings with reticulate thickenings. Inner walls of exotestal cells variously thickened; cell walls of remaining layers thickened and lignified. Endotestal cell walls sometimes thickened and lignified. Perisperm not developed. Endosperm usually copious (sometimes sparse; absent in Monttea, Asepalum and Cyclocheilon), usually oily (often starchy). Embryo small to minute, often undifferentiated when mature (globular or ovoid, lacking histogens except dermatogen), usually straight, without chlorophyll. Cotyledons two (often reduced). Germination phanerocotylar or cryptocotylar, often through germination tube.

Cytology n = (6) 7 – 21 (or more?) – Polyploidy occurring; endomitotic polyploidization in inner tapetal cells (not in Pedicularis and Melampyrum); genome size decreasing after polyploidization. Protein bodies in cell nucleus lamellar.

DNA Deletion in plastid gene matK. Larger part of small single copy region of plastid DNA deleted in Epifegus. Plastid inverted repeat absent from Conopholis and one inversion present in its cpDNA. Plastid genome in Striga asiatica with three inversions. Numerous plastid genes (e.g. rps16 and infA) and introns lost in Epifegus and Conopholis.

Phytochemistry Group I carbocyclic iridoids (aucubin, catalpol, daphylloside), Group X secoiridoids (nepeta lactones, iridoidpyridine alkaloids), pyrrolizidine alkaloids as macrocyclic diesters (at least in Castilleja and Melampyrum), caffeic acid esters (bartsioside, orobanchin, verbascosides), and silicic acid present. Flavonols, 6- and/or 8-hydroxylated flavone glycosides, ellagic acid, tannins, proanthocyanidins, saponins, and cyanogenic compounds not found. Carbohydrates stored as hexites and other oligosaccharides (e.g. mannitol in some species). Many substances are transported from host to parasite, not manufactured by the parasite.

Use Ornamental plants?

Systematics Orobanchaceae are sister-group to Rehmanniaceae.

Lindenbergieae T. Yamaz. in Fl. Cambodge, Laos et Vietnam 21: 20. 25 Jan 1985

1/12. Lindenbergia (12; northeastern Africa, tropical Asia to the Philippines, with their highest diversity in India). – Bracts foliaceous. Floral prophylls (bracteoles) usually absent. Anther thecae inserted on connective arms. Pollen grains tricolporate. Testal surface usually with hook-shaped thickenings. n = 16.

Orobanchoideae Eaton, Bot. Dict., ed. 4: 32. Apr-Mai 1836 [‘Orobancheae’]

97/1.820–2.100. Cymbarieae D. Don in Edinburgh New Philos. j. 19: 112. Jul 1835. Schwalbea (1; S. americana; eastern United States), Siphonostegia (3; S. laeta: eastern Mediterranean; S. syriaca: eastern Greece, southern Turkey; S. chinensis: East Asia), Monochasma (2; M. savatieri, M. sheareri; East Asia), Bungea (2; B. trifida: southwestern Asia; B. vesiculifera: Tien Shan), Cymbaria (5; C. borysthenica, C. chaneti, C. daurica, C. linearifolia, C. mongolica; Ukraine and southern Russia to Central and East Asia). – Brandisia (11; Burma, China). – Orobancheae Lam. et DC., Syn. Pl. Fl. Gall.: 214. 30 Jun 1806. Kopsiopsis (2; K. hookeri, K. strobilacea; western North America), Xylanche (1; X. himalaica; Himalayas, China to Taiwan), Boschniakia (1; B. rossica; temperate Asia to Japan, northwestern North America), Conopholis (3; C. alpina, C. americana, C. panamensis; southeastern United States, Mexico to Panamá; in Boschniakia?), Epifagus (1; E. virginiana; North America; in Boschniakia?); ‘Orobanche’ (c 100; temperate and subtropical regions of both hemispheres; paraphyletic), Boulardia (1; B. latisquama; Spain, the Baleares), Diphelypaea (2–4; D. boissieri, D. coccinea, D. helenae, D. tournefortii; southeastern Europe, Turkey, the Caucasus), Phelipanche (c 50; Europe, the Mediterranean, Macaronesia, western and southwestern Asia), Aphyllon (21; southern Canada to Peru), Mannagettaea (2; M. hummelii, M. labiata; eastern Siberia to western China), Cistanche (16; the Mediterranean, Ethiopia to western India and northwestern China). – Rhinantheae Lam. et DC., Syn. Pl. Fl. Gall.: 208. 30 Jun 1806. Pterygiella (5; P. bartschioides, P. cylindrica, P. duclouxii, P. nigrescens, P. suffruticosa; southern China); Melampyrum (c 35; temperate regions on the Northern Hemisphere), Rhinanthus (c 45?; temperate regions on the Northern Hemisphere), Lathraea (4; L. clandestina, L. japonica, L. rhodopaea, L. squamaria; Europe, temperate Asia), Bartsia (1; B. alpina; cold-temperate and alpine regions in Europe and North America), Rhynchocorys (8; southern Balkan Peninsula, central Mediterranean to Iran), Euphrasia (170–350; temperate regions on the Northern Hemisphere, alpine regions in New Guinea, eastern Australia and New Zealand), Tozzia (1; T. alpina; the Alps, the Carpathians), Odontitella (1; O. virgata; the Iberian Peninsula), Nothobartsia (2; N. aspera, N. spicata; western Mediterranean), Hedbergia (1; H. abyssinica; mountains in tropical Africa), ‘Bartsia’ (c 60; Europe, the Mediterranean, mountains in tropical Africa, temperate Asia, North America to southern South America, with their largest diversity in the Andes; paraphyletic), Parentucellia (3; P. floribunda, P. latifolia, P. viscosa; the Mediterranean to southwestern Asia), Macrosyringion (2; M. glutinosum, M. longiflorum; the Mediterranean), Odontites (c 30; Europe, the Mediterranean to the Himalayas). – Buchnereae Benth. in Edwards’s Bot. Reg. 21: ad t. 1770. 1 Feb. 1835. Cyclocheilon (3; C. kelleri, C. physocalyx, C. somaliense; Somalia, Ethiopia, the Arabian Peninsula), Asepalum (1; A. eriantherum; Ethiopia, Kenya, Tanzania, Yemen), Sopubia (41; tropical and southern Africa, the Himalayas to Indochina and Taiwan), Graderia (4; G. fruticosa, G. linearifolia, G. scabra, G. subintegra; Africa, Socotra), Nesogenes (8; Tanzania, Madagascar, islands in the Indian and Pacific oceans), Radamaea (5; R. latifolia, R. montana, R. perrieri, R. prostrata, R. rupestris; Madagascar), Bardotia (1; B. ankaranensis; northern Madagascar), Rhamphicarpa (5; R. aquatica, R. brevipedicellata, R. capillacea, R. fistulosa, R. medwedewii; Russia, tropical and southern Africa, Turkey, India, tropical Australia), Sieversandreas (1; S. madagascarianus; southern Madagascar), Xylocalyx (5; X. aculeolatus, X. asper, X. carterae, X. hispidus, X. recurvus; Somalia, Socotra), Striga (40–45; tropical, subtropical and southern Africa, southern Asia, northern Australia), Cycnium (18; tropical East Africa to South Africa), Buchnera (c 100; tropical and subtropical regions on both hemispheres), Centranthera (c 13; China and south to northern and northeastern Australia), Alectra (c 40; tropical and southern Africa, tropical Asia), Melasma (25–35; southern Africa, tropical and subtropical America), Escobedia (c 10; Mexico, Central America, tropical South America), Hyobanche (9; southern Africa), Harveya (c 30; tropical and southern Africa, the Mascarene Islands), Aeginetia (8; India, Sri Lanka and Burma to China, Japan and Southeast Asia, Malesia), Christisonia (16–20; southwestern China, Southeast Asia, Malesia). – Pedicularideae Duby, Bot. Gall. 1: 351. 12-14 Apr 1828. Pedicularis (500–600; temperate and arctic-alpine regions on the Northern Hemisphere and south to the Andes in Colombia, with their largest diversity in Central Asia to western China), Phtheirospermum (5; P. glandulosum, P. japonicum, P. muliense, P. parishii, P. tenuisectum; East Asia), Lamourouxia (28; Mexico, Central America, tropical South America to Peru), Esterhazya (7; E. andina, E. caesarea, E. eitenorum, E. macrodonta, E. nanuzae, E. splendida, E. triflora; Brazil, Bolivia), Agalinis (c 70; United States, Mexico, Central America, the West Indies, tropical South America), Aureolaria (8; eastern United States, Mexico), Seymeria (c 25; southern United States, Mexico), Cordylanthus (19; western North America), Orthocarpus (8; southwestern Canada, western United States, northwestern Mexico), Castilleja (>200; temperate regions on the Northern Hemisphere to Central America and the Andes in northern South America), Triphysaria (6; T. chinensis, T. eriantha, T. floribunda, T. micrantha, T. pusilla, T. versicolor; British Columbia to California). – Unplaced Orobanchoideae Baumia (1; B. angolensis; Angola), Brachystigma (1; B. wrightii; Arizona, New Mexico, northwestern Mexico), Buttonia (3; B. hildebrandtii, B. natalensis, B. superba; tropical and southern Africa), Chloropyron (4; C. maritimum, C. molle, C. palmatum, C. tecopense; western United States, northwestern Mexico), Dasistoma (1; D. macrophylla; southeastern United States), Eremitilla (1; E. mexicana; Mexico), Gerardiina (2; G. angolensis, G. kundelungensis; tropical and southern Africa), Ghikaea (1; G. speciosa; northeastern Africa), Gleadovia (3; G. banerjiana, G. mupinensis, G. ruborum; western Himalayas, western China), Hiernia (1; H. angolensis; Angola, Namibia), Leptorhabdos (1; L. parviflora; the Caucasus, Iran to Central Asia and the Himalayas), Leucosalpa (3; L. grandiflora, L. madagascariensis, L. poissonii; Madagascar), Macranthera (2; M. flammea, M. fuchsioides; southeastern United States), Magdalenaea (1; M. limae; southeastern Brazil), Micrargeria (3; M. barteri, M. filiformis, M. wightii; tropical Africa, India), Micrargeriella (1; M. aphylla; Congo, Zambia), Nothochilus (1; N. coccineus; Brazil), Omphalotrix (1; O. longipes; northeastern Asia), Paraharveya (1; P. alba; Central and tropical East Africa), Parasopubia (3; P. bonatii, P. delphiniifolia, P. hofmannii; Southeast Asia), Petitmenginia (2; P. comosa, P. matsumurae; southern China, Southeast Asia), Phacellanthus (1; P. tubiflorus; China, the Korean Peninsula, Japan, the Russian Far East), Physocalyx (3; P. aurantiacus, P. major, P. scaberrimus; Brazil), Pseudomelasma (1; P. peduncularioides; Madagascar), Pseudosopubia (7; tropical Africa), Pseudostriga (1; P. cambodiana; Southeast Asia), Rhaphispermum (1; R. gerardioides; central Madagascar), Seymeriopsis (1; S. bissei; Cuba), Silviella (2; S. prostrata, S. serpyllifolia; Mexico), Tetraspidium (1; T. laxiflorum; Madagascar), Thunbergianthus (2; T. quintasii: São Tomé; T. ruwenzoriensis: Congo, tropical East Africa), Vellosiella (2; V. dracocephaloides, V. spathacea; Brazil). – Subcosmopolitan. Root hemi- or holoparasitic herbs (sometimes shrubs). Stomata not closing. Sepals sometimes free or almost free. Corolla tube development sometimes intermediate. Filaments in Eremitilla free from petals. Stamens in Castillejinae unithecal or unequally bithecal. Anthers often hairy and with tails or basal awns. Pollen grains often with starch. Cells of testal wings sometimes with reticulate thickenings on radial walls. Endosperm sometimes with thickened cell walls. Polysaccharides sometimes mannose-rich. – Schwalbea is probably basal in Orobanchoideae. It forms a clade – together with Bungea, Cymbaria, Monochasma and Siphonostegia – possibly being sister-group to the remainder. Cyclocheilaceae form an ingroup within Orobanchoideae. Nesogenes is sometimes identified as sister, with large support, to Radamaea.

The positions of Orobancheae and Rhinantheae relative to Buchnereae and Pedicularideae are ambiguous. Rhinantheae have been basal to Orobancheae in some earlier analyses, and Buchnereae are sometimes basal to both Rhinantheae and Pedicularideae.

Holoparasitism has evolved several times from hemiparasitism.

The position of Brandisia is uncertain. According to, Bennett & Mathews 2006 it may be sister to the core clade of hemiparasitic Orobanchoideae.Brandisia consists of shrubs, sometimes parasitic, with usually stellate hairs. The leaves are opposite, with entire or serrate margin. The flowers are axillary, solitary or pairwise, or arranged in racemes, and have two floral prophylls (bracteoles). The calyx is campanulate, two-lipped or almost actinomorphic, with two or five lobes. The corolla has a slightly curved tube and a two-lipped limb with bilobate upper lip and trilobate lower lip.The four stamens are didynamous, with filaments adnate to base of corolla tube. Staminodia are absent. The superior ovary is bilocular, the style is simple and the stigma capitate. The ovules are numerous per carpel. The fruit is a loculicidal capsule. The seed coat has reticulate pattern (cf. Paulownia in Paulowniaceae) and the testa is provided with membranous wings (cf. Paulowniaceae). Phenylethanoids are present.

Phylogeny (simplified) of Orobanchaceae based on DNA sequence data (Wolfe & al. 2005; Morawetz & al. 2010; etc.).

Phylogeny (simplified) of Orobanchaceae based on DNA sequence data (McNeal & al. 2013).

PAULOWNIACEAE Nakai

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Nakai in J. Jap. Bot. 24: 13. 10 Dec 1949

Genera/species 1–2/7–18

Distribution Warm-temperate, subtropical and tropical regions in continental China to northern Indochina, the Korean Peninsula, Japan, Taiwan.

Fossils Uncertain. Fossils assigned to Paulownia columbiana have been described from the Miocene/Pliocene transition zone of Germany.

Habit Bisexual, deciduous trees.

Vegetative anatomy Phellogen ab initio outer-cortical. Medulla septated by diaphragms. Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits? Imperforate tracheary xylem elements ? with simple or bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular. Axial parenchyma aliform, lozenge-aliform, winged-aliform, confluent or banded. Tyloses abundant. Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace (Paulownia). Crystals? C4 physiology present in Paulownia.

Trichomes Hairs multicellular, uniseriate or branched, or absent.

Leaves Opposite, simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular. Venation pinnate. Stomata anomocytic? Cuticular wax crystalloids? Leaf margin entire. Extrafloral nectaries often present on lamina.

Inflorescence Terminal.

Flowers Zygomorphic, large. Hypogyny. Sepals five, with valvate aestivation, persistent, more or less connate (in Paulownia at base), campanulate, in Paulownia covered by long brown hairs; median sepal adaxial. Petals five, with ascending-cochlear aestivation, deciduous, connate inte bilabiate or hypocrateriform corolla (in Paulownia covered by simple hairs with tapering terminal cell). Nectaries vascularized. Disc intrastaminal.

Androecium Stamens two longer and two shorter (in terminal flowers often five), haptostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers with separate or confluent thecae (in Paulownia positioned head-to-head), basifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits), in Paulownia with massive endothecium surmounting connective. Tapetum secretory? Placentoid present (Paulownia). Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains (2–)3(–4)-colporate (or –colpate), shed as monads, bicellular? at dispersal. Exine semitectate, with columellate? infratectum, foveolate-reticulate to microreticulate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular. Style singe, simple, hollow, sometimes widened at apex. Stigma punctate, hollow, papillate?, type? Pistillodium absent.

Ovules Placentation axile, protruding. Ovules numerous per carpel, anatropous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular? Endothelial cells thickened laterally and at endosperm. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal (Paulownia) or loculicidal-septicidal capsule with persistent woody calyx tube.

Seeds Aril absent? Seed pedestals present. Testa winged, with several membranous sinuous wings. Exotestal cells wide, with complex reticulate wall thickenings. Endotesta? Perisperm not developed. Endosperm smooth, uni- or multilayered, oily? Embryo straight?, well differentiated?, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = (19) 20 (Paulownia)

DNA Mitochondrial coxI intron present.

Phytochemistry Insufficiently known. Flavones (luteolin), Group I iridoids (catalpol), iridoid glycosides (mussaenoside), sitosterolglycosides (daucosterol), sitosterols (gentiobioside), and verbascosides present.

Use Ornamental plants, medicinal plants, timber.

Systematics Paulownia (6–17; warm-temperate, subtropical and tropical regions in continental China to northern Indochina, the Korean Peninsula, Japan, Taiwan; incl. Shiuyinghua?), Shiuyinghua (1; S. silvestrii; central China; in Paulownia?).

Paulowniaceae are sister-group to the clade [Rehmanniaceae+Orobanchaceae]. However, there is medium support for a sister-group relationship between Paulownia and Lamiaceae (Olmstead & al. 2000), and a somewhat higher support for Paulownia being sister to Orobanchaceae.

PEDALIACEAE R. Br.

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Brown, Prodr. Fl. Nov.-Holl.: 519. 27 Mar 1810 [’Pedalinae’], nom. cons.

Pedaliales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 246. Jan-Apr 1820 [‘Pedalinae’]; Sesamales Bercht. et J. Presl, Přir. Rostlin: 246. Jan-Apr 1820 [‘Sesamoideae’]; Sesamaceae Horan, Prim. Lin. Syst. Nat.: 74. 2 Nov 1834

Genera/species 14/77–80

Distribution Tropical and southern Africa, Madagascar, southwestern Asia, Malesia to New Guinea, northern and arid Australia.

Fossils Unknown.

Habit Bisexual, usually perennial or annual herbs (sometimes deciduous trees, shrubs or suffrutices; in Sesamothamnus and Uncarina with succulent stems; several genera with water-storing tuberous root; in Pterodiscus with caudex consisting of succulent sub- and supraterranean stem and root segments). Many species are xerophytic. Often with strong smell.

Vegetative anatomy Phellogen ab initio superficial. Cortical cells in some succulents with chloroplasts. Medulla in Pedalium sometimes septated by diaphragms? Cambium storied. Wood elements usually non-storied. Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements usually libriform fibres (in Uncarina fibre tracheids) with simple pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal banded or usually paratracheal scanty vasicentric. Fibres sometimes few. Sieve tube plastids Ss type. Nodes? Pericycle also with sclereids; medulla and cortex in Sesamothamnus with sclereids.Wood ray cells sometimes with single rhomboidal calciumoxalate crystals.

Trichomes Hairs eglandular uniseriate (rarely branched), or glandular capitate-peltate/stellate mucilage hairs consisting of unicellular or multicellular uniseriate stalk and usually peltate (sometimes stellate) and quadricellular apical head; outer cell walls of glandular head extremely thickened; cell walls of mucilage glands when moistened dissolved and modified into mucilage.

Leaves Usually opposite (sometimes, e.g. in Sesamum alternate, spiral), simple, entire or pinnately lobed (sometimes palmately compound), with ? ptyxis. Stipules and leaf sheath absent. Petiole sometimes modified into spines. Petiole vascular bundle transection disrupted annular. Venation usually pinnate (sometimes palmate). Stomata usually anomocytic or anisocytic (rarely diacytic or paracytic) or absent. Cuticular wax crystalloids? Leaf margin serrate, lobate or entire; lobe apex in Uncarina with hydathode.

Inflorescence Flowers usually axillary solitary (probably corresponding to reduced cymes; sometimes in few-flowered axillary dichasial inflorescences; in Sesamothamnus raceme-like). In axils of floral prophylls (bracteoles) two (to five) aborted lateral floral buds modified into axillary vascularized nectariferous glands (not in Uncarina).

Flowers Zygomorphic, often large. Extrafloral nectaries often present on pedicel. Hypogyny. Sepals five, with imbricate aestivation, unequal in size, persistent, more or less connate; median sepal adaxial. Petals five, with imbricate aestivation, bilabiate (with two upper and three lower lobes) or somewhat non-uniformly quinquelobate connate into tubular or infundibuliform corolla (rarely with adaxial spur at base). Nectariferous disc intrastaminal?, annular, often asymmetrical.

Androecium Stamens usually four, two longer and two shorter (didynamous; often with staminodial adaxial additional stamen), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed or basifixed, often pairwise connivent, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); thecae usually confluent, inserted at right angles to filaments; connective usually with apical gland. Tapetum secretory, with uninucleate to quadrinucleate cells. Staminodium one or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (3–)5–13(–15)-stephanocolpate, usually shed as monads (in Sesamothamnus as tetrads), tricellular (bicellular?) at dispersal. Exine intectate?, with columellate? infratectum, pilate.

Gynoecium Pistil composed of usually two (in Josephinia four) connate carpels (rarely distinctly unequal in size). Ovary superior, usually bilocular (in Josephinia octalocular); locules often entirely or partially bipartite by secondary septa. Style single, simple, filiform. Stigma usually bilobate, often sensitive, papillate, Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules two to numerous per carpel (in Josephinia one ovule per locule), anatropous, pendulous, horizontal or ascending, unitegmic, tenuinucellar. Integument seven to 20 cell layers thick. Archespore unicellular or bicellular. Hypostase present. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis onagrad.

Fruit Usually a loculicidal (in Uncarina loculicidal-septicidal) capsule (sometimes a nut or schizocarp) with persistent calyx and hardening (indurating) stylar base, and with beak-shaped outgrowths from paracarp sterile upper part of ovary. Exocarp and mesocarp decaying during maturation. Endocarp sclerenchymatous, with bristles, sharp edges, prickles, hooks, wings or horn-shaped fibrous processes.

Seeds Aril? Testa multiplicative, often with wings. Outer epidermis tanniniferous. Exotestal cells palisade with lignified walls or with annular wall thickenings. Exotesta and mesotesta sometimes with stellate calciumoxalate crystals. Endotesta? Perisperm not developed. Endosperm very thin, oily, or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, often rich in lipids and amyloid (xyloglucans). Germination phanerocotylar.

Cytology n = 8, 13, 16, 26, 32 – Polyploidy frequently occurring. Protein bodies present in cell nucleus?

DNA Mitochondrial coxI intron present. Deletion in plastid gene matK?

Phytochemistry Flavone-C-glycosides, Route II iridoids (also 10-hydroxylated carboxylic iridoids), Group II carbocyclic iridoids (harpagide, harpagioside, procumbide), iridoid glycosides, and verbascosides (i.a. orobanchin) present. Ellagic acid, proanthocyanidins, saponins, and cyanogenic compounds not found. Seed oils in, e.g., Sesamum containing up to 2,5% lignans (e.g. sesamin and sesamolin).

Use Ornamental plants, edible seeds, seed oils (Sesamum), medicinal plants (Harpagophytum).

Systematics Pedalieae Dumort., Anal. Fam. Plant.: 22. 1829. Uncarina (14; Madagascar), Rogeria (2; R. adenophylla, R. longiflora; tropical and southwestern Africa), Holubia (1; H. saccata; northeastern South Africa, Botswana, Zimbabwe), Harpagophytum (2; H. procumbens, H. zeyheri; Namibia, Botswana, northern South Africa, Madagascar), Pterodiscus (14; tropical and southern Africa), Pedaliodiscus (1; P. macrocarpus; Kenya, Tanzania), Pedalium (1; P. murex; northeastern Africa, probably introduced and naturalized in tropical and subtropical regions in the Old World). – Sesamothamneae Ihlenf. in Mitt. Staatsinst. Allg. Bot. Hamburg 12: 75. 1967. Sesamothamnus (6; S. benguellensis, S. busseanus, S. guerichii, S. leistneranus, S. lugardii, S. rivae; northeastern tropical Africa to southwestern and southeastern subtropical Africa). – Sesameae Dumort., Anal. Fam. Plant.: 22. 1829. ‘Sesamum’ (c 20; tropical and southern Africa, the Mascarene Islands, the Arabian Peninsula, southwestern Asia to India and Sri Lanka; non-monophyletic), Ceratotheca (5; C. integribracteata, C. reniformis, C. saxicola, C. sesamoides, C. triloba; tropical and Southern Africa), Dicerocaryum (4; D. eriocarpum, D. forbesii, D. senecioides, D. zanguebarium; eastern and southern Africa, Madagascar), Josephinia (3–6; J. africana, J. eugeniae, J. imperatricis; Kenya, Somalia, Malesia, tropical and arid regions in Australia). – Unplaced Pedaliaceae Linariopsis (2; L. chevalieri, L. prostrata; western and southwestern tropical Africa), Dewinteria (1; D. petrophila; Namibia).

Pedaliaceae may be sister to Acanthaceae, although their sister-group relationship is unresolved. According to Gormley & al. (2015), Pedaliaceae are possibly sister-group to Bignoniaceae.

Maximum likelihood tree of Pedaliaceae based on DNA sequence data (Gormley & al. 2015).

PHRYMACEAE Schauer

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Schauer in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 11: 520. 25 Nov 1847, nom. cons.

Wightiaceae Bo Liu, Bing Liu, Su Liu & Y. H. Tan, J. Syst. Evol. 58: 12. https://doiorg/10.1111/jse.12513 17 May 2019.

Genera/species 2–15/c 135

Distribution Tropical and southern Africa, Madagascar, East Asia to Manchuria and Japan, South and Southeast Asia, Malesia, Australia, Tasmania, New Zealand, America.

Fossils Unknown.

Habit Bisexual, usually perennial or annual herbs, often with rhizome (rarely lignified in lower part). Wightia also contains evergreen trees, shrubs and lianas. Young stem quadrangular in cross-section. Some species are aquatic.

Vegetative anatomy Phellogen? Primary vascular tissue a cylinder of bundles (one bundle at each edge of young stem). Vessel elements with simple? perforation plates; lateral pits? Imperforate tracheary xylem elements ? with simple pits, septate? Wood rays absent? Axial parenchyma paratracheal? Sieve tube plastids S type. Nodes 1:3, unilacunar with three leaf traces (Phryma). Crystals?

Trichomes Hairs multicellular, uniseriate (in Wightia branched stellate); glandular hairs often present.

Leaves Alternate (spiral) or opposite, simple, entire or lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually anomocytic (rarely anisocytic or diacytic). Cuticular wax crystalloids? Leaf margin usually serrate (in Wightia entire).

Inflorescence Terminal or axillary, cymose, raceme or spike (rarely capitate or flowers solitary axillary).

Flowers Usually zygomorphic (rarely almost actinomorphic). Hypogyny. Sepals (three to) five, with valvate aestivation (Mimulus), persistent, connate into tubular or bilabiate (3:2, three upper and two lower lobes) calyx. Petals (two to) four or five, connate into tubular or bilabiate (2:3, two upper and three lower lobes; Mimulus douglasii with two upper lobes only) corolla, sometimes with spur. Nectaries present or absent. Disc absent.

Androecium Stamens usually two longer and two shorter (rarely two), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers often with confluent (in Mimulus sometimes connate) thecae, dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate or <10-colpodiorate (each colpus with two ora; rarely spiraperturate, penta- to heptastephanocolpate etc.), shed as monads, bicellular or tricellular at dispersal. Exine tectate, not thickest close to apertures, with columellate infratectum, often microreticulate, granulate or spinulate.

Gynoecium Pistil composed of two connate carpels (abaxial carpel often reduced and sterile). Ovary superior, usually unilocular (pseudomonomerous; sometimes bilocular). Style single, simple. Stigma usually broadly bilobate (sometimes capitate or faintly bilobate), without asymmetrically swollen stigmatoid tissue, in Mimulus sensitive, type? Pistillodium absent.

Ovules Placentation usually axile (sometimes subbasal, rarely parietal). Ovules one or several per carpel, orthotropous or hemitropous, ascending, apotropous or epitropous, unitegmic, tenuinucellar. Integument three to seven cell layers thick. Hypostase present. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus (Phryma). Antipodal cells persistent (Phryma), in Mimulus two, one of which binucleate. Endosperm development cellular. Endosperm haustorium chalazal. Embryogenesis solanad.

Fruit A usually loculicidal (rarely septicidal) capsule or a one-seeded nut, enclosed by persistent and sometimes accrescent calyx (dorsal calyx teeth often modified into prickles or hooks; rarely a fleshy fruit or a schizocarp).

Seeds Aril absent? Seed pedestals present. Exotesta? Endotesta? Perisperm not developed. Endosperm copious (Mimulus), sparse or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, convolute, inrolled. Germination cryptocotylar. Radicula ephemeral (Phryma).

Cytology n = 7–10, 14, 22, 23, 27, 30, 32 – Polyploidy frequent. Protein bodies in cell nuclei lamellar.

DNA Deletion in plastid gene matK?

Phytochemistry Iridoids (Route I carbocyclic iridoids?) present (Phryma) or absent (Mimulus). Lignans (phrymarolin-I, phrymarolin-II, haedoxane A, leptostachyolacetate) present. Proanthocyanidins, alkaloids, saponins, and cyanogenic compounds not found.

Use Ornamental plants, insecticides.

Systematics Mimulus (c 120; tropical and southern Africa, Madagascar, India to Manchuria and Japan, Southeast Asia, Malesia to New Guinea, Australia, Tasmania, New Zealand, North America, Mexico, Central America, the Andes in South America); Wightia (2; W. speciosissima, W. borneensis; warm-temperate to tropical regions in the Himalayas, northeastern India, northern Burma, East and Southeast Asia and Malesia).

Phrymaceae are sister-group to the clade [Paulowniaceae+[Rehmanniaceae+Orobanchaceae]].

Mimulus’ and ’Phryma’ in their traditional sense are paraphyletic. Moreover, ‘Mimulus’ is obviously paraphyletic relative to ‘Phryma’, and ‘Phryma’ is here included in Mimulus.

Phrymaceae often appear in the same clade as Paulowniaceae, the Rehmannia clade, Cyclocheilaceae and Orobanchaceae. Phrymaceae have a tubular dentate calyx, usually a loculicidal capsule (rarely a berry), and a bilamellate stigma, receptive only inside and closing when touched.

Cyrtandromoea was fairly highly supported as sister-group to the clade [Mimulus+Uvedalia+[Glossostigma+Peplidium]] in the analyses by Liu & al. (2020), whereas Wightia appeared as sister to all genera of Phrymaceae.

Cyrtandromoea (11–12; Southeast Asia, Malesia) was ambiguously placed in Gesneriaceae-Epithematoideae, identified as sister-group to the clade [Monophyllaea+Rhynchoglossum] in ndhF analyses by Smith, Brown & al. (1997). Cyrtandromoea has endosperm and exotesta with U-shaped laminated thickenings in cross-section. The cotyledons are isocotylous (Weber & al. 2013). Moreover, it has iridoids and other substances similar to the Plantaginaceae and “higher” Plantaginales.

Wightia consists of evergreen to deciduous trees, shrubs or lianas. The hairs are stellate. The leaves are opposite, sometimes with ‘glands’ (domatia?) at vein axils on abaxial side, and have entire margin. The flowers are arranged in lateral racemes or thyrses consisting of cymes. The calyx is campanulate, trilobate or quadrilobate or entire. The corolla limb is 2-lipped, with bilobate upper lip and trilobate lower lip. The four stamens are didynamous, with filaments adnate to the tube base. The anthers are basifixed and the thecae are parallel and fused at apex. Staminodia are absent. The pollen grains are usually tricolporate (sometimes di- or tetracolporate). The style is elongate and simple, and the stigma is punctate. The fruit is a septicidal capsule with persistent central columella. The ovules are numerous in each carpel and the testa has membranous wings. Wightiaceae, sister-group to Phrymaceae and comprising the genus Wightia, were described by Liu & al. (2019).

Cladogram of Phrymaceae based on DNA sequence data (Beardsley & Olmstead 2002).

PLANTAGINACEAE Juss.

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de Jussieu, Gen. Plant.: 89. 4 Aug 1789 [’Plantagines’], nom. cons.

Globulariaceae DC. in de Lamarck et A. P. de Candolle, Fl. Franç., ed. 3, 3: 427. 17 Sep 1805 [’Globulariae’], nom. cons.; Antirrhinaceae Pers., Syn. Plant. 2: 154, 640. Sep 1807 [‘Antihirrhineae’, ’Antirrhineae’]; Veronicaceae Cassel, Lehrb. Nat. Pflanzenord.: 366. Apr-Mai 1817 [’Veronicae’]; Hippuridaceae Vest, Anleit. Stud. Bot.: 265, 278. 1818 [’Hippuroideae’], nom. cons.; Callitrichales Bercht. et J. Presl, Přir. Rostlin: 271. Jan-Apr 1820 [‘Callitricheae’]; Chelonaceae Augier ex Martinov, Tekhno-Bot. Slovar: 124. 3 Aug 1820 [’Cheloneae’]; Digitalidaceae Augier ex Martinov, Tekhno-Bot. Slovar: 202. 3 Aug 1820 [’Digitales’]; Gratiolaceae Martinov, Tekhno-Bot. Slovar: 294. 3 Aug 1820 [’Gratioleae’]; Linariaceae Bercht. et J. Presl, Přir. Rostlin: 243. Jan-Apr 1820 [‘Linariae’]; Callitrichaceae Link, Enum. Hort. Berol. Alt. 1: 7. 16 Mar-30 Jun 1821 [‘Callitrichinae’]), nom. cons.; Littorellaceae Gray, Nat. Arr. Brit. Pl. 2: 290, 294. 10 Jan 1822 [’Littorellideae’]; Globulariales Link, Handbuch 1: 675. 4-11 Jul 1829 [’Globulariaceae’]; Hippuridales Link, Handbuch 1: 288. 4-11 Jul 1829 [‘Hippurideae’]; Psylliaceae Horan., Prim. Lin. Syst. Nat.: 69. 2 Nov 1834; Aragoaceae D. Don in Edinburgh New Philos. J. 19: 109, 113. Jul 1835; Sibthorpiaceae D. Don in Edinburgh New Philos. J. 19: 114. Jul 1835; Plantaginopsida Meisn., Plant. Vasc. Gen.: Comm.: 226. 18-24 Jul 1841 [’Plantagoideae’]; Antirrhinineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1136, 1137. 1846 [‘Antirrhineae‘]; Veronicineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1136, 1140. 1846 [‘Veroniceae‘]; Antirrhinales Döll, Fl. Baden 2: 723. med. 1858 [’Antirrhineae’]; Oxycladaceae (Miers) Schnizlein, Iconogr. Fam. Regni Veg. 2: ad t. 151**. 1857-1870 [’Oxycladeae’]; Erinaceae Duvau ex L. K. G. Pfeiffer, Nomencl. Bot. 1(2): 1236. 16 Jan 1874; Stellariaceae C. MacMill., Metasp. Minnesota Valley: 344. 1892, nom. illeg. – non Stellariaceae Bercht. et J. Presl; Hippuridineae Engl., Syllabus, ed. 5: 175. 20-22 Jul 1907; Callitrichineae Engl. et Gilg in H. G. A. Engler, Syllabus, ed. 7: 240. Oct 1912-Mar 1913; Ellisiophyllaceae Honda in M. Honda et M. Sakisaka, Daikou Nippon Shokubutsu Bunruigaku [Syst. Plant. Japon.]: 373. 25 Apr 1930; Trapellaceae Honda et Sakisaka, Daikou Nippon Shokubutsu Bunruigaku [Syst. Pl. Japon.]: 378. 25 Apr 1930; Hemimeridaceae Doweld, Tent. Syst. Plant. Vasc.: xlix. 23 Dec 2001

Genera/species 92/1.785–1.850

Distribution Cosmopolitan except polar areas (mainly temperate regions).

Fossils Fruits of Trapella have been found in Miocene and Pliocene layers in Europe. Fossil pollen possibly originating from Plantago have been found in Miocene and younger layers in Europe.

Habit Usually bisexual (rarely monoecious, gynomonoecious, dioecious, or gynodioecious), usually perennial, biennial or annual herbs (rarely shrubs or suffrutices). Some genera are aquatic or semi-aquatic. Numerous representatives are xerophytes. At least one species of Philcoxia has carnivorous subterranean leaves.

Vegetative anatomy Phellogen ab initio epidermal to pericyclic; cork tissue usually not developed. Medullary vascular bundles present or absent. Secondary lateral growth normal or absent. Endodermis often prominent. Vessel elements with simple perforation plates; lateral pits alternate, simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids or libriform fibres with simple and/or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually paratracheal scanty vasicentric, or absent (rarely apotracheal diffuse). Cambium sometimes storied (in, e.g., Plantago, Penstemon). Wood elements sometimes storied. Sieve tube plastids S type. Nodes usually 1:1 or 3, unilacunar with one or three leaf traces (sometimes >3:>3, multilacunar with more than three traces). Sclereids sometimes present in stem and roots. Cystoliths present or absent. Calciumoxalate crystals as druses.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, dendritic, sometimes peltate, lepidote or arachnoid (sometimes calcified or silicified); glandular hairs with unicellular stalk and multicellular head (sometimes peltate), sometimes with vertically dividing cells.

Leaves Alternate (spiral) or opposite (rarely verticillate), simple or compound, entire or lobed (in Trapella and closely allied genera anisophylly), rarely coriaceous (in, e.g., Globularieae), with conduplicate or ? ptyxis. Stipules absent; leaf sheath usually absent (petiole base rarely sheathing stem and decurrent). Petiole vascular bundles? Venation usually pinnate or palmate (rarely parallel; leaves sometimes uninerved). Stomata usually anomocytic (sometimes diacytic, anisocytic, paracytic, tetracytic, or cyclocytic), sometimes on adaxial side of lamina only. Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts. Leaf margin serrate, crenate or entire; teeth sometimes with hydathodes. Extrafloral nectaries rarely present on lamina.

Inflorescence Terminal or axillary, cymose or racemose of various shape (usually raceme or spike, rarely capitate; flowers sometimes solitary axillary; paired cymes present in, e.g., Penstemon), sometimes pseudanthia with involucre consisting of bracts. Floral prophylls (bracteoles) often absent in Antirrhinoideae.

Flowers Usually zygomorphic (rarely secondarily actinomorphic: Sibthorpia, Aragoa, Plantago). Usually hypogyny (in Hippuris, Trapella and closely allied genera epigyny). Sepals (two to) four or five (to eight in Sibthorpia), usually with imbricate or valvate aestivation, often persistent, more or less connate (in Trapella free?; absent in Callitriche); median sepal usually adaxial. Petals (two to) four or five (to eight; absent in Callitriche, Hippuris and sometimes in Synthyris and Veronica), usually with imbricate or valvate (rarely descending cochlear) aestivation, connate into hypocrateriform, infundibuliform, campanulate or bilabiate corolla (often with two upper and three lower lobes; sometimes with a spur). Nectariferous disc intrastaminal, annular, unilateral, glandular, or absent.

Androecium Stamens usually two longer and two shorter (rarely five to eight or two [in, e.g., Veronica, Gratiola and Trapella two adaxial-lateral stamens] or one [e.g. Callitriche and Hippuris]), haplostemonous, antesepalous, alternipetalous. Filaments free, usually adnate to corolla tube (epipetalous). Anthers usually with parallel thecae (thecae sometimes entirely fused) or sagittate, dorsifixed, sometimes versatile, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; in Globularioideae with short apical slit on fused thecae); connective in Trapella and closely allied genera large and peltate. Tapetum secretory, with binucleate or multinucleate cells. Staminodia one (sometimes in, e.g., Cheloneae and Antirrhinoideae), two (adaxial staminal pair; in Trapella and closely allied genera two abaxial-lateral stamens modified into staminodia) or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually (2–)3(–6)-colpate or (2–)3(–6)-colporate (in Plantago 3–15-polypantoporate; in Trapella and closely allied genera tricolpor[oid]ate; in Callitriche often inaperturate), usually shed as monads (in Anticharis as tetrads), usually bicellular at dispersal (sometimes tricellular; in Callitriche, Hippuris, and Trapella tricellular). Exine tectate or semitectate, with columellate? infratectum, perforate, reticulate, striate, rugulate or spinulate.

Gynoecium Pistil composed of usually two connate carpels (rarely three; in Hippuris one carpel). Ovary usually superior (rarely inferior), usually bilocular (rarely uni- or trilocular or pseudomonomerous; in Callitriche quadrilocular by secondary septa; abaxial locule degenerating in Trapella and allied genera). Stylodia two, partially free or entirely connate (in Callitriche two, completely free from each other, with stigmatic areas along their entire length). Stigma capitate or bilobate (in Trapella and allied genera non-uniformly bilabiate), papillate or non-papillate, usually Dry (sometimes Wet) type. Female flowers sometimes with pistillodium?

Ovules Placentation usually axile (sometimes axile to apical; rarely intrusively parietal; in Plantago basal to axile; in Globularioideae apical). Ovules one or two (in Trapella and closely allied genera in adaxial locule only) to numerous per carpel, usually anatropous or hemianatropous (rarely amphitropous or campylotropous), pendulous or ascending, apotropous or epitropous, unitegmic, tenuinucellar (reduced, with meiocyte semi-inferior). Integument three to 22 cell layers thick. Megagametophyte usually monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Endothelial cells often small, irregularly arranged and with evenly thick walls. Embryogenesis usually onagrad (in, e.g., Ellisiophyllum pinnatum solanad).

Fruit Usually a septicidal capsule (in, e.g., Gratiola and Veronica septicidal and/or loculicidal; in many Antirrhinoideae poricidal; rarely a pyxidium [in, e.g., Plantago], berry, nut or drupe, sometimes with persistent calyx; in Callitriche a schizocarp with four one-seeded drupaceous or nutlike mericarps; in Trapella and closely allied genera a nut with persistent calyx with five basal alternisepalous vascularized often long and hook-tipped appendages). Placenta sometimes with cushion-shaped scars of detached seeds.

Seeds Aril absent. Seed pedestals often present. Testa in Trapella and closely allied genera with thin cell walls. Exotestal cells with inner walls thickened (in Plantago large, mucilaginous). Endotestal cells in Plantago persistent. Perisperm not developed. Endosperm usually copious (sometimes sparse), oily. Embryo large or small, usually straight (rarely curved), usually well differentiated, usually without chlorophyll. Cotyledons two. Germination phanerocotylar. Radicula in Trapella and closely allied genera ephemeral.

Cytology n = 3–10, 12, 14, 16, 17, 19–21, 24, 26–28 (32) (34) (56) (n = c. 25 in Trapella and closely allied genera) – Polyploidy occurring. Protein bodies in cell nuclei in Gratioloideae lamellar or fibrillar; in Russelioideae lamellar or tubular; in Chelonoideae amorphous; in Antirrhinoideae amorphous, crystalline or tubular; in Globularioideae lamellar; in Digitalidoideae amorphous, crystalline or tubular, sometimes absent.

DNA Deletion in the plastid gene matK. Mitochondrial coxI intron present.

Phytochemistry Flavonols (kaempferol), 6- and/or 8-hydroxyflavones and their glycosides or 6-methoxyflavones, Route II decarboxylated iridoids, Group I carbocyclic iridoids (aucubin, catalpol, 6-O-esters of catalpol, daphylloside, monotropein), Group II carbocyclic iridoids (antirrhinoside, linarioside), Group X secoiridoids (loganin, antirrhide, gardoside, iridoid pyridine alkaloids), iridoid glucosides (e.g. ajugol; absent from some lineages, e.g. Digitalis with digitalis glucosides), terpenes, cardenolides (in, e.g., Digitalis), ursolic acid and caffeic acid esters (cornosides in, e.g., Digitalis and Veronica, verbascosides), saponins, polyols, and shikimic acid derived arthroquinones present. Ellagic acid, tannins, proanthocyanidins, and cyanogenic compounds not found. Carbohydrates stored as stachyose and other oligosaccharides (e.g. saccharose; sometimes also mannitol and sorbitol in Plantago).

Use Ornamental plants, medicinal plants (Digitalis, seeds of Plantago psyllium, etc.).

Systematics (under construction) Plantaginaceae are sister-group to the remaining Plantaginales “above” the Gesneriaceae clade.

Gratioloideae Luerss., Handb. Syst. Bot. 2: 993. Sep 1882 [‘Gratioleae’]

33/385–395. Gratioloideae are sister to the remaining Plantaginaceae, according to some analyses. In other studies (e.g. Rahmanzadeh & al. 2005) Gratioloideae are positioned outside of Plantaginaceae in a polytomy.

Angelonieae Pennell in Proc. Acad. Nat. Sci. Philadelphia 71: 227. 11 Mar 1920

6/c 75. Ourisia (18; Tasmania, New Zealand incl. Stewart Island, the Andes), Angelonia (c 45; Mexico, Central America, the West Indies, tropical South America), Basistemon (6; B. bogotensis, B. intermedius, B. klugii, B. peruvianus, B. silvaticum, B. spinosus; the Andes in Venezuela to Bolivia, western Brazil, Paraguay and northwestern Argentina), Monopera (2; M. micrantha, M. perennis; South America), Monttea (3; M. aphylla, M. chilensis, M. schickendantzii; Chile, western Argentina), Melosperma (1; M. andicola; Chile, western Argentina). – Tropical America. Usually shrubs. Integument five to twelve cell layers thick. Seeds few, with special oil attracting pollinators (at least in Angelonia, Basistemon and Monttea).

Gratioleae Benth. in Edwards’s Bot. Reg. 21: ad t. 1770. 1 Feb 1835

27/310–320. Bacopa (65–70; tropical and subtropical regions on both hemispheres), Maeviella (1; M. cochlearia; northeastern Brazil), Boelckea (1; B. beckii; Bolivia), Benjaminia (1; B. reflexa; Venezuela, Brazil), Braunblanquetia (1; B. litoralis; Venezuela to Argentina), Sophronanthe (1; S. hispida; southeastern United States), Trapella (1; T. sinensis; temperate to tropical regions in eastern Asia), Gratiola (c 25; temperate regions on the Northern Hemisphere, East Malesia to New Guinea, southern and eastern Australia, Tasmania, mountains in South America), Mecardonia (12; southeastern United States, Mexico, Central America, the West Indies, tropical and subtropical South America), Scoparia (10; pantropical), Deinostema (2; D. adenocaula, D. violacea; East Asia), Dopatrium (14; tropical and southern Africa, tropical Asia to New Guinea, tropical Australia), Hydrotriche (3; H. galiifolia, H. hottoniiflora, H. mayacoides; Madagascar), Limnophila (37; tropical and subtropical regions in the Old World), Philcoxia (3; P. bahiensis, P. goiasensis, P. minensis; Brazil), Adenosma (27; China, tropical Asia, northern Australia), Otacanthus (7; O. azureus, O. caeruleus, O. caparaoensis, O. crenatus, O. fernandesii, O. platychilus, O. villosus; Brazil), Achetaria (10; tropical South America), Tetraulacium (1; T. veroniciforme; Brazil), Dizygostemon (2; D. angustifolium, D. floribundum; Brazil), Stemodia (c 60; tropical and subtropical regions on both hemispheres), Darcya (3; D. costaricensis, D. mutisii, D. reliquiarum; Central America), Cheilophyllum (8; the West Indies), Schistophragma (4; S. intermedium, S. mexicanum, S. polystachyum, S. pusillum; western United States, Mexico, Central America), Conobea (16; tropical America), Leucospora (2; L. coahuilensis, L. multifida; eastern North America, Mexico), Schizosepala (1; S. glandulosa; Brazil). – Tropical and subtropical regions on both hemispheres, with their highest diversity in tropical and subtropical America. Integument three to six cell layers thick. Endothelial cells transversely elongate in six to eight vertical rows and thickened only near endosperm. Seeds with longitudinal ridges. Exotestal cells with hook-shaped thickenings. – Philcoxia comprises three Brazilian terrestrial species with subterranean stems and peltate leaves with circinate ptyxis. The adaxial side of their leaves are provided with stalked capitate glands. Protease activity and presence of phosphatases as well as digestion of nematodes and nutrient uptake have been detected. The rare case of circinate leaves in these carnivorous plants are remarkable, since the narrow leaves in both Byblis (Byblidaceae) and Drosophyllum (Drosophyllaceae) have circinate vernation.

Trapella sinensis is an aquatic rhizomatous and stoloniferous herb. The flowers are somewhat zygomorphic and epigynous with two fertile stamens (adaxial pair) and two staminodia. The anther has an apically peltate connective. The pollen grains are tricellular at dispersal. The abaxial ovary loculus degenerates. The stigma is bilobate, with unequal lobes. There are two ovules per carpel and the placentation is apical. The fruit is indehiscent and spiny, with persistent calyx and an appendage from below sepal apex. The testal cells are thin-walled. n = c. 25. Trapella is sister to Gratiola and Amphianthus, according to Gormley & al. (2015).

Russelieae Pennell in Proc. Acad. Nat. Sci. Philadelphia 71: 226. 11 Mar 1920

2/51–55. Tetranema (3; T. evolutum, T. megaphyllum, T. roseum; Central America), Russelia (48–52; Mexico, Central America, Cuba, Colombia). – Mexico, Central America, Cuba, Colombia. Heads of glandular hairs with vertically dividing cells. Fruit in Russelia a unique type of hard and dry capsule. – Russelieae are perhaps sister to Chelonoideae.

Chelonoideae Luerss., Handb. Syst. Bot. 2: 993. Sep 1882 [’Cheloneae’]

9/295–320. Collinsia (c 20; southern Canada, United States, Mexico, with their highest diversity in western United States), Tonella (2; T. floribunda, T. tenella; southwestern Canada, western United States), Keckiella (7; K. antirrhinoides, K. breviflora, K. cordifolia, K. corymbosa, K. lemmonii, K. rothrockii, K. ternata; western United States, northwestern Mexico), Chionophila (2; C. jamesii, C. tweedyi; Rocky Mountains), Uroskinnera (4; U. almedae, U. flavida, U. hirtiflora, U. spectabilis; Mexico, Central America), Brookea (4; B. albicans, B. auriculata, B. dasyantha, B. tomentosa; Borneo), Nothochelone (1; N. nemorosa; northwestern North America), Chelone (5; C. caeruleum, C. cuthbertii, C. glabra, C. lyonii, C. obliqua; southeastern Canada, eastern United States), Penstemon (250–275; North America, northern Mexico). – Borneo, North America, Mexico, Central America. Stem with conspicuous medulla. Hairs simple. Inflorescence cymose. Staminodia present. – Collinsia has papilionoid flowers resembling some species of Lupinus, with velum formed by two adaxial petals and carina consisting of median abaxial petal.Uroskinnera should probably be included in Chelonoideae or Russelieae; it is here tentatively placed in Chelonoideae.

Antirrhinoideae Kostel., Allg. Med.-Pharm. Fl. 3: 874. Apr-Dec 1834 (Benth. in Lindl., Veg. Kingd.: 684. Jan-Mai 1846 [‘Antirrhinideae’])

26/290–300. Antirrhinum (c 20; the Mediterranean, with their largest diversity in Spain), Chaenorhinum (26; the Mediterranean, southwestern Asia), Holzneria (2; southwestern Asia), Albraunia (3; southwestern Asia), Schweinfurthia (8; S. apterum, S. imbricata, S. latifolia, S. papilionacea, S. pedicellata, S. pterosperma, S. sphaerocarpa, S. spinosa; arid and semiarid regions in northeastern Africa to India), Acanthorrhinum (1; A. ramosissimum; northwestern Africa), Misopates (9; Europe, Macaronesia, the Mediterranean to Ethiopia and northwestern India), Linaria (95–100; Europe, the Mediterranean, northern Africa, temperate and subtropical Asia), Nuttallanthus (4; N. canadensis, N. floridanus, N. subandinus, N. texanus; southern Canada, United States, Mexico, western South America), Sairocarpus (12; southwestern United States, northwestern Mexico), Howelliella (1; H. ovata; eastern California), Neogaerrhinum (2; N. filipes, N. kelloggii; southwestern United States, northwestern Mexico), Pseudorontium (1; P. cyathiferum; southwestern United States, northwestern Mexico), Mohavea (2; M. breviflora, M. confertiflora; southwestern United States), Asarina (1; A. procumbens; southern France, northeastern Spain), Cymbalaria (10; Central Europe, the Mediterranean to Iran), Maurandya (8; southwestern United States, Mexico, Central America, the West Indies, tropical South America), Maurandella (1; M. antirrhiniflora; southwestern United States, northwestern Mexico), Epixiphium (1; E. wislizeni; southwestern United States, northwestern Mexico), Mabrya (6; M. acerifolia, M. coccinea, M. erecta, M. flaviflora, M. geniculata, M. rosei; southwestern United States, northwestern Mexico), Holmgrenanthe (1; H. petrophila; southwestern United States), Lophospermum (8; Mexico, Guatemala), Rhodochiton (3; R. atrosanguineus, R. hintonii, R. nubicola; Mexico, Central America), Galvezia (5; G. fruticosa, G. grandiflora, G. juncea, G. lanceolata, G. leucantha; Ecuador, the Galápagos Islands, coastal areas in Peru), Gambelia (4; G. glabrata, G. juncea, G. rupicola, G. speciosa; southwestern United States, northwestern Mexico), Kickxia (24; Europe, Macaronesia, the Mediterranean, northern and northeastern Africa, southwestern and southern Central Asia), Nanorrhinum (29; tropical and subtropical regions in the Old World), Anarrhinum (7; A. bellidfolium, A. corsicum, A. duriminium, A. forsskaolii, A. fruticosum, A. longipedicellatum, A. pedatum; Central and South Europe, the Mediterranean, Ethiopia). – Temperate regions on the Northern Hemisphere, northern and northeastern Africa, southwestern Asia, Mexico to western South America and the West Indies, few species in tropical regions of the Old World. Floral prophylls often absent. Flowers zygomorphic. Fruit a unique type of poricidal capsule. Antirrhinosides (unique group of iridoids) present. – Antirrhinoideae are sister to the clade [Globularioideae+Digitalidoideae].

Globularioideae Luerss., Handb. Syst. Bot. 2: 1038. Sep 1882 [‘Globulariaceae’]

3/52. Campylanthus (18; the Canary Islands, northeastern Africa, the Arabian Peninsula, Pakistan), Globularia (31; Europe, Madeira, the Canary Islands, Cape Verde Islands, the Mediterranean, Turkey, North and northeastern Africa), Poskea (3; P. africana, P. newbouldii, P. socotrana; Somalia, Socotra). – Europe, Macaronesia, the Mediterranean to Pakistan, northern and northeastern Africa, the Arabian Peninsula, Socotra. Phellogen epidermal to pericyclic. Tracheids present (fibre tracheids and libriform fibres are absent). Leaves with conduplicate ptyxis. Ovules one or two per carpel (one pendulous and one erect) or, alternatively, only abaxial carpel developed. Stigma Dry type. Fruit a nutlet with membranous pericarp. n = 8–10, 16 (19?). – Globularioideae are sister-group to Digitalidoideae.

Digitalidoideae Luerss., Handb. Syst. Bot. 2: 994. Sep 1882 [’Digitaleae’]

16/650–665

Digitalideae Dumort., Anal. Fam. Plant.: 24. 1829

3/c 28. Digitalis (c 25; Europe, Madeira, the Canary Islands, the Mediterranean to Central Asia), Erinus (2; E. alpinus: the Alps, the Pyrenees; E. thiabaudii: Morocco), Lafuentea (1; L. rotundifolia; southern Spain). – Europe, Macaronesia, the Mediterranean to Central Asia, Morocco. Corolla with descending-cochlear aestivation. Cornosides present (Digitalis).

Plantagineae Dumort., Anal. Fam. Plant.: 25. 1829

3/175–180. Aragoa (19; the Andes in Colombia and Venezuela), Plantago (155–160; cosmopolitan except polar regions), Littorella (1–3; L. uniflora: Europe, the Azores; L. americana: temperate North America; L. australis: temperate South America). – Almost cosmopolitan. Nodes 3:3. Leaves alternate (spiral). Leaf margin serrate or entire. Flowers actinomorphic, small. Sepals and petals usually four (occasionally three; in Aragoa five). Corolla with descending-cochlear aestivation. Pistil composed of two connate carpels. Ovary unilocular or bilocular. Style long-branched, stigmatic its entire length. Stigma Dry type. Placentation usually parietal (sometimes basal, with ovule single, epitropous). Fruit a pyxidium or a nut. Exotestal cells large, mucilaginous. Endotestal cells persistent. Endosperm present. Embryo usually straight (sometimes curved). Iridoid glucosides with 8,9 double bonds and sorbitol present. – Aragoa and Plantago are sister-groups. The substitution frequency in synonymous positions of the mitochondrial genome in Plantago is three to four thousand times higher than in nearly all other angiosperms (Mower & al. 2007). Three or more mitochondrial genes were shown to have been transferred from Cuscuta to the Plantago coronopus lineage (Mower & al. 2010).

Sibthorpieae Benth. in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 10: 189, 424. 8 Apr 1846

2/6. Ellisiophyllum (1; E. pinnatum; India, Southeast and East Asia to Japan and east to the Philippines, Taiwan and New Guinea), Sibthorpia (5; S. africana, S. conspicua, S. europaea, S. peregrina, S. repens; Europe, the Azores, Madeira, mountains in tropical East Africa, tropical America). – Europe, Macaronesia, East African mountains, India to Japan, the Philippines, Taiwan and New Guinea, tropical America. Flowers actinomorphic. Sepals in Sibthorpia up to eight. Megasporangium development unique type in Plantaginaceae. Embryogenesis in Ellisiophyllum solanad. – Sibthorpieae lack the iridoids typical of Plantaginales.

Veroniceae Duby, Bot. Gall. 1: 355. 12-14 Apr 1828

c 7/440–450. Kashmiria (1; K. himalaica; the Himalayas), Lagotis (c 20; eastern Europe, the Caucasus, northern and Central Asia, the Himalayas to western China), Picrorhiza (3; P. kurroa, P. minima, P. scrophulariiflora; the Himalayas), Scrofella (1; S. chinensis; northwestern China), Veronica (410–420; temperate and alpine regions on both hemispheres, tropical African mountains, New Zealand and surrounding islands, Tierra del Fuego, western Patagonia north to 45°53’ south latitude, the Falkland Islands), Wulfenia (4; W. baldacii, W. blechicii, W. carinthiaca, W. orientalis; southeastern Europe, Turkey), Wulfeniopsis (1; W. amherstiana; the Himalayas in eastern Afghanistan, Pakistan, northern India and Nepal). – Temperate and alpine regions on both hemispheres. Corolla with descending-cochlear aestivation. Stamens in Veronica two. Iridoid glucosides with 8,9 double bonds present.

Hemiphragmateae Rouy, Consp. Fl. France: 172. 15 Aug 1927 [‘Hemiphragmeae’]

1/1. Hemiphragma (1; H. heterophyllum; the Himalayas in northern India, Nepal, Sikkim, Assam and Bhutan to China, Taiwan, the Philippines, Sulawesi). – Perennial herb. Flowers zygomorphic. Corolla quinquelobate, campanulate. Thecae apically connate. Fruit a berry-like capsule.

Callitrichoideae Arn., Botany: 110. 9 Mar 1832 [‘Callitricheae’]

2/c 60. Callitriche (c 60; nearly cosmopolitan incl. the Falkland Islands, South Georgia Islands, Auckland Islands, Campbell Island, Antipodes and Antarctic islands), Hippuris (1; H. vulgaris; nearly cosmopolitan). – Subcosmopolitan. Aquatic, unisexual (monoecious). Flowers actinomorphic (zygomorphic by reduction), minute. Petals absent. Pollen grains tricellular at dispersal. Ovules two per carpel. Callitriche: Hairs stellate-peltate. Sepals absent. Stamens one (to three). Pollen grains often inaperturate. Exine intectate, reticulate or echinate (sometimes absent). Pistil composed of two transverse carpels. Ovary locules with secondary septa. Stylodia thin, separate. Integument thin. Fruit a schizocarp with four nutlike mericarps. Exotesta possibly persistent. n = 3–20. – Hippuris: Usually bisexual (sometimes unisexual). Hairs peltate, glandular. Leaves 4–16-verticillate. Epigyny. Calyx bilobate to quadrilobate or entire. Stamen single. Pollen grains colpoidate. Pistil composed of a single carpel. Style single, with stigmatic area extending its entire length. Stigma Dry type. Ovules pendulous, apotropous. Chalazogamy. Fruit a drupe or achene-like. Endosperm thin, starchy. n = 8, 15, 16. Flavones not found.

Trapelleae Stapf in Engl. et Prantl, Nat. Pflanzenfam. IV, 3b: 260, 265. Mai 1893

1/1–2. Trapella (1–2; temperate to tropical regions in eastern Asia). – Aquatic rhizomatous and stoloniferous herb. Flower somewhat zygomorphic. Epigyny. Stamens two (adaxial pair). Anthers with apically peltate connective. Staminodia two. Pollen grains tricellular at dispersal. Abaxial ovary loculus degenerating. Stigma bilobate, with unequal lobes. Placentation apical. Ovules two per carpel. Fruit indehiscent, spiny, with persistent calyx and an appendage from below sepal apex. Testal cells thin-walled. n = c. 25.

Phylogeny of Plantaginaceae based on DNA sequence data (Schäferhoff & al. 2010).

Genus incertae sedis

Cubitanthus (1; C. alatus; Bahia in Brazil) is a perennial herb with multicellular hairs, four-winged stem, opposite leaves with serrate margins, solitary axillary flowers without bracteoles, persistent calyx with free sepals, bilabiate corolla limb with trilobite lower lip, four didynamous stamens adnate to corolla base, annular nectariferous disc, superior ovary with parietal placentae, capitate stigma, septicidal bivalvular capsule, and striate seed surface. – It was placed in Gesneriaceae, although it is more similar to Plantaginaceae or Scrophulariaceae.

PLOCOSPERMATACEAE Hutch.

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Hutchinson, Fam. Fl. Pl. ed. 3, 1: 469. Jan-Jul 1973

Genera/species 1/1

Distribution Southern Mexico, Guatemala, Costa Rica.

Fossils Unknown.

Habit Functionally unisexual (cryptic dioecy), evergreen trees or shrubs.

Vegetative anatomy Phellogen? Prominent groups of fibres present in outer cortex, in places where leaf traces leave stem. Vessels present in radial multiples. Vessel elements with simple perforation plates; lateral pits alternate. Vestured pits? Imperforate tracheary xylem elements thick-walled fibre tracheids with bordered pits, septate or non-septate. Wood rays usually uniseriate, heterocellular. Axial parenchyma paratracheal scanty, lignified. Sieve tube plastids S type? Nodes 1:1?, unilacunar with one? leaf trace. Styloids present. Parenchyma with druses and single (prismatic?) crystals.

Trichomes Hairs unicellular, simple, filled with calciumcarbonate and accompanied by cystoliths in adjacent basal epidermal cells (or only cystoliths); glandular hairs bicellular, clavate; cells in head of glandular hair not entirely with vertical walls.

Leaves Opposite (often almost verticillate), simple, entire, coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Colleters absent. Petiole articulated at base. Petiole vascular bundle transection annular. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids absent. Styloids present. Leaf margin entire or somewhat serrate. Hairs long, unicellular, simple.

Inflorescence Axillary, raceme-like (flowers sometimes solitary axillary). Floral prophylls (bracteoles) absent.

Flowers Somewhat zygomorphic. Hypogyny. Sepals five (or six), with imbricate or open aestivation, persistent, connate at base. Petals five (or six), with imbricate (contorted?) aestivation, equal or slightly unequal in size, connate into campanulate or infundibuliform corolla. Nectariferous disc at ovary base (hence only in female flowers).

Androecium Stamens five (or six), unequal in size, haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers basifixed (dorsifixed?), versatile, tetrasporangiate, extrorse (to latrorse?), longicidal (dehiscing by longitudinal slits). Placentoid? Tapetum secretory? Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate? infratectum, reticulate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, unilocular, short-stalked (gynophore?). Stylodia two, bifid, caducous, without indusium, sometimes slightly connate at base. Stigmas clavate, not expanded, type? Male flowers with pistillodium.

Ovules Placentation basal to parietal and/or apical to parietal. Ovules two or four per ovary, ascending and/or pendulous, two parietal placentae each with one or two basal erect ovules or with two basal erect ovules on one placenta and two subapical pendulous ovules on the other, unitegmic, tenuinucellar? Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A usually one-seeded (rarely two- to four-seeded) loculicidal capsule with two valves.

Seeds Aril? Chalazal end of seed with dense coma consisting of long multicellular hairs. Testa? Perisperm not developed. Endosperm sparse, thin, fleshy. Embryo straight, large, well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = ?

DNA

Phytochemistry Insufficiently known. Caffeic acid and ursolic acid esters (caffeoyl phenylethanoid glucosides, e.g., echinacoside, compounds of the verbascoside group, cornoside, a quinole glucoside), and lugrandoside (a phenylpropanoid glucoside) present. Iridoids not found.

Use Unknown.

Systematics Plocosperma (1; P. buxifolium; southern Mexico, Guatemala, Costa Rica).

Plocosperma is sister to the remaining Plantaginales.

REHMANNIACEAE Kunkel ex Reveal

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Reveal in Kew Bull. 66: 47. Mar 2011

Genera/species 2/8

Distribution China, the Korean Peninsula.

Fossils Unknown.

Habit Bisexual, perennial or annual herbs.

Vegetative anatomy Phellogen? Vessel elements with simple? perforation plates; lateral pits? Vestured pits? Imperforate tracheary xylem elements? with simple? pits. Wood rays absent. Axial parenchyma? Sieve tube plastids S type? Nodes in Rehmannia 1:3, unilacunar with three leaf traces. Crystals?

Trichomes Hairs unicellular or multicellular; brown to white glandular hairs present in Rehmannia; gead of glandular hairs without vertical septa (not dividing vertically).

Leaves Alternate (spiral), simple, entire or pinnately lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate; petiole with wing bundles. Venation pinnate. Stomata anomocytic? Cuticular wax crystalloids? Leaf margin serrate, crenate, lobate or entire.

Inflorescence Axillary, few-flowered, cymose, or solitary (Rehmannia), or terminal raceme (Rehmannia, Triaenophora). Floral prophylls (bracteoles) one or two, lateral, present at pedicel base, immediately above foliaceous bract, or absent (aborted in most species of Rehmannia).

Flowers Zygomorphic. Hypogyny. Sepals five (to seven; calyx lobes in Rehmannia entire; in Triaenophora each calyx lobe trilobate), with ? aestivation, connate. Petals five, with imbricate quincuncial aestivation (posterior corolla lobes in bud covered by lateral lobes, abaxial lobe situated outside remaining lobes), connate into infundibular bilabiate corolla. Nectariferous disc intrastaminal, at base of ovary.

Androecium Stamens four, antesepalous, alternipetalous, initiated one-way from abaxial to adaxial side, didynamous (adaxial stamen absent). Filaments free from each other, adnate to corolla tube. Anthers dorsifixed, versatile?, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal lobes). Tapetum secretory? Staminodia usually absent (sometimes one).

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, in Rehmannia usually bilocular (rarely unilocular), in Triaenophora bilocular. Style single, simple. Stigma bilamellate, sensitive, type? Pistillodium absent.

Ovules Placentation axile. Ovules numerous per ovary, anatropous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria micropylar and chalazal. Embryogenesis?

Fruit A loculicidal capsule with persistent calyx.

Seeds Aril absent. Exotesta cells with with complex reticulate wall thickenings. Endotesta? Perisperm not developed. Endosperm? Embryo?, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = ? – Protein bodies in cell nuclei lamellar.

DNA Mitochondrial coxI intron present.

Phytochemistry Group I carbocyclic iridoids (catalpol, ajugol, 6-feruloylajugol, aucubin), iridoid glucosides, caffeoyl phenylethanoid glycosides, and ionone glucosides present. Harpagide and 6-rhamnopyranosyl-catalpol and their esters not found. Mannitol present. Sorbitol not found.

Use Ornamental plants, medicinal plants (Rehmannia).

Systematics Rehmannia (6; R. chingii, R. elata, R. glutinosa, R. henryi, R. piasezkii, R. solanifolia; China, the Korean Peninsula), Triaenophora (2; T. integra, T. rupestris; northeastern China).

Rehmanniaceae are sister-group to Orobanchaceae.

SCHLEGELIACEAE (A. H. Gentry) Reveal

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Reveal in Phytologia 79: 74. 29 Apr 1996

Genera/species 4/34–39

Distribution Tropical America.

Fossils Unknown.

Habit Bisexual, evergreen trees or shrubs (sometimes twining, often epiphytic). Bark often whitish.

Vegetative anatomy Phellogen? Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits? Imperforate tracheary xylem elements libriform fibres? with simple or bordered pits, septate. Wood rays uniseriate or multiseriate, homocellular. Axial parenchyma paratracheal vasicentric. Sieve tube plastids S type? Nodes 1:3, unilacunar with three leaf traces. Pericycle with sclereids. Acicular crystals, styloids, crystal sand and other types of calciumoxalate crystals often present.

Trichomes Eglandular hairs?; glands sometimes present on lamina.

Leaves Opposite, simple, entire (in Gibsoniothamnus anisophyllous), often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection solid to almost annular; petiole with wing bundles, without pericyclic lignification. Venation pinnate. Stomata in Schlegelia anomocytic or paracytic, in Gibsoniothamnus anisocytic or cyclocytic. Cuticular wax crystalloids? Petiole in Schlegelia with sclereids. Leaf margin serrate (in Synapsis spinose-dentate) or almost entire. Lamina sometimes with small abaxial glandular hairs with radially arranged head cells.

Inflorescence Terminal or axillary, usually cymose (thyrse; in Exarata racemose).

Flowers Zygomorphic, often large. Hypogyny. Sepals (three to) five, with imbricate aestivation, connate, in Schlegelia with vascularized multicellular nectariferous glands inserted in groups and sunken into epidermis on abaxial surface of calyx. Petals five, with ascending-cochlear aestivation, connate into tubular bilabiate (upper lip bilobate, lower lip trilobate) corolla. Nectariferous disc present, vascularized from carpellary bundles, or absent.

Androecium Stamens two longer and two shorter (didynamous), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia present or absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate or triporate, shed as monads, ?-cellular at dispersal. Exine tectate to semitectate, with columellate infratectum, reticulate to finely reticulate or psilate-foveolate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular? Style single, simple. Stigma capitate, bilobate or trilobate, papillate?, Wet type? Pistillodium absent.

Ovules Placentation axile; placentae sometimes widened. Ovules ? per carpel, anatropous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Fruit A baccate fruit with lignified pericarp and persistent calyx.

Seeds Aril absent. Exotestal cells with scalariform thickenings on inner periclinal walls or mucilaginous without outer periclinal walls. Endotesta? Perisperm not developed. Endosperm usually present. Embryo stout, straight, well differentiated, chlorophyll? Cotyledons two, lobate. Germination phanerocotylar.

Cytology n = 20

DNA

Phytochemistry Unknown.

Use Timber?

Systematics Gibsoniothamnus (12; southern Mexico, Central America, northern Colombia), Schlegelia (20–25; southern Mexico, Central America, the West Indies, tropical South America), Synapsis (1; S. ilicifolia; eastern Cuba), Exarata (1; E. chocoensis; Colombia, Ecuador).

Schlegeliaceae are sister-group to Martyniaceae.

SCROPHULARIACEAE Juss.

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de Jussieu, Gen. Plant.: 117. 4 Aug 1789 [’Scrophulariae’], nom. cons.

Myoporaceae R. Br., Prodr. Fl. Nov.-Holl.: 514. 27 Mar 1810 [‘Myoporinae’], nom. cons.; Caprariaceae Martinov, Tekhno-Bot. Slovar: 102. 3 Aug 1820 [‘Caprariae’]; Myoporales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 245. Jan-Apr 1820 [‘Myoporinae’]; Verbascaceae Bercht. et J. Presl, Přir. Rostlin: 243. Jan-Apr 1820 [’Verbaceae’]; Selaginaceae Choisy [Mém. Sélag. 19] in Mém. Soc. Phys. Genève 2: 89. 1823 [‘Selagineae’], nom. cons.; Scrophulariineae Link, Handbuch 1: 531. 4-11 Jul 1829 [’Scrofularinae’]; Scrophulariales Lindl., Nix. Plant.: 20. 17 Sep 1833 [’Scrophulales’]; Bontiaceae Horan., Prim. Lin. Syst. Nat.: 77. 2 Nov 1834 [’Bontiaceae (Myoporin.)’]; Hebenstretiaceae Horan., Prim. Lin. Syst. Nat.: 76. 2 Nov 1834 [‘Hebenstreitiaceae (Selagineae)’]; Selaginales Choisy in C. F. P. von Martius, Consp. Regn. Veg.: 19. Sep-Oct 1835 [‘Selagineae’]; Selaginopsida Brongn., Enum. Plant. Mus. Paris: xix, 64. 12 Aug 1843 [’Selaginoideae’]; Verbascineae J. Presl in Nowočeská Bibl. [‘Wšobecný Rostl.] 7: 1136, 1137. 1846 [‘Verbasceae‘]; Limosellaceae J. Agardh, Theoria Syst. Plant.: 340. Apr-Sep 1858 [‘Limoselleae’]; Spielmanniaceae J. Agardh, Theoria Syst. Plant.: 194. Apr-Sep 1858 [’Spielmannieae’], nom. illeg.; Verbascales Döll, Fl. Baden 2: 750. med. 1858 [’Verbasceae’]; Buddlejaceae K. Wilh., Samenpflanzen: 90. Oct 1910 [‘Buddleiaceae’], nom. cons.; Oftiaceae Takht. et Reveal in Phytologia 74: 284. 28 Apr 1993

Genera/species 53/1.830–1.850

Distribution Mainly subtropical and warm-temperate regions on the Northern and Southern Hemispheres, with their largest diversity in South Africa and Australia; some representatives in tropical East Africa, Madagascar, tropical Asia, New Guinea, tropical Australia and tropical America.

Fossils Unknown.

Habit Usually bisexual (rarely functionally dioecious), perennial or annual herbs, evergreen or deciduous shrubs (sometimes suffrutices or biennial herbs, rarely trees or lianas). Many species are xerophytes.

Vegetative anatomy Phellogen ab initio superficial, or in inner cortex or pericycle. Primary medullary rays narrow or alternately narrow and wide. Endodermis sometimes prominent (Scrophularia). Vessel elements with simple perforation plates; lateral pits alternate, simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres usually with simple (rarely bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular, or absent. Axial parenchyma paratracheal scanty vasicentric, confluent, or banded, or absent. Wood elements sometimes storied (at least in Myoporeae). Intraxylary phloem present in young stems of Oftia (and Teedia?). Sieve tube plastids S type. Nodes 1:1 or 1:3, unilacunar with one or three leaf traces, often girdling bundle. Primary cortex and medulla in Myoporeae usually with secretory cavities containing oils and resins. Prismatic and acicular calciumoxalate crystals, styloids and crystal sand present in some species.

Trichomes Hairs simple, unicellular or multicellular, or branched, sometimes dendritic, stellate, lepidote, peltate or with globular terminal cell (sometimes calcified/silicified); glandular hairs often present.

Leaves Alternate (spiral), opposite or verticillate, simple, entire or lobed, sometimes coriaceous, sometimes ericoid, often with flat ptyxis. Stipules usually absent (in Buddleja sometimes with interpetiolar stipule-like lobes at base, rarely foliaceous); leaf sheath absent. Petioles sometimes fused in pairs at base. Petiole vascular bundles?; petiole often with wing bundles. Venation pinnate or palmate. Stomata usually anomocytic (sometimes anisocytic or paracytic). Cuticular wax crystalloids? Secretory cavities containing oil or resin present in Myoporeae; lamina often with translucent or raised dots when secretory cavities large. Idioblasts (tanniniferous etc.) present in, e.g., Scrophularia and Verbascum. Leaf margin serrate, crenate or entire. Extrafloral nectaries often present.

Inflorescence Terminal or axillary, usually cymose (sometimes racemose), often panicle, thyrsoid, raceme-, spike- or headlike (flowers rarely solitary axillary).

Flowers Usually zygomorphic (sometimes actinomorphic). Hypogyny. Sepals (two to) five, often with open or imbricate aestivation, often unequal in size, often persistent, usually connate (in, e.g., Myoporeae often free); median sepal adaxial. Petals (four or) five, usually with imbricate (sometimes valvate) aestivation, connate into usually hypocrateriform, infundibuliform or tubular (sometimes campanulate or bilabiate) corolla (rarely with one or two spur-like appendages). Nectary small or absent. Disc intrastaminal. Many species with floral hairs secreting oils attracting pollinators.

Androecium Stamens usually two longer and two shorter (sometimes two or four stamens equal in length [e.g. Buddleja] or five stamens [e.g. Capraria and most species of Verbascum]), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, usually adnate to corolla tube (epipetalous). Anthers with thecae often confluent at apex, head-to-head, U-shaped or parallel, dorsifixed, sometimes versatile, usually tetrasporangiate (rarely disporangiate), introrse, usually longicidal (dehiscing by longitudinal slits, in Myoporeae sometimes by transverse slits). Tapetum secretory. Staminodia one (adaxial-median), two (adaxial-lateral) or three (one adaxial-median and two adaxial-lateral) or absent; female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colpate or (2–)3(–5)-colporate (colpi sometimes with two ora), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, smooth (Buddleja).

Gynoecium Pistil composed of usually two connate carpels (rarely one carpel). Ovary superior, usually bilocular (rarely unilocular, pseudomonomerous, or quadrilocular to decemlocular due to in-growing secondary septa). Style single, simple, sometimes persistent. Stigma usually capitate (sometimes bifid, rarely lingulate), papillate, Dry or Wet type. Male flowers with pistillodium.

Ovules Placentation usually axile (sometimes, e.g. in many Limoselleae apical; in Limosella free central in upper part). Ovules usually numerous (in many Limoselleae one; in Oftia and Tetraselago four; in Myoporeae sometimes one or two) per carpel, anatropous to hemianatropous, pendulous or ascending, apotropous, epitropous or pleurotropous, unitegmic, tenuinucellar. Integument five to twelve cell layers thick. Hypostase present in Buddleja. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes persistent. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Some endothelial cells, bothroblasts, protruding into endosperm (surface then becoming alveolated). Embryogenesis onagrad (Buddlejeae, Myoporeae) or solanad?

Fruit A usually septicidal and septifragal (sometimes also apically loculicidal) capsule (rarely a berry, drupe [Oftia, some Myoporeae and Buddleja], nutlet or schizocarp with one or two drupaceous mericarps [some Myoporeae and Limoselleae]). Placentae often with cushion-shaped scars of detached seeds.

Seeds Aril absent. Seed pedestals often present. Testa sometimes winged, sometimes multiplicative. Exotestal and endotestal cells usually with thickened inner walls (exotestal cells sometimes longitudinally prolonged, in Buddleja with thickened inner walls). Perisperm not developed. Endosperm copious, oily, sparse or absent (sometimes ruminate due to invaginations [unequal radial elongation] of endotestal cells). Embryo usually straight (sometimes curved), well differentiated, usually without chlorophyll. Cotyledons two. Germination phanerocotylar. Radicula in Limosella ephemeral.

Cytology n = 6–10, 12–20, 23–27 (n = 15, 18, 27, 34 in Myoporeae; n = 19 in Oftia) – Polyploidy occurring. Protein bodies in cell nuclei in Verbascum usually amorphous, in Teedieae and Scrophularieae usually lamellar, in Limoselleae and Hemimerideae lamellar, or absent.

DNA Deletion in plastid gene matK. Mitochondrial coxI intron present in Scrophularia and in Celsia clade of Verbascum.

Phytochemistry Flavonols (kaempferol), 6- and/or 8-hydroxylated flavone glycosides, 6- or 8-hydroxyflavones or 6-methoxyflavones, Group I carbocyclic iridoids (aucubin, catalpol, 6-O-rhamnopyranosylcatalpol), Group II carbocyclic iridoids (harpagide, harpagioside [8β-8α-methyl substituted iridoids]), Group X secoiridoids (nepeta lactones), sesquiterpenes, alkaloids, saponins, phenylalanine-derived cyanogenic compounds, shikimic acid derived arthroquinones, and caffeic acid esters of phenylethanoid glucosides (caffeoyl phenylethanoid glucosides: verbascosides, cornosides), and acetylenes (in Myoporeae) present. Ellagic acid, tannins and proanthocyanidins not found. Carbohydrates stored as stachyose and other oligosaccharides (not as mannitol).

Use Ornamental plants, medicinal plants, timber (Myoporum).

Systematics (under construction) Scrophulariaceae are sister-group to the remaining Plantaginales “above” Plantaginaceae.

The clade [Buddleja+Teedieae+Camptoloma+Phygelius] may be sister to [Scrophularieae+Limoselleae].

Colpias

1/1. Colpias (1; C. mollis; Northern Cape). – Small shrub. Leaves alternate (spiral). Leaf margin serrate. Flowers zygomorphic. Calyx quinquelobate. Corolla quinquelobate (2:3), with tube somewhat curved. Corolla at base with two pouches beset with oil-secreting glandular hairs. Fruit a septicidal capsule. n = 10. – Colpias may be sister to the remaining Scrophulariaceae.

[Hemimerideae+Myoporoideae+[[Buddleja+Teedieae+Camptoloma+Phygelius]+[Scrophularieae+Limoselleae]]]

Hemimerideae Benth. in Edwards’s Bot. Reg. 21: ad t. 1770. 1 Feb 1835

5/160–165. Diascia (c 70; southern Africa), Nemesia (60–65; tropical and southern Africa, especially Western Cape), Alonsoa (c 12; Western and Eastern Cape, Mexico to Chile), Hemimeris (7; H. centrodes, H. gracilis, H. montana, H. nana, H. pachyceras, H. racemosa, H. sabulosa; Northern, Western and Eastern Cape), Diclis (10; tropical and southern Africa, Madagascar). – Tropical and southern Africa, Madagascar, Mexico to Chile. Flowers in Hemimeris inverted. Corolla spurred (in Diascia with two spurs). Flowers with oil-secreting hairs. Protein bodies in cell nuclei lamellar.

Myoporoideae Arn., Botany: 123. 9 Mar 1832 [‘Myoporinae’]

10/295–300. Lamina with pellucid gland dots. Flowers usually zygomorphic (in Eremophila tubular).

[Aptosimeae+[Leucophylleae+Myoporeae]]

Aptosimeae Benth. et Hook. f., Gen. Plant. 2: 915, 916. 1-16 Mai 1876

3/41. Anticharis (14; Africa, the Arabian Peninsula to Malesia), Aptosimum (c 20; tropical and southern Africa), Peliostomum (7; P. calycinum, P. leucorrhizum, P. lugardiae, P. oppositifolium, P. origanoides, P. virgatum, P. viscosum; tropical and southern Africa). – Africa, the Arabian Peninsula to Malesia.Aptosimeae are sister to the [Leucophylleae+Myoporeae] clade.

[Leucophylleae+Myoporeae]

Pollen grains tricolpate. Colpi with two ora.

Leucophylleae Miers in Ann. Mag. Nat. Hist., ser. 2, 5: 252. Apr 1850

3/15. Eremogeton (1; E. grandiflorus; southern Mexico, Guatemala), Leucophyllum (12; southwestern United States, Mexico), Capraria (2; C. biflora, C. mexicana; Mexico, Central America). – Southwestern United States to South America. Leaves in Capraria with abundant pellucid glands, in Leucophyllum with one pellucid gland at apex, and in Eremogeton without pellucid glands. Stamens in Capraria five. – Leucophylleae are sister to Myoporeae.

Myoporeae Rchb., Handb. Nat. Pfl.-Syst.: 196. 1-7 Oct 1837 [‘Myoporinae’]

4/c 240. Androya (1; A. decaryi; southern Madagascar); Eremophila (210–215; Australia, one species, E. debilis, also in New Zealand, with their highest diversity in southwestern Western Australia), Bontia (1; B. daphnoides; the West Indies, tropical South America), Myoporum (c 30; Mauritius, Southeast Asia to China, East Malesia to New Guinea, Australia, New Zealand, Kermadec Islands, Polynesia including the Hawaiian Islands). – Mauritius, East and Southeast Asia, East Malesia to New Guinea, Australia, New Zealand to the Hawaiian Islands, the West Indies, tropical South America. Usually shrubs (sometimes herbs). Wood elements often storied. Primary cortex and medulla usually with secretory cavities containing oils and resins. Secretory oil- or resin-containing cavities present. Sepals often free. Anthers sometimes dehiscing by transverse slits. Ovules few (sometimes one or two) per carpel, epitropous. Fruit a drupe or a schizocarp. Endosperm sparse. n = 15, 18, 27, 34. Acetylenes present. – Myoporeae are sister to Leucophylleae. Androya is probably sister to the remaining Myoporeae.

[Teedieae+Buddleja+Camptoloma+Phygelius+[Limoselleae+Scrophularieae]]

Teedieae Benth. in Edwards’s Bot. Reg. 21: ad t. 1770. 1 Feb 1835

6/18–19. Ranopisoa (1; R. rakotosonii; Madagascar), Dermatobotrys (1; D. saundersii; Eastern Cape, KwaZulu-Natal), Freylinia (9; tropical and southern Africa), Teedia (2; T. lucida, T. pubescens; southern Africa), Oftia (3; O. africana, O. glabra, O. revoluta; Northern, Western and Eastern? Cape), Ameroglossum (1; A. pernambucense; Brazil)? – Tropical and southern Africa, Madagascar, Brazil? Young stems and branches in Oftia with intraxylary phloem. Inflorescence in Oftia raceme. Ovules in Oftia four per carpel. Fruit in Oftia a drupe. Endosperm in Oftia copious. n = 19 (Oftia). Protein bodies in cell nuclei usually lamellar. – Teedieae may be sister to Buddleja. The position of the Brazilian Ameroglossum is highly uncertain.

Buddlejeae Bartl., Ord. Nat. Plant.: 172. Sep 1830 [‘Buddlejea’]

1/c 140. Buddleja (c 140; warm-temperate to tropical regions in Africa, Madagascar, Asia and America, with their highest diversity in East Asia). – Shrubs or small treews. Interpetiolar stipule-like lobes (rarely foliaceous) sometimes present at base. Flowers often actinomorphic. Corolla sometimes quadrilobate. Stamens sometimes four. Fruit sometimes a drupe.

Camptoloma

1/3. Camptoloma (3; C. canariense: Gran Canaria; C. lyperiiflorum: Somalia, southern Yemen, Socotra, Kuria Muria Islands; C. rotundifolium: western deserts in Angola, northern Namibia). – Small shrubs. Leaves opposite or spiral, glandular. Leaf margin serrate. Flowers zygomorphic. Calyx quinquelobate. Corolla quinquelobate, with tube curved. Stamens four. Ovules numerous per carpel. Fruit a bivalvular capsule.

Phygelius

1/2. Phygelius (2; P. aequalis, P. capensis; southern Africa). – Perennial herbs. Leaves opposite. Flowers zygomorphic. Corolla quinquelobular, narrowly tubular. Fruit a capsule.

Scrophularioideae Beilschm. in Flora 16(Beibl. 7): 67. 14 Jun 1833 [‘Scrofularomae’]

Limoselleae Dumort., Fl. Belg.: 32. 1827

25/640–650. Jamesbrittenia (c 85; Angola, Zambia to South Africa, one species also in Egypt, Sudan and southwestern Asia to India); Manuleopsis (1; M. dinteri; Namibia), Barthlottia (1; B. madagascariensis; Madagascar), Limosella (c 15; cosmopolitan), Manulea (74; southern Africa, India, with their highest diversity in Western Cape), Sutera (3; S. cooperi, S. griquensis, S. foetida; southern Africa from the Orange Free State and Transvaal to the Cape Provinces), Chaenostoma (46; Africa south of the Cunene and Zambezi rivers, with their highest diversity in the Cape Provinces, Natal and Transvaal), Lyperia (6; L. antirrhinoides, L. formosa, L. lychnidea, L. tenuiflora, L. tristis, L. violacea; Namibia to Western Cape), Glekia (1; G. krebsiana; Eastern Cape, Lesotho), Trieenea (10; Western and Eastern Cape), Hebenstretia (c 40; tropical and southern Africa north to Eritrea), Dischisma (11; Namibia, Western and Eastern Cape), Chenopodiopsis (3; C. chenopodioides, C. hirta, C. retrorsa; Northern and Western Cape), Pseudoselago (29; Northern, Western and Eastern Cape), Zaluzianskya (c 60; southern Africa, Mount Elgon at the Kenya/Uganda border), 'Phyllopodium' (c 23; Namibia, Northern, Western and Eastern Cape, KwaZulu-Natal; polyphyletic; incl. Polycarena?), Polycarena (17; Northern and Western Cape; in Phyllopodium?), Glumicalyx (6; G. apiculatus, G. flanaganii, G. goseloides, G. lesuticus, G. montanus, G. nutans; Drakensberg in Lesotho, Eastern Cape, KwaZulu-Natal and Free State), Strobilopsis (1; S. wrightii; Drakensberg in KwaZulu-Natal and Lesotho), Tetraselago (4; T. longituba, T. natalensis, T. nelsonii, T. wilmsii; KwaZulu-Natal, Northern Province, Mpumalanga, Swaziland), Melanospermum (1; M. transvaalense; northern South Africa to western Zimbabwe), Selago (c 190; tropical and southern Africa, Madagascar, with their highest diversity in Western and Eastern Cape), Cromidon (12; Namibia, Northern and Western Cape, Free State), Microdon (6; M. capitatus, M. dubius, M. nitidus, M. orbicularis, M. parviflorus, M. polygaloides; Northern and Western Cape), Globulariopsis (7; G. adpressa, G. montana, G. obtusiloba, G. pumila, G. stricta, G. tephrodes, G. wittebergensis; Western Cape), Gosela (1; G. eckloniana; Western Cape). – Cosmopolitan, with their highest diversity in the Cape Provinces in South Africa. Placentation often apical (in Limosella free central in apical part). Ovule often one per carpel. Radicula in Limosella ephemeral. Protein bodies in cell nuclei lamellar. – Limoselleae may be sister-group to Scrophularieae. Jamesbrittenia may be sister to the remaining Limoselleae.

Scrophularieae Dumort., Fl. Belg.: 35. 1827

6/c 570. Antherothamnus (1; A. pearsonii; southwestern Zimbabwe, southern Namibia, Botswana, northern South Africa); Verbascum (c 360; Europe, the Mediterranean, western Asia, mountains in Ethiopia and tropical East Africa), Rhabdotosperma (2; R. brevipedicellatum, R. scrophulariifolia; Cameroon, Congo, tropical East Africa, Ethiopia, the Arabian Peninsula), Nathaliella (1; N. alaica; Kyrgyzstan, western China), Scrophularia (c 200; Europe, Macaronesia, the Mediterranean, temperate Asia, the Himalayas and Tibet, North and tropical America). – Leaves in at least Scrophularia and Verbascum with (tanniniferous etc.) idioblasts. Stamens in Verbascum usually five. Protein bodies in cell nuclei usually lamellar (in Verbascum usually amorphous). – Scrophularieae are possibly sister-group to Limoselleae.

Phylogeny (simplified) of Scrophulariaceae from Bayesian and Jacknife analyses based on DNA sequence data (Kornhall 2004; Kornhall & Bremer 2004).

Genus incertae sedis

Peltanthera 1/1. Peltanthera (1; P. floribunda; Guatemala, Panamá, Ecuador, Peru, Bolivia). – Bisexual evergreen small tree or shrub. Phellogen? Vessel elements? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma? Sieve tube plastids S type? Nodes 3:3, trilacunar with three leaf traces. Crystals? Hairs branched moniliform. Leaves opposite, simple, with involute ptyxis. Petiole vascular bundle transection flattened annular; petiole with mid-vein bundles (and sometimes medullary bundles). Stipules and leaf sheath absent. Venation pinnate. Stomata? Cuticular wax crystalloids? Leaf margin serrate. Inflorescence axillary, thyrsoid. Flowers somewhat zygomorphic. Hypogyny. Sepals five, with valvate aestivation, connate at base. Petals five, with valvate aestivation, connate. Nectariferous disc intrastaminal. Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers basifixed, versatile?, tetrasporangiate, introrse?, longicidal, with confluent thecae. Tapetum? Staminodia absent. Pistil composed of two connate carpels. Ovary superior, bilocular. Style single, entire. Stigma bifid, type? Pistillodium absent. Placentation axile. Ovules numerous per carpel, anatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis? Fruit a loculicidal capsule. Aril absent. Exotestal cells? Endotestal cells? Perisperm not developed. Endosperm? Embryo? Cotyledons? Germination? n = ? DNA? Verbascosides and cornoside derivatives present. – Peltanthera is possibly allied to Buddleja, although this has to be investigated.

STILBACEAE Kunth

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Kunth, Handb. Bot.: 393. 1831 [’Stilbineae’], nom. cons.

Retziaceae (Bartl.) Choisy in Mém. Soc. Phys. Hist. Nat. Genève 6: 400. 11 Aug 1834; Stilbales Kunth in C. F. P. von Martius, Consp. Regn. Veg.: 19. Sep-Oct 1835 [’Stilbinae’]

Genera/species 12/c 38

Distribution Tropical and southern Africa, Madagascar, islands in the Indian Ocean, southwestern Arabian Peninsula, with their largest diversity in Western Cape in South Africa.

Fossils Unknown.

Habit Bisexual, evergreen trees or shrubs, or perennial herbs. Some species with lignotuber. Most species are xerophytes.

Vegetative anatomy Phellogen ab initio present immediately outside pericycle. Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements usually libriform fibres (sometimes fibre tracheids) with simple pits, usually septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma absent or very rare (paratracheal scanty vasicentric). Wood elements sometimes storied (Retzia). Sieve tube plastids S type? Nodes 1?:1?, unilacunar? with one? leaf trace. Silica absent. Calciumoxalate usually absent (rarely as acicular crystals, styloids or crystal sand).

Trichomes Hairs simple, sometimes capitate; glandular hairs sometimes present.

Leaves Usually verticillate (sometimes opposite, rarely alternate), simple, entire, coriaceous, often ericoid, often with revolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate or leaves one-veined. Stomata anomocytic. Cuticular wax crystalloids as rodlets or threads. Leaf margin usually entire (rarely slightly serrate).

Inflorescence Usually terminal spike, head or axillary panicle, thyrsoid, spike- or head-like (flowers sometimes solitary axillary). Floral prophylls (bracteoles) as long as sepals.

Flowers Usually zygomorphic (sometimes actinomorphic). Hypogyny. Sepals four or five (to seven), usually with valvate (sometimes imbricate) aestivation, persistent or caducous, usually connate into tubular, campanulate or bilabiate calyx (petals sometimes free); median sepal adaxial. Petals four or five (to seven), with imbricate and/or valvate (sometimes induplicate-valvate) aestivation, equal or unequal in size, connate into tubular or infundibuliform corolla. Nectariferous disc small, intrastaminal, or absent. Oil-secreting glandular hairs often present.

Androecium Stamens four or five (to seven), as many as or one (adaxial median) fewer than sepals (adaxial median stamen rarely reduced and staminodial), haplostemonous, antesepalous, alternipetalous. Filaments filiform, free from each other, adnate to corolla tube (epipetalous). Anthers with thecae often confluent at apex or parallel (with seprate slits), dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodium one or absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, perforate, with various sculpturing.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular, with one locule often sterile, or unilocular (septum absent) or bilocular in lower part (septum incomplete) and unilocular in apical part. Style single, filiform, simple or somewhat bifid at apex, sometimes persistent. Stigma capitate, punctate or slightly bilobate, type? Pistillodium absent.

Ovules Placentation usually axile or basal (in Nuxia axile-peltate; in Thesmophora apical, with two transversely arranged collateral carpels each containing one descending ovule, abaxial carpel with secondary septum?). Ovules one or two ascending-apotropous and/or pendulous per carpel (when ovary unilocular) or numerous per carpel (when ovary bilocular), anatropous, apotropous or epitropous, unitegmic, tenuinucellar. Integument several cell layers thick. Hypostase well developed at base of megagametophyte. Megagametophyte monosporous, Polygonum type. Two antipodal cells developing (at least in Retzia). Endosperm development probably cellular. Endosperm haustoria chalazal and strongly micropylar (Retzia). Embryogenesis?

Fruit A usually loculicidal (sometimes also septicidal) capsule, with calyx and corolla persistent (sometimes a nutlet). Placenta sometimes with cushion-shaped scars of detached seeds.

Seeds Aril? Seed pedestals present. Exotesta? Endotesta? Perisperm not developed. Endosperm copious, usually oily (sometimes starchy). Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = 10, 12, 19 – Protein bodies in cell nuclei crystalline (Halleria) or absent.

DNA

Phytochemistry Insufficiently known. Route II iridoids (decarboxylated and C8-iridoids), Group IV carbocyclic iridoids, C8-iridoid glycosides (e.g. unedoside, stilbericoside, capensioside, holmioside, s-deoxyholmioside, and thunbergioside; extremely rare in other angiosperms), and verbascoside (a C10-iridoid glycoside) present.

Use Ornamental plants.

Systematics Charadrophila (1; C. capensis; around Stellenbosch and on Hangklip Peninsula in Western Cape), Halleria (4; H. elliptica, H. ligustrifolia, H. lucida, H. ovata; Ethiopia to Angola and South Africa, Madagascar, Yemen); Ixianthes (1; I. retzioides; Western Cape), Anastrabe (1; A. integerrima; Mozambique, KwaZulu-Natal, Eastern Cape), Bowkeria (3; B. citrina, B. cymosa, B. verticillata; South Africa, Swaziland, Lesotho); Nuxia (c 15; tropical Africa, Madagascar, the Comoros, the Mascarene Islands, southwestern Arabian Peninsula), Euthystachys (1; E. abbreviata; central Western Cape), ‘Stilbe’ (7; S. albiflora, S. ericoides, S. gymnopharyngia, S. overbergensis, S. rupestris, S. serrulata, S. vestita; Western Cape; non-monophyletic; incl. Campylostachys?), Campylostachys (1; C. cernua; Western Cape; in Stilbe?), Retzia (1; R. capensis; mountains in Western Cape), Kogelbergia (2; K. phylicoides, K. verticillata; southwestern Western Cape), Thesmophora (1; T. scopulosa; Ceres Mountains in Western Cape).

Stilbaceae are sister-group to a clade comprising the two large subclades [Lamiaceae+[Mazaceae+[Phrymaceae+[Paulowniaceae+[Rehmanniaceae+Orobanchaceae]]]]] and [Thomandersia to Lentibulariaceae].

Most asterids have C10- or C11-iridoid glucosides, wheras C8-iridoid glucosides (C8-iridoids) are found in Nuxia, Retzia and Stilbe (Stilbaceae), Thunbergia (Acanthaceae), and Arbutus/Arctostaphylos (Ericaceae). Unedoside is present in some genera of Stilbaceae, as well as in Loasaceae and Hydrangeaceae (Loasales).

Cladogram of Stilbaceae based on DNA sequence data (Kornhall 2004).

TETRACHONDRACEAE Wettst.

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Wettstein, Handb. Syst. Bot., ed. 3, (2:) 807. Jun-Sep 1924

Polypremaceae Reveal in Kew Bull. 66: 47. Mar 2011

Genera/species 2/3

Distribution New Zealand, southeastern United States to northern South America, the West Indies, Paraguay?, southern Patagonia, Tierra del Fuego,

Fossils Unknown.

Habit Bisexual, perennial or annual herbs, prostrate to erect. Aquatic or helophytic. Succulent (Tetrachondra). Young stems in Polypremum quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio superficial (Polypremum). Lateral vascular bundles split at nodes. Vessel elements with simple? perforation plattor; lateral pits? Imperforate tracheary xylem elements fibre tracheids (Polypremum) with simple? pits. Wood rays absent? Axial parenchyma? Sieve tube plastids S type? Nodes ?:?, split laterals. Etheral oils present in Tetrachondra. Crystals?

Trichomes Hairs unicellular or multicellular, in Polypremum moniliform; glandular hairs absent.

Leaves Opposite, simple, entire, coriaceous (Tetrachondra), with ? ptyxis. Stipules and leaf sheath absent. Petiole bases fused or connected by membranous interstipular sheathing structure. Petiole vascular bundles? Venation pinnate. Stomata anomocytic? Cuticular wax crystalloids? Lamina covered by sunken multicellular hairs consisting of basal cell, stalk cell and terminal head of at least four cells; lamina in Tetrachondra with numerous aromatic glands. Leaf margin serrate? or entire.

Inflorescence Terminal or axillary, few-flowered cymose (Polypremum), or flowers paired or solitary, axillary (Tetrachondra). Two or more pairs of floral prophylls (bracteoles) present.

Flowers Actinomorphic, small. Hypogyny (Tetrachondra) or partial epigyny (Polypremum). Sepals usually four (in Polypremum sometimes five), with valvate aestivation, persistent, connate in lower part into campanulate calyx; median sepal abaxial (Tetrachondra); sepals (Polypremum) diagonally arranged/initiated. Petals usually four (in Polypremum sometimes five), with imbricate aestivation, connate in lower part, orthogonally arranged/initiated (Polypremum). Nectary absent. Disc absent (at least in Polypremum).

Androecium Stamens usually four (in Polypremum sometimes five), haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Placentoid? Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate (6-sulcate?), shed as tetrads, bicellular? at dispersal. Exine tectate, with columellate infratectum, microperforate, psilate.

Gynoecium Pistil composed of two connate carpels (in Tetrachondra fused only in lower parts), transversely orientated. Ovary superior or semi-inferior, bilocular (Polypremum), or secondarily quadrilocular by false septa due to ingrowth of ovary wall (Tetrachondra). Style single, simple, gynobasic (Tetrachondra) or absent to minute (Polypremum). Stigma capitate or sometimes slightly bilobate (Polypremum), or small and somewhat truncate (Tetrachondra), type? Pistillodium absent.

Ovules Placentation basal (Tetrachondra), or basal to axile with ovules on peltate placentae inserted at base of septum (Polypremum). Ovules two ascending (Tetrachondra) or numerous (Polypremum) per carpel, anatropous, unitegmic, tenuinucellar. Integument three or four cell layers thick (Polypremum). Endothelium present. Megagametophyte monosporous, Polygonum type. Micropylar end of megagametophyte penetrating megasporangial epidermis to ovule surface; micropylar end not surrounded by integument (Polypremum). Endosperm development cellular (Polypremum). Endosperm haustorium micropylar or chalazal (Polypremum). Embryogenesis onagrad (Polypremum).

Fruit A schizocarp divided by secondary septum and with four single-seeded nutlike mericarps (Tetrachondra), or a loculicidal (and sometimes septicidal) many-seeded capsule (Polypremum), with green persistent calyx.

Seeds Aril? Seed pedestals present. Testa thin. Endothelial cells with persistent thickened inner walls. Perisperm not developed. Endosperm copious, oily. Embryo straight (Polypremum), well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = 10, 11 (Polypremum) – Protein bodies in nucleus?

DNA

Phytochemistry Insufficiently known. Caffeic acid esters (cornosides, verbascosides), scutellarin, essential oils, and sorbitol present. Iridoids not found. Shikimic acid derived anthraquinones?

Use Unknown.

Systematics Tetrachondra (2; T. hamiltonii: southern New Zealand; T. patagonica: southern Patagonia, Tierra del Fuego), Polypremum (1; P. procumbens; southeastern United States, Mexico, Central America, the West Indies, northern South America, Paraguay?).

Tetrachondraceae are sister to the remaining Plantaginales except Plocospermataceae and the [Oleaceae+Carlemanniaceae] clade.

THOMANDERSIACEAE Sreem.

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Sreemadhavan in Phytologia 37: 412. 22 Oct 1977

Genera/species 1/6

Distribution Tropical West and Central Africa.

Fossils Unknown.

Habit Bisexual, evergreen shrubs or small trees (sometimes lianas). Stem and branches not articulated.

Vegetative anatomy Phellogen? Secondary phloem stratified into hard fibrous and soft parenchymatous layers. Vessel elements with simple? perforation plates; lateral pits? Vestured pits? Imperforate tracheary xylem elements ? Wood rays? Axial parenchyma paratracheal? Pericyclic fibres short?, massively thickened. Sieve tube plastids S type? Nodes 1:3, unilacunar with three leaf traces. Cystoliths and acicular crystals absent.

Trichomes Hairs unicellular? or absent.

Leaves Opposite, anisophyllous, simple, usually entire (sometimes lobed), with ? ptyxis. Stipules and leaf sheath absent. Petiole swollen at base and apex. Petiole vascular bundle transection annular or incurved C-shaped. Venation pinnate, camptodromous. Stomata anisocytic. Cuticular wax crystalloids? Domatia in vein axils or absent. Leaf margin entire or serrate. Lamina with large and flat abaxial glands, blackening when dry. Extrafloral nectaries present.

Inflorescence Terminal or axillary, raceme-like cymose.

Flowers Zygomorphic. Hypogyny. Sepals five, persistent, connate, with imbricate? aestivation, on outer side between two abaxial calyx lobes with extranuptial vascularized multicellular nectaries (each one up to 3 mm wide and surrounded by swelling) consisting of layers or secretory cells. Petals five, connate into a bilabiate corolla (upper lip bilobate, lower lip trilobate), with irregular aestivation. Nectaries intrastaminal, vascularized by carpellary traces. Disc absent.

Androecium Stamens two longer and two shorter, haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodium one, adaxial-median.

Pollen grains Microsporogenesis simultaneous? Pollen grains penta- or hexacolpate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate? infratectum, punctate.

Gynoecium Pistil composed of two connate carpels. Gynoecial vascular tissue 8-shaped in cross-section. Ovary superior, bilocular, with nectariferous tissue at base. Style single, simple. Stigma cylindrical or bilobate, non-papillate?, Dry type? Pistillodium absent.

Ovules Placentation axile, with expanded placentae. Ovules one to three per carpel, hemianatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Hypostase present at chalazal end. Megagametophyte monosporous, Polygonum type. Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Fruit A woody loculicidal, non-explosive, capsule with persistent and accrescent calyx. Each seed situated on flattened elongate hook-shaped lignified ejaculator, retinaculum, expanded modified funicle.

Seeds Aril absent. Hilum large. Seed coat with ascending-imbricate scales or spirally arranged warts (with up to six cell layers in warts). Exotesta palisade, non-lignified. Endotesta? Perisperm not developed. Endosperm absent. Embryo large?, strongly curved, without chlorophyll? Cotyledons two, thin-foliaceous, complexly folded. Germination phanerocotylar?

Cytology n = ?

DNA

Phytochemistry Insufficiently known. Terpenoids (e.g. thomandertriole) and 2-indolinone alkaloids (e.g. thomandersine and isothomandersine) present. Iridoids?

Use Unknown.

Systematics Thomandersia (6; T. anachoreta, T. butayei, T. congolana, T. hensii, T. laurentii, T. laurifolia; southern Liberia and Côte d’Ivoire, southeastern Nigeria, southern Cameroon, Gabon, Equatorial Guinea, Congo Brazzaville, Congo Kinshasa, southern Central African Republic).

The sister-group relationship of Thomandersia is unsolved. The lineage is sometimes placed, with weak support, as sister to Schlegeliaceae in DNA analyses. Other alternative sister-groups of Thomandersia are Bignoniaceae or Verbenaceae. On the other hand, the pollen morphology in Thomandersia resembles the one in Pedaliaceae.

Thomandersia has a large complex calyx nectary, extended placentae, special corolla aestivation, and scaly spherical seeds without endosperm. Like the majority of Acanthaceae (Acanthoideae), Thomandersia has lignified retinacula (this may be a parallelism), but contrary to Acanthaceae they lack an explosion capsule. Retinacula are wide and flat in Thomandersia, whereas in Acanthaceae they are narrower and U-shaped in cross-section (Wortley & al.2005). In Thomandersia the retinaculum is compressed between the seed and the carpellary wall; in Acanthaceae the seeds are usually discoid and the retinaculum, which is present along one of the edges of the seed, is formed like a channel. Whereas the retinacula in Acanthaceae probably have an important role in connection with seed dispersal from the explosion capsule, the role of the retinacula in Thomandersia rather seems to be keeping the seed inside the capsule after the dehiscence (Wortley & al.2005).

VERBENACEAE J. St.-Hil.

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Saint-Hilaire, Expos. Fam. Nat. vol. 1(2): 245. Feb-Apr 1805, nom. cons.

Lantanaceae Martinov, Tekhno-Bot. Slovar: 357. 3 Aug 1820 [’Lantanae’]; Verbenales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 245. Jan-Apr 1820 [‘Verbenaceae’]; Durantaceae J. Agardh, Theoria Syst. Plant.: 295. Apr-Sep 1858; Petreaceae J. Agardh, Theoria Syst. Plant.: 364. Apr-Sep 1858 [’Petraeaceae’]; Verbenineae Engl., Syllabus, ed. 2: 177. Mai 1898.

Genera/species 33–35/1.120–1.190

Distribution Southeastern North America, Mexico, the West Indies, southern Brazil to Argentina, Atlantic islands, Europe, northern and northeastern Africa, Southwest and Central Asia, Taiwan, the Korean Peninsula, Japan.

Fossils Pollen grains are reported from Neogene strata.

Habit Usually bisexual (some species of Citharexylum dioecious), evergreen or deciduous trees, shrubs or lianas, perennial or annual herbs (Pitraea tuberous perennial). Stem and branches often quadrangular in cross-section. Often aromatic.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements often very long libriform fibres with simple pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (sometimes homocellular). Axial parenchyma paratracheal scanty vasicentric, aliform, lozenge-aliform, winged-aliform, confluent, or banded, or absent. Sieve tube plastids S type. Nodes 1:≥1, unilacunar with one or several leaf traces. Heartwood with resins. Calciumoxalate sometimes as styloids and acicular, stellate or cubical crystals.

Trichomes Eglandular hairs unicellular or multicellular, uniseriate or branched, often dendritic, sometimes capitate, hook-tipped, furcate or stellate (sometimes calcified/silicified); glandular hairs capitate, peltate or lepidote, often secreting resin.

Leaves Usually opposite (rarely verticillate or alternate), simple or pinnately compound, entire or lobed (sometimes scale-like), with imbricate? ptyxis. Stipules and leaf sheath absent. Petiole bases fused by line across node. Petiole vascular bundle transection arcuate; petiole sometimes also with medullary bundles, associated with median bundle. Venation pinnate. Stomata diacytic or anomocytic. Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts (sometimes with groups of brachysclereids). Acicular (sometimes stellate or cubical) crystals abundant. Leaf margin serrate, crenate, lobate or entire. Extrafloral nectaries present in many species on petiole and abaxial side of lamina.

Inflorescence Terminal or axillary, raceme-, spike- or head-like, simple or compound cymose; sometimes pseudanthium with involucre consisting of often large and sometimes petaloid bracts.

Flowers More or less zygomorphic. Hypogyny. Sepals (four or) five, with usually imbricate quincuncial aestivation, campanulate, persistent, connate; median sepal adaxial. Petals (four or) five, with usually imbricate quincuncial aestivation, connate into an infundibuliform or hypocrateriform corolla (sometimes bilabiate). Nectariferous disc intrastaminal, annular.

Androecium Stamens usually two longer and two shorter (didynamous; in Hierobotana and Stachytarpheta two fertile stamens; adaxial stamen sometimes staminodial; fertile stamens in Verbena five), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, usually adnate to corolla tube (epipetalous). Anthers basifixed or dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective sometimes shrinked or expanded, occasionally with glandular appendage. Tapetum secretory, with binucleate to quadrinucleate cells. Staminodia usually absent (sometimes one or two).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–5)-colporate (in Petrea and Stachytarpheta tricolpate; in Citharexylum sometimes 3(–4)-porate), shed as monads, bicellular or tricellular? at dispersal. Exine tectate to semitectate, with columellate infratectum, perforate to reticulate, rugulate-reticulate, psilate, spinulate or echinate (rarely verrucate).

Gynoecium Pistil composed of one or two connate, collateral, transverse carpels (often gradually bipartite by in-growing secondary septa; one carpel often aborted resulting in unicarpellate, pseudomonomerous, ovary; carpels in Duranta four). Ovary superior, bilocular or secondarily quadrilocular (by inflexion of carpellary margins; in Duranta secondarily octalocular; in Bouchea pseudomonomerous, with one carpel modified into carpophore). Style single, simple or bilobate, often persistent. Stigma usually capitate (rarely somewhat quadrangular), with asymmetrically swollen stigmatoid tissue and glandular surface, papillate, Wet type. Pistillodium absent.

Ovules Placentation usually basal or axile (rarely parietal), attached directly to margins of secondary carpellary septa (cf. Lamiaceae). Ovules usually two (ovule rarely one) per carpel, usually anatropous (rarely hemianatropous or orthotropous), usually ascending (rarely pendulous), apotropous, unitegmic, tenuinucellar. Integument five to nine cell layers thick. Obturator present. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent, sometimes multinucleate. Endosperm development ab initio cellular. Endosperm haustorium chalazal and/or micropylar. Embryogenesis onagrad.

Fruit A dry or fleshy schizocarp with two or four one-seeded mericarps or two two-seeded mericarps, or a drupe with one, two or four pyrenes (bilocular mericarps or pyrenes may contain ovules from both carpels) with persistent calyx. Fruits occasionally winged.

Seeds Aril absent. Testa thin-walled. Exotesta present. Endotesta? Perisperm not developed. Endosperm usually absent (present in Lantaneae). Embryo straight, oily, without chlorophyll. Cotyledons two, spatulate. Germination phanerocotylar or cryptocotylar?

Cytology n = 5–12, 17–20 (sometimes more) – Polyploidy occurring.

DNA Deletion the plastid gene matK.

Phytochemistry 6- or 8-hydroxyflavones or 6-methoxyflavones and 6-methoxyflavone glycosides, Route-I-iridoids (some species of Verbena), Route II decarboxylated iridoids (4-carboxyiridoids), Group II carbocyclic iridoids (lamiide), Group III carbocyclic iridoids (cornine, hastatoside), iridoid glycosides, ursolic acid, caffeic acid esters (caffeoyl phenylethanoid glucosides, e.g. verbascoside, orobanchin; cornoside present in Phyla), naphthoquinones, toxic triterpene esters, and triterpene saponins present. Ethereal oils present in Lantaneae. Flavonols, ellagic acid, proanthocyanidins, and cyanogenic compounds not found. Carbohydrates usually stored as stachyose and other oligosaccharides.

Use Ornamental plants, medicinal plants, timber, carpentries (e.g. Citharexylum).

Systematics

The sister-group relationships of Verbenaceae are not clarified.

A probable topology is the following (Thode & al. 2013): [Petreeae+[Duranteae+[[Casselieae+Citherexyleae]+[Priveae+[Rhaphithamnus+[Neospartoneae+[Verbeneae+Lantaneae]]]]]]].

Petreeae Briq. in Engler et Prantl, Nat. Pflanzenfam. IV, 3a: 144, 157. 26 Feb 1895 [‘Petraeeae’]

2/12. Petrea (11; Mexico, Central America), Xolocotzia (1; X. asperifolia; Mexico). – Mexico to Amazonian Brazil. Petrea consists of lianas, whereas Xolocotzia is a shrub or small tree. Fruit a drupe consisting of two pyrenes derived from secondarily unicarpellate ovary (one carpel aborted). Calyx large, showy, exceeding corollas. – Petreeae are sister to the remaining Verbenaceae. Petrea has been identified as sister to Bignoniaceae in some previous analyses.

[Duranteae+[[Casselieae+Citherexyleae]+[Priveae+[Rhaphithamnus+[Neospartoneae+[Verbeneae+Lantaneae]]]]]]

Duranteae Benth. in Ann. Mag. Nat. Hist., ser. 1, 2: 448. Feb 1839

5/195–205. Duranta (c 20; the West Indies, South America), Bouchea (18; Texas, Mexico, Central America, tropical South America, one species, B. pterygocarpa, in Ethiopia), Chascanum (27–32; Africa, Madagascar, the Arabian Peninsula to western India), Recordia (1; R. boliviana; Bolivia), Stachytarpheta (120–125; Florida, Mexico, Central America, the West Indies, tropical South America). – Africa, Madagascar, southwestern Asia to India, southeastern United States to Argentina. Trees, shrubs or herbs. Eglandular hairs multicellular. Inflorescence terminal spike or compound raceme with terminal and axillary flowering shoots. Two floral prophylls (bracteoles) present in Bouchea, Recordia, species of Duranta and species of Chascanum. Two fertile stamens present in Stachytarpheta. Carpels in Duranta four. Ovary in Duranta secondarily octalocular, in Bouchea pseudomonomerous, with one carpel modified into carpophore. Calyx usually persistent and enclosing fruit (not in Bouchea). – Duranteae are sister to the remaining Verbenaceae except Petreeae.

[[Casselieae+Citherexyleae]+[Priveae+[Rhaphithamnus+[Neospartoneae+[Verbeneae+Lantaneae]]]]]

[Casselieae+Citherexyleae]

Casselieae (Schauer) Troncoso in Darwiniana 18: 385. Mar 1974

3/19. Casselia (11; tropical South America), Parodianthus (2; P. capillaris, P. ilicifolius; Argentina), Tamonea (6; T. boxiana, T. curassavica, T. euphrasiifolia, T. juncea, T. spicata, T. subbiflora; southern Mexico, Central America, tropical South America). – Mexico, Central America and the West Indies to Argentina. Inflorescence consisting of lateral racemes. Pistil composed of a single carpel. Secondarily bicarpellate ovaries with secondary septa fused to carpel walls only at base and apex. Ovule inserted in upper part of locule, attached to margins. Placental line broad; placental bundles entering ovules in upper part of locule. – Casselieae are sister to Citharexyleae.

Citharexyleae Briq. in Engler et Prantl, Nat. Pflanzenfam. IV, 3a: 144, 158. 26 Feb 1895

2–3/100–105. Citharexylum (100–105; Florida, Mexico, Central America, the West Indies, tropical South America to Argentina; incl. Verbenoxylum?), Verbenoxylum (1; V. retizii; Brazil; in Citharexylum?), Rehdera (2; R. penninervia, R. trinervis; Mexico, Central America). – Tropical America to Argentina. Staminodium single. Pollen grains in Citharexylum sometimes 3(–4)-porate. Bicarpellate ovary usually developing into a drupaceous to subdrupaceous fruit with two two-seeded mericarps, with minute often deciduous floral bracts and shortly pedicellate flowers.

[Priveae+[Rhaphithamnus+[Neospartoneae+[Verbeneae+Lantaneae]]]]

Priveae Briq. in Engler et Prantl, Nat. Pflanzenfam. IV, 3a: 144, 155. 26 Feb 1895

1–2/c 20. Priva (c 20; tropical and subtropical regions in Africa, southern Asia and southwestern United States to northern Argentina; incl. Pitraea?), Pitraea (1; P. cuneato-ovata; temperate South America; in Priva?). – Perennial herbs. Rhachis, stem, leaves and peduncles often with uncinate hairs. Fruit a dry schizocarp splitting into two two-seeded mericarps derived from a bicarpellate ovary.

[Rhaphithamnus+[Neospartoneae+[Lantaneae+ Verbeneae]]]

Rhaphithamnus clade

1/2. Rhaphithamnus (2; R. spinosus, R. venustus; Chile, Juan Fernandez, Argentina). – Spinescent shrubs. Inflorescence an axillary, one- to five-flowered raceme. Corolla tube long. Ovary bicarpellate. Fruit drupaceous.

[Neospartoneae+[Lantaneae+Verbeneae]]

Neospartoneae Olmstead et N. O’Leary in Amer. J. Bot. 97: 1653. 1 Oct 2010

3/10. Neosparton (4; N. aphyllum, N. darwinii, N. ephedroides, N. patagonicum; temperate regions in Chile and Argentina), Lampayo (5; L. aratae, L. castellani, L. hieronymi, L. officinalis, L. schickendantzii; arid regions in Bolivia, Chile and Argentina), Diostea (1; D. juncea; Chile, Argentina). – Bolivia, Chile, Argentina. Shrubs (sometimes Ephedra-like). Hairs absent. Inflorescence a usually terminal spike. Corolla tube much longer than calyx. Staminodium sometimes present. Carpel one. Fruit a drupe with two-seeded pyrene.

[Lantaneae+Verbeneae]

Staminodia absent.

Lantaneae Endl., Gen. Plant.: 635. Aug 1838

10/415–420. Coelocarpum (7; C. africanum, C. glandulosum, C. haggierense, C. humbertii, C. madagascariense, C. socotranum, C. swinglei; Somalia, Madagascar, Socotra); ‘Acantholippia’ (6; A. deserticola, A. riojana, A. salsoloides, A. seriphioides, A. tarapacana, A. trifida; arid regions in South America; polyphyletic), ‘Aloysia’ (30–35; United States, Mexico, Central America, South America; non-monophyletic), Xeroaloysia (1; X. ovalifolia; northwestern Argentina; in Aloysia),‘Lippia’ (c 200; tropical Africa, tropical America; paraphyletic), Phyla (c 11; tropical and subtropical regions on both hemispheres), Diphyllocalyx (7; D. armata, D. cayensis, D. galanus, D. myrtifolia, D. nipensis, D. urquiolae, D. variifolia; Cuba), Isidroa (1; I. spinifera; Hispaniola), Nashia (1; N. inaguensis; Puerto Rico, St Croix in the Virgin Islands),‘Lantana’ (c 150; tropical and southern Africa, southern Mexico, Central America, the West Indies, tropical South America; polyphyletic). – Tropical and subtropical Africa, Madagascar, Socotra, tropical and subtropical Asia, America. Stomata anisocytic. Inflorescence capitate. Ovary usually secondarily unicarpellate (one carpel aborted; in Coelocarpum bicarpellate). Ovule in Lantana one per carpel. Fruit a two-seeded drupe or a dry schizocarp splitting into two one-seeded mericarps. Fruit in Coelocarpum a fleshy drupe splitting into two two-seeded mericarps. Calyx persistent and more or less enclosing fruit. Endosperm present in Lantana. Ethereal oils present. – Lantaneae are sister to Verbeneae. Coelocarpum may be more allied to the Verbeneae or sister to [Lantaneae+Verbeneae] (Lu-Irving & Olmstead 2013). ‘Lantana’ is nested inside ‘Lippia’, according to Lu-Irving & Olmstead (2013).

Verbeneae Dumort., Anal. Fam. Plant.: 22. 1829

6/350–400. Dipyrena (1; D. glaberrima; Argentina); Mulguraea (11; arid regions in Peru, Chile and Argentina), ‘Verbena’ (200–250; temperate and subtropical regions in North and South America, 2–3 species in the Old World; non-monophyletic; incl. Hierobotana?), Hierobotana (1; H. inflata; Colombia, Ecuador, Peru; in Verbena?), ‘Junellia’ (c 40; South America; non-monophyletic), ‘Glandularia’ (c 95; temperate to tropical America; polyphyletic; in Junellia?). – North and South America, a few species in northern Africa, Europe and Asia. Fertile stamens in Verbena five, in Hierobotana two. Ovary bicarpellate. Fruit a dry schizocarp splitting into four single-seeded mericarps. – Dipyrena (leaves alternate; ovary bicarpellate and developing into a subdrupaceous fruit subdivided into two two-seeded mericarps) is sister to the remaining Verbeneae. Hierobotana may be included in Verbena. The following topology is thus possible: [Dipyrena+[Mulguraea+[Junellia+[Verbena+Glandularia]]]]. A number of species of ‘Glandularia’ are nested in Verbena, according to analyses using plastid DNA data (O’Leary & al. 2009).

Cladogram (simplified) of Verbenaceae based on DNA sequence data (Marx & al. 2010).


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