DIPSACALES Dumortier Main Tree, Synapomorphies

Habit Usually bisexual (rarely polygamomonoecious or dioecious), annual, biennial or perennial herbs, evergreen or deciduous trees or shrubs, lianas (sometimes suffrutices, rarely woody and creeping).

Vegetative anatomy Phellogen ab initio superficial or pericyclic. Pericyclic fibres absent. Secondary lateral growth normal or absent. Vessel elements with simple or scalariform (sometimes reticulate or foraminate) perforation plates; lateral pits alternate, scalariform or opposite, usually bordered (sometimes simple) pits. Imperforate tracheary xylem elements tracheids, libriform fibres or fibre tracheids with simple and/or bordered pits, septate or non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular, or absent. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, confluent, vasicentric, unilateral, banded, or absent. Wood elements sometimes partially storied. Tile cells often present. Intraxylary phloem sometimes present. Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, unilacunar with one trace, or 5:5, pentalacunar with five traces). Oil cells and/or mucilage cells frequent. Prismatic or rhomboidal calciumoxalate crystals or crystal sand often frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or branched (often stellate or peltate, rarely hair tufts); glandular hairs often present.

Leaves Usually opposite (rarely verticillate or alternate), simple or compound (pinnately compound or trifoliolate), entire or lobed, with involute, supervolute or conduplicate (rarely plicate) ptyxis. Stipules usually absent (stipitate extrafloral nectaries sometimes present in stipular position); leaf sheath absent. Petiole vascular bundle transection arcuate, petiole sometimes also with adaxial inverted bundles (sometimes closed C-shaped and petiole with wing bundles). Foliaceous to scale-like appendages or nectariferous glands sometimes present on petiole or leaf base. Venation pinnate or palmate. Stomata usually anomocytic (sometimes anisocytic or paracytic). Cuticular wax crystalloids as rodlets or tubuli, or absent. Domatia usually absent. Leaf margin and leaflet margins serrate (sometimes glandular serrate) or entire; glandular leaf teeth with main vein and two accessory veins, or one accessory vein proceeding above tooth.

Inflorescence Terminal or axillary, dichasial, cymose, head-, spike-, raceme- or umbel-like, corymbose, paniculate, or thyrsoid (flowers often paired, sometimes solitary axillary), sometimes pseudanthia with large sterile marginal flowers. Supernumerary bracts often present (number of bracts larger than number of flowers due to floral abortions, forming epicalyx).

Flowers Actinomorphic or zygomorphic (sometimes asymmetric). Epicalyx (consisting of supernumerary bracts) sometimes present. Usually epigyny (sometimes partial epigyny). Sepals (two or) three to five, with imbricate or open aestivation, small, usually persistent, usually connate; median sepal adaxial. Petals (three or) four or five (or six), with imbricate or valvate aestivation, caducous, connate into a disc-shaped, infundibuliform, campanulate or tubular and bilabiate corolla. Unicellular or multicellular nectariferous hairs or glands often present near corolla lobe bases (sometimes at stylar base). Disc usually absent (sometimes intrastaminal).

Androecium Stamens (one to) three to five, more or less bifid, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals, or epipetalous. Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely split into two disporangiate halves), extrorse or introrse, longicidal (dehiscing by longitudinal slits). Tapetum usually amoeboid-periplasmodial (sometimes secretory). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually (2–)3(–4)-colporate (sometimes colpate or porate, rarely pororate), shed as monads, usually tricellular (rarely bicellular) at dispersal. Exine tectate or semitectate, with columellate (rarely acolumellate) infratectum, perforate, reticulate or microreticulate, spinulate (sometimes echinate or psilate, rarely intectate).

Gynoecium Pistil composed of two to five (to eight) connate carpels, one or several of which often sterile and degenerating. Ovary usually inferior (sometimes semi-inferior), (unilocular or) bilocular to quinquelocular; ovaries often more or less connate. Style single, simple, or stylodia three to five, free (sometimes absent). Stigmas entire or lobate, papillate or non-papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile to apical. Ovules one to numerous per carpel, anatropous, pendulous, apotropous or epitropous, unitegmic, usually tenuinucellar (sometimes crassinucellar through multiple divisions of parietal cell). Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, or tetrasporous, Adoxa type). Endosperm development ab initio cellular. Endosperm haustoria sometimes present. Embryogenesis asterad, solanad or piperad.

Fruit A berry-like drupe, many-seeded berry, drupaceous syncarp, septicidal capsule or cypseloid nut, often with persistent calyx and/or epicalyx. Endocarp often with two layers of different cell types, with fibrous layers perpendicular to each other. Endocarp sclereids thick-walled, crystalliferous.

Seeds Aril absent. Testa sometimes vascularized, sometimes crushed. Exotestal cells often palisade, often enlarged, cuboud to rectangular, often lignified and with theoidal thickenings. Endotesta? Perisperm not developed. Endosperm usually copious (sometimes sparse or absent), often with thin-walled undifferentiated cells, rich in fatty oils, sometimes ruminate. Embryo large or small, straight, well differentiated, with or without chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA Several CYCLOIDEA and DIVARICATA genes duplicated (duplications possibly correlated with evolution of zygomorphy, according to Howarth & Donoghue 2005, 2009, and Howarth & al. 2011). Mitochondrial intron coxII.i3 often absent.

Phytochemistry Flavonols (kaempferol, quercetin), biflavonoids, cyanidin, Route I secoiridoids, Group V carbocyclic iridoids (valtrate, villoside, patrinoside), Group VI secoiridoids (secologanin, morroniside, kingiside), Group VII secoiridoids (sweroside, gentiopicroside), Group X secoiridoids (e.g. loganin, adoxoside, iridoid pyridine alkaloids), iridoid glucosides (i.a. dimethyl-acetal of cantleyoside), valepotriates (triesters of Route I secoiridoids), oleanolic acid derivatives, monoterpenoids, sesquiterpenoids (foetid ethereal oils), ursolic acid, caffeic acid, chlorogenic acid, indole alkaloids, triterpene saponins, cyanogenic glycosides (holocalin, sambunigrin, zierin), phenylalanine- or tyrosine-derived cyanogenic compounds, ethereal oils (rare), arbutin, eleostearic acid, valerianic acid, davidigenin, and actinidin present. Ellagic acid not found.

The clade [Diervillaceae+[Caprifoliaceae+[Linnaeaceae+[Morinaceae+[Dipsacaceae+ Valerianaceae]]]]], in other words Dipsacales except Adoxaceae, is supported by the potential synapomorphies: bark papery-flaky; phellogen deeply seated; pericyclic fibres moderately developed; wood often fluorescent; glandular foliar tooth with mid-vein and two lateral veins or lateral vein running above tooth; corolla zygomorphic, often two-lipped, tubular; epigyny; nectary as densely spaced adaxial unicellular or multicellular glandular hairs near base of corolla; tapetum amoeboid-periplasmodial; pollen grains large, spheroidal; tectum echinate; lateral and dorsal carpellary bundles absent, ventral bundles in axis; style long; stigma capitate, Wet type; chromosomes small; duplication of nuclear genes dipsCYC2 and dipsCYC3; horizontal transfer of mitochondral gene rps11; and presence of O-methylated flavonols and flavones (Stevens 2001 onwards).

The clade [Linnaeaceae+[Morinaceae+[Dipsacaceae+Valerianaceae]]] has the following potential synapomorphies, according to Stevens (2001 onwards): supernumerary bracts (bracts more numerous than flowers) due to numerous aborted flowers; stamens four, didynamous; pistil composed of three carpels, two of which aborted; fruit a cypsela; seed coat flattened; and exotesta not lignified.

The clade [Morinaceae+[Dipsacaceae+Valerianaceae]] is characterized by the synapomorphies: perennial herbs with taproot; presence of flank bridge in stem between lateral vascular bundles (two bundles on each side seem to form split lateral branches and one branch from each bundle fusing in middle, forming bridging bundles); bundles innervating petiole arise from bridging bundle (not in Morina); presence of libriform fibres; absence of bud scales; nectary present at corolla base; filaments glabrous; anthers dorsifixed; ovary with much reduced sterile loculi; placentation apical; ovule one per carpel; endothelium prominent, with crystalliferous layer; chalazal nuclei proliferating; exotesta not thickened; endosperm sparse; embryo large; plastid inverted repeat significantly expanded; presence of monoterpenoids, cathecolic tannins and alkaloids; and absence of ellagic acid.

Finally, Dipsacaceae and Valerianaceae share the following potential synapomorphies: fairly small flowers; uniform staminal length (stamens equal in length); tapetum with cells at least quadrinucleate; exine with prominent and branched infratectal columellae; stigma Dry type; megagametophyte with binucleate to quadrinucleate chalazal cells; embryo with chlorophyll; presence of saccharose; and absence of quercetin.

Sambucaceae Batsch ex Borkh., Bot. Wörterb. 2: 322. 1797 [‘Sambuci’]; Sambucales Bercht. et J. Presl, Přir. Rostlin: 258. Jan-Apr 1820 [‘Sambucinae’]; Tinaceae Martinov, Tekhno-Bot. Slovar: 635. 3 Aug 1820 [‘Tiniaceae’]; Viburnaceae Raf. in Ann. Gén. Sci. Phys. Bruxelles 6: 87. Oct-Dec 1820 [’Viburnidia’]; Viburnales Dumort., Anal. Fam. Plant.: 29. 1829 [’Viburnarieae’]; Adoxales Nakai in J. Jap. Bot. 24: 14. Dec 1949

Distribution Mainly temperate regions on the Northern Hemisphere; some species in subtropical regions, few species on tropical mountains, southeastern Australia and Tasmania, with their largest diversity in Himalayas and China.

Fossils Numerous Cenozoic fossilized pollen, fruits and seeds of Sambucus and Viburnum are known from Europe and North America. Endocarps of Viburnum are known from the Miocene onwards and seeds of Sambucus are recorded from Eocene and later strata. Pollen fossils of Viburnum are known from Eocene to Miocene layers in, e.g., North America and Iceland (Manchester & al. 2015).

Habit Usually bisexual (in Viburnum rarely polygamomonoecious), perennial herbs (Adoxa, Sinadoxa, Tetradoxa, some species of Sambucus), evergreen or deciduous trees or shrubs (Sambucus, Viburnum).

Vegetative anatomy Phellogen ab initio superficial. Pericyclic fibrous envelope poorly developed or absent. Secondary lateral growth normal or absent. Vessel elements usually with scalariform (in Sambucus simple) perforation plates; lateral pits alternate, scalariform or opposite, bordered pits. Imperforate tracheary xylem elements tracheids, libriform fibres (Sambucus) or fibre tracheids (Viburnum) with simple or bordered pits, septate or non-septate; tracheids absent in Adoxa, Sinadoxa and Tetradoxa. Vestured pits absent. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, confluent, vasicentric, unilateral, or banded. Wood elements in Sambucus partially storied. Tile cells present. Intraxylary phloem present in Sambucus. Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, unilacunar with one trace, or 5:5, pentalacunar with five traces). Coenocytic canals in internodes in Sambucus secreting tannins. Oil cells and/or mucilage cells frequent. Prismatic calciumoxalate crystals abundant. Crystal sand frequent in Sambucus. Calcium oxalate (crystal sand and druses) abundant in Sambucus and Viburnum.

Trichomes Hairs unicellular or multicellular, uniseriate or branched (in Viburnum often stellate or peltate); glandular hairs abundant. Hairs absent in Adoxa, Sinadoxa and Tetradoxa.

Leaves Usually opposite (rarely verticillate or alternate spiral), simple or compound (pinnately compound or trifoliolate), entire or lobed, usually with involute or conduplicate (rarely plicate) ptyxis. Stipules absent (Sambucus usually with stipitate extrafloral nectaries in stipular position); leaf sheath absent. Petiole vascular bundle transection arcuate (sometimes closed C-shaped); petiole also with adaxial inverted bundles (sometimes with wing bundles). In Sambucus and Viburnum foliaceous to scale-like appendages or nectariferous glands sometimes present on petiole or leaf base. Venation pinnate or palmate. Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids? Leaf margin and leaflet margins usually serrate. Mesophyll without mucilage cells.

Inflorescence Usually terminal (rarely axillary), dichasial, of various shape: in Adoxa capitate; in Sinadoxa spicate; in Tetradoxa raceme-like; in Sambucus and Viburnum umbel-like, corymbose or paniculate, thyrsoid; in Viburnum sometimes pseudanthium with large sterile marginal flowers.

Flowers Usually actinomorphic (sometimes slightly zygomorphic), small. Epigyny to partial epigyny. Sepals (two or) three to five, with open aestivation, very small, usually persistent, connate; median sepal adaxial. Petals (three to) five (or six), with imbricate or valvate aestivation, caducous, connate into disc-shaped, infundibuliform, campanulate or tubular corolla. Nectary in Viburnum epigynous disc-like structure on ovary (absent in Sambucus and some species of Viburnum). Nectaries in Adoxa, Sinadoxa and Tetradoxa multicellular nectariferous glands at corolla lobe bases. Disc absent.

Androecium Stamens three to five, more or less bifid (in Sinadoxa bifid to base), haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals or adnate to corolla tube (epipetalous, in Sambucus adnate to base of corolla tube). Anthers basifixed or dorsifixed (Viburnum), non-versatile, usually tetrasporangiate (in Adoxa, Sinadoxa and Tetradoxa more or less split into two disporangiate monothecal anther halves, in Tetradoxa peltate), extrorse (Adoxa, Sinadoxa, Tetradoxa, Sambucus) or introrse (Viburnum), longicidal (dehiscing by longitudinal slits). Tapetum usually secretory (at least in Adoxa and sometimes in Viburnum amoeboid-periplasmodial). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tricolpate or triporate), shed as monads, tricellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, or intectate (Viburnum).

Gynoecium Pistil composed of two to five connate carpels (often only one carpel developed; in Viburnum three, two of which usually sterile). Ovary inferior to semi-inferior (in Tetradoxa superior?), usually bilocular to quinquelocular (in Sinadoxa unilocular, in Viburnum unilocular to trilocular); ovary cells in Sambucus with calciumoxalate druses. Stylodia in Adoxa and Tetradoxa three to five, free; absent in Sinadoxa; in Sambucus and Viburnum very short or absent. Stigmas capitate or lobate, papillate, Dry or Wet type. Pistillodia absent.

Ovules Placentation axile to apical. Ovule one median per carpel, anatropous, pendulous, apotropous (Viburnum) or epitropous (Sambucus), unitegmic, usually tenuinucellar (in Viburnum crassinucellar by division of parietal cell). Integument ? cell layers thick. Parietal tissue present. Endothelium present (in Adoxa poorly developed). Megagametophyte usually tetrasporous, octanucleate, Adoxa type (in Viburnum monosporous, Polygonum type, disporous, Allium type, or tetrasporous, Adoxa type). Endosperm development cellular. Endosperm haustoria? Embryogenesis asterad.

Fruit A berry-like drupe with one (Viburnum) or three to five (Sambucus) single-seeded pyrenes, or a drupaceous syncarp (Adoxa, Sinadoxa, Tetradoxa), often with persistent calyx.

Seeds Aril absent. Seeds often ruminate. Exotestal cells often palisade, unlignified. Endotesta? Perisperm not developed. Endosperm copious, oily, in Viburnum with small amorphous crystals. Embryo large or small, straight, usually well differentiated, with (Sambucus) or without (Viburnum) chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = (8) 9 (10, 18) – Chromosomes large, continuously condensed from prophase to metaphase.

Phytochemistry O-methylated flavonols, Group VI secoiridoids present in Adoxa and Viburnum (secologanin, morroniside), Group X secoiridoids (in Viburnum adoxoside), Group III, V and VII secoiridoids present in Viburnum, alkaloids, caffeic acid, p-coumaric acid, sinapic acid, ferulic acid, ursolic acid (Sambucus), tannins, and cyanogenic compounds present. Ellagic acid and proanthocyanidins not found.

Use Ornamental plants, medicinal plants, liqueurs, wines, tea and jam (fruits of Sambucus nigra), timber.

Systematics Adoxaceae are sister to the remaining Dipsacales.

Viburnum is sister to the remaining Adoxaceae and Sambucus sister to the clade [Sinadoxa+[Adoxa+Tetradoxa]].

Viburnum

1/150–175. Viburnum (150–175; temperate regions on the Northern Hemisphere, tropical mountains; absent from Africa). – Usually shrubs (rarely trees). Hairs often stellate or peltate. Foliaceous to scale-like appendages or nectariferous glands sometimes present on petiole or leaf base. Nectary epigynous disc-like epithelial structure on ovary. Anthers dorsifixed, introrse. Pistil composed of three connate carpels, with odd carpel abaxial, and two carpels aborting. Ovary unilocular to trilocular. Ovule apotropous, crassinucellar by division of parietal cell. Parietal tissue approx. one cell layer thick. Fruit a single-seeded baccate drupe. Endocarp with fibrous layers orientated in parallel, with one or more sclereidal layers. Endosperm crystalliferous, with differentiated outer layer and thickened cell walls. – Viburnum clemensae on Borneo is obviously sister to the other species in Viburnum.

Adoxoideae Syme in Smith et Soweby, Engl. Bot., ed. 3(B), 4: 197. 1865 [‘Adoxeae’]

3–4/29–30. Sambucus (c 25; temperate regions on both hemisphere, few species in subtropical regions and tropical mountains in South America, southeastern Australia and Tasmania), Adoxa (2; A. moschatellina,A. xizangensis; temperate regions on the Northern Hemisphere south to the Himalayas and central United States; incl. Tetradoxa?), Tetradoxa (1; T. omeiensis; China; in Adoxa?), Sinadoxa (1; S. corydalifolia; China). – Temperate regions on the Northern Hemisphere, few species in subtropical regions and on tropical mountains. Shrubs or herbs. Cambium in Sambucus storied. Vessel elements with simple perforations. Cortex and phloem in Sambucus with crystal sand. Nodes in Sambucus 1:1. Leaves usually compound (sometimes simple and deeply lobed). Sambucus usually with stipitate extrafloral nectaries in stipular position. Nectary epigynous, on corolla associated with multicellular hairs, or absent. Anthers extrorse (sometimes eight, monothecal). Pistil composed of one or five connate antepetalous fertile carpels. Stylodia free. Stigmas capitate. Megagametophyte tetrasporic, 8-nucleate, Adoxa type. Endocarp with three layers of various cell types: two fibrous cell layers and one sclereid cell layer. Embryo in Sambucus large.

Cladogram of Adoxaceae based on DNA sequence data (Winkworth & al. 2008).

Loniceraceae Vest, Anleit. Stud. Bot.: 272, 296. 1818 [’Loniceroideae’]; Caprifoliales Bercht. et J. Presl, Přir. Rostlin: 257. Jan-Apr 1820 [‘Caprifoliae’]; Caprifoliopsida Endl., Gen. Plant.: 520. Jun 1838 [’Caprifolia’]; Loniceropsida Brongn., Enum. Plant. Mus. Paris: xvii, 47. 12 Aug 1843 [’Loniceroideae’]; Lonicerales T. Liebe, Grund. Spec. Bot.: 71. 31 Jan-1 Feb 1866 [’Loniceroideae’]

Distribution Mainly temperate regions on the Northern Hemisphere, with their highest diversity in the Himalayas and China; few species in tropical mountains south to Mexico and West Malesia.

Fossils Pollen of Lonicera were reported from the Late Eocene of western North America.

Habit Usually bisexual (rarely polygamomonoecious), evergreen or deciduous shrubs or lianas (rarely trees; in Triosteum perennial rhizomatous herbs).

Vegetative anatomy Phellogen ab initio usually pericyclic (in Heptacodium in centre of cortex). Medulla in some species of Lonicera septated by diaphragms. Cortical fibres in Heptacodium well developed. Stem endodermis absent. Vessel elements with simple or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, scalariform or opposite, usually bordered (sometimes simple) pits. Imperforate tracheary xylem elements tracheids with simple and/or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric or unilateral, or absent. Tile cells? Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (rarely 1:1?, unilacunar with one? trace). Prismatic calciumoxalate crystals (druses) often abundant.

Trichomes Hairs unicellular or multicellular, usually simple (sometimes stellate); sometimes peltate-lepidote glandular hairs.

Leaves Usually opposite (rarely verticillate), simple, usually entire (in, e.g., Leycesteria and often in Lonicera pinnately lobed), usually with supervolute or conduplicate (in Heptacodium involute) ptyxis. Stipules usually absent (present in, e.g., Leycesteria); leaf sheath absent. Petiole vascular bundles? Venation camptodromous or brochidodromous, pinnate or palmate (lamina in Heptacodium three-veined from base). Stomata usually anomocytic (rarely paracytic). Cuticular wax crystalloids in Leycesteria, Lonicera and Symphoricarpos as tubuli (sparsely branched; in some species of Lonicera platelets), dominated by nonacosan-10-ol (absent in Heptacodium and Triosteum). Domatia usually absent (rarely as pits, pockets or hair tufts?). Leaf margin serrate or entire. Extrafloral nectaries present or absent.

Inflorescence Terminal or axillary, dichasial or corymbose, raceme-, spike- or head-like; flowers often paired (in Triosteum solitary axillary). Supernumerary bracts absent (cf. Linnaeaceae).

Flowers Usually zygomorphic (rarely actinomorphic). Epigyny. Sepals four or five, with imbricate or open aestivation, small, usually persistent, connate; median sepal adaxial. Petals four or five, with imbricate aestivation, caducous, connate to tubular to infundibuliform corolla (corolla often bilabiate, in Lonicera with quadrilobate upper and unilobate lower lip). Corolla at base with swelling covered by nectariferous unicellular hairs. Disc present or absent.

Androecium Stamens four or five, approximately equal in length, haplostemonous, antesepalous, alternipetalous. Filaments glabrous, free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum usually amoeboid-periplasmodial. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate, -colpate or -porate?, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, microreticulate, spinulate.

Gynoecium Pistil composed of usually two to five (to eight?) connate carpels, all or two, three or four of which fertile; odd carpel adaxial (in Leycesteria antepetalous). Ovary inferior, usually bilocular to quinquelocular (to octalocular; ovaries in Lonicera often connate); one or two locules often compressed and sterile. Style single, simple. Stigma capitate to truncate, or bilobate to quinquelobate, papillate, Wet type. Pistillodium absent.

Ovules Placentation axile to apical. Ovules one to numerous per carpel, anatropous, pendulous, apotropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type, or disporous, Allium type, or tetrasporous, Adoxa type. Endosperm development cellular. Endosperm haustoria? Embryogenesis asterad (also other types?).

Fruit A one- to three-seeded drupe (in Symphoricarpus externally with thick layer of narrow vertical fibres and internal to this a layer of horizontal fibres), or a many-seeded berry or syncarp (Lonicera).

Seeds Aril absent. Testa in Symphoricarpus crushed. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, oily. Embryo straight, without chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA Plastid gene infA lost/defunct. Deletion in plastid gene clpP present in at least some species of Lonicera. Mitochondrial intron coxII.i3 lost. Duplication of nuclear genes dipsCYC2 and dipsCYC3?

Phytochemistry Flavonols (kaempferol, quercetin; O-methylated?), flavones (dominating), Group VI secoiridoids (secologanin, morroniside, kingiside), Group VII secoiridoids (sweroside), and Group X secoiridoids (loganin) present. Proanthocyanidins (cyanidin), alkaloids, and phenylalanine-derived cyanogenic compounds sometimes present. Coumarins present in Lonicera and Symphoricarpos. Ellagic acid not found.

Systematics Heptacodium (1; H. miconioides; central and eastern China); Triosteum (6; T. angustifolium, T. aurantiacum, T. himalayanum, T. perfoliatum, T. pinnatifidum, T. sinuatum; eastern Himalayas to China, the Korean Peninsula and Japan, eastern United States), Lonicera (c 180; temperate and subtropical regions on the Northern Hemisphere south to the Philippines and Mexico), Leycesteria (6–7; L. crocothyrsos, L. formosa, L. glaucophylla, L. gracilis, L. sinensis, L. stipulata, L. thibetica; western Himalayas to southwestern China), Symphoricarpos (16; Canada, United States, Mexico, Central America, one species, S. sinensis, in western China).

Caprifoliaceae are sister to [Linnaeaceae+[Morinaceae+[Dipsacaceae+Valerianaceae]]].

Heptacodium is sister to the remaining Caprifoliaceae.

Cladogram of Caprifoliaceae based on DNA sequence data (Winkworth & al. 2008). In analyses by Theis & al. (2008) Leycesteria is sister to Symphoricarpos and Lonicera sister to [Leycesteria+Symphoricarpos].

Fossils Weigela fossils have been found in Miocene and Pliocene layers in Europe and Siberia.

Vegetative anatomy Phellogen ab initio pericyclic. Vessel elements usually with scalariform (sometimes reticulate) perforation plates; lateral pits alternate or scalariform?, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty or banded, or absent. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Crystals?

Leaves Opposite, simple, entire, with involute ptyxis (Weigela). Stipules and leaf sheath absent. Petiole vascular bundle arcuate. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids as platelets (Weigela) or absent (Diervilla). Leaf margin serrate or entire.

Inflorescence Terminal or axillary, cymose; flowers often paired or solitary axillary. Supernumerary bracts usually absent (present in some species).

Flowers Zygomorphic (in Weigela almost actinomorphic). Epigyny. Sepals four or five, with open or imbricate? aestivation, free (Weigela) or connate (Diervilla); median sepal adaxial. Petals four or five, with imbricate aestivation, connate to tubular to infundibuliform corolla (in Diervilla with quadrilobate upper lip). Corolla at base with swelling covered by nectar secreting unicellular hairs. Disc absent.

Androecium Stamens four or five, antesepalous, alternipetalous. Filaments hairy, free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial? Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tripororate, shed as monads, tricellular at dispersal. Exine tectate, with acolumellate to columellate infratectum (with little developed columellae), spinulate.

Gynoecium Pistil composed of two connate carpels, one of which often aborted and sterile. Ovary inferior, unilocular or bilocular. Style single, simple. Stigma capitate, papillate, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules numerous per carpel, anatropous, pendulous, apotropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development cellular. Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule (with beak-shaped processes and two valves), dehiscing downwards along its sides.

Seeds Aril absent. Testa with (Weigela) or without (Diervilla) comb-shaped wing. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, oily. Embryo straight, well developed, without chlorophyll. Cotyledons two. Germination phanerocotylar?

DNA Deletion in plastid gene clpP? Duplication of nuclear genes dipsCYC2 and dipsCYC3?

Phytochemistry Insufficiently known. Flavonols (O-methylated?; dominating) and Group VI secoiridoids (secologanin) present. Coumarins present. Ellagic acid not found?

Scabiosaceae Martinov, Tekhno-Bot. Slovar: 563. 3 Aug 1820 [’Scabioseae’]; Triplostegiaceae (Höck) A. E. Bobrov ex Airy Shaw in Kew Bull. 18: 269. 8 Dec 1965; Dipsacineae Shipunov in A. Shipunov et J. L. Reveal in Phytotaxa 16: 63. 4 Feb 2011

Distribution Temperate and subtropical regions of Eurasia, North Atlantic islands, and Africa, with their largest diversity in the Mediterranean and western Asia; some species on mountains in southern India, Sri Lanka, Southeast Asia, Central Malesia, and New Guinea; Triplostegia: eastern Himalayas, southern China, Taiwan, Sulawesi, New Guinea.

Habit Bisexual, usually perennial, biennial or annual herbs (sometimes suffrutices, rarely shrubs).

Vegetative anatomy Phellogen ab initio usually pericyclic (at least in Knautia often superficial). Pericyclic fibrous envelope often well developed. Flank bridge in stem between lateral vascular bundles? Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids with bordered pits. Wood rays uniseriate or multiseriate, heterocellular, or absent. Axial parenchyma? Sieve tube plastids S type. Endodermis sometimes with thick-walled cells (e.g. Dipsacus). Nodes 3: 3, trilacunar with three leaf traces (lateral traces bifurcating, median trace not ramifying). Calcium oxalate crystals usually in clusters (wood ray cells each with one rhomboidal calciumoxalate crystal).

Trichomes Hairs unicellular, usually lignified, or multicellular (in Cephalaria hair tufts); glandular hairs often present (in Knautia capitate glandular hairs).

Leaves Usually opposite (rarely verticillate), simple or pinnately compound, entire or pinnately lobed, sometimes coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles?; vascular flank bridge connecting vascular strands leading into petioles. Leaf bases sometimes connate. Venation pinnate. Stomata usually anomocytic (sometimes anisocytic). Cuticular wax crystalloids? Leaf margin and leaflet margins serrate or entire.

Inflorescence Terminal, usually head-like cymose, usually as pseudanthia (polythelic capitula), often with involucre consisting of bracts (in Triplostegia few-flowered open paniculate thyrse with glanduliferous branches). Supernumerary bracts (forming epicalyx) present (number of bracts larger than number of flowers, due to numerous floral abortions). Usually persistent (in Cephalaria, Dipsacus, and Knautia caducous) eight-ribbed epicalyx, consisting of usually two connate floral prophylls (bracteoles), usually surrounding calyx (epicalyx absent from some species of Cephalaria and Succisa; in Triplostegia two epicalyces present: one outer/lower epicalyx, consisting of four basally connate glandular acute bracts, and one inner/upper epicalyx, consisting of cupularly connate floral prophylls with eight ridges); epicalyx tube with usually eight ribs (in Knautia four ribs).

Flowers Usually zygomorphic (corolla in Triplostegia actinomorphic), usually small. Epigyny. Sepals usually five (often bilobate, in Triplostegia four), with valvate aestivation, usually bristle-like or membranous, usually persistent, free (when bristle-like) or connate (two sepals often entirely connate). Petals four or five (in Triplostegia four), with imbricate aestivation, unequal in size, caducous, connate into tubular or infundibuliform, often bilabiate corolla (two petals often entirely connate; lateral abaxial corolla lobes overlapping adaxial lobes). Nectaries usually present at stylar base (in Triplostegia at abaxial-basal part of corolla tube), as unicellular hairs. Disc intrastaminal.

Androecium Stamens usually four (rarely three; in Pterocephalidium diandrum two adaxial), antesepalous, alternipetalous; adaxial stamen absent; two abaxial stamens often shorter. Filaments free from each other, adnate to corolla lobe (epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial, with multinucleate cells. Staminodia usually absent (sometimes one or two?).

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–4)-colpate or 3(–4)-porate (in Triplostegia tricolpor[oid]ate), with costae surrounding apertures, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum (with branched columellae), perforate to microperforate, spinulate (with spines or microspines) or echinate.

Gynoecium Pistil composed of usually two (in Triplostegia three) connate carpels, one of which (adaxial-lateral) fertile. Ovary inferior, unilocular, pseudomonomerous. Style single, simple, narrow. Stigma entire or bilobate (with unequally sized lobes; in Triplostegia small), capitate, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation apical. Ovule one per ovary, anatropous, pendulous, apotropous, unitegmic, tenuinucellar. Micropyle very long. Integument ten to 15 cell layers thick, non-vascularized? Hypostase usually present (absent from Triplostegia), with cavities containing yellowish liquid. Megagametophyte monosporous, Polygonum type. Chalazal cells binucleate to quadrinucleate, polyploid. Endosperm development cellular. Endosperm haustoria present in Triplostegia. Embryogenesis piperad.

Fruit A single-seeded cypseloid nut with persistent epicalyx and with remnants of calyx as bristles or prickles (in Triplostegia with two epicalyces).

Seeds Aril absent. Elaiosome present in Knautia. Testa membranous, consisting of outer epidermis and hypodermis, remaining parts crushed. Exotestal cells? crushed? Endotesta? Perisperm not developed. Endosperm sparse, oily (starch almost absent), not ruminate. Embryo large, straight, well differentiated, with chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA Elongation of plastid inverted repeat. Duplication of nuclear genes dipsCYC2 and dipsCYC3. Horizontal transfer of mitochondrial gene rps11.

Phytochemistry Flavonoids, Group VI secoiridoids (secologanin), Group VII secoiridoids (gentiopicroside), monoterpenoid glucoindole alkaloids, triterpenoid glycosides, alkaloids, coumarins, phenolic compounds, caffeoyl guinic acid derivatives, lignin glycoside, and saponins present. Valepotriate iridoids present in Triplostegia. Quercetin, ellagic acid, proanthocyanidins and cyanogenic compounds not found. Inulin? (in Cephalaria?) Catechol tannins? O-methylated flavones?

Systematics Dipsacaceae are sister-group to Valerianaceae.

Triplostegia is sister to the remaining Dipsacaceae.

Triplostegia

1/2. Triplostegia (2; T. glandulifera, T. grandiflora; eastern Himalayas, southern China, Taiwan, Sulawesi, New Guinea). – Herbs. Inflorescence few-flowered open paniculate thyrse with glanduliferous branches. Flowers tetramerous, with floral prophylls (bracteoles) and double epicalyx: one outer/lower epicalyx, consisting of four basally connate glandular acute bracts, and one inner/upper epicalyx, consisting of cupularly connate floral prophylls with eight ridges. Corolla actinomorphic. Stamens four. Pollen grains tricolpor[oid]ate, apertures with halo and granular colpus. Pistil composed of three carpels. Hypostase absent. Endosperm haustoria present. Fruit surrounded by capitate-glandular outer epicalyx and with hooked tips. Endosperm reduced. n = ? Valepotriate iridoids present. – Valerianaceae also have pollen aperture with halo and granulate colpus, and valepotriates are present (apart from several other morphological similarities).

Dipsacoideae Eaton, Bot. Dict., ed. 4: 36. Apr-Mai 1836 [‘Dipsaceae’]

12/300–305. Bassecoia (3; B. bretschneideri, B. hookeri, B. siamensis; the Himalayas, western China, Thailand); Pseudoscabiosa (5; P. africana, P. diandra, P. grosii, P. saxatilis, P. limoniifolia; western and central Mediterranean), Succisa (3; S. pratensis: Europe, the Mediterranean, North Africa, western Siberia; S. pinnatifida: northwestern Iberian Peninsula; S. trichotocephala: Cameroon), Succisella (5; S. andreae-molinae, S. carvalhoana, S. inflexa, S. microcephala, S. petteri; the Iberian Peninsula, eastern Europe from the Balkan Peninsula to Lithuania); Knautia (c 60; Europe, the Mediterranean), Pterocephalidium (1; P. diandrum; central Spain, Portugal), Pterothamnus (1; P. centennii; Mozambique); Cephalaria (c 80; Europe, the Mediterranean to Central Asia, Ethiopia, southern Africa), Dipsacus (c 20; Europe, the Mediterranean, mountains in tropical Africa, temperate Asia, southern India, Sri Lanka); Lomelosia (>50; the Mediterranean to the Caucasus), Pterocephalus (c 30; Macaronesia, the Mediterranean to Central Asia, the Himalayas and western China, tropical Africa), Scabiosa (c 40; Europe, the Mediterranean, temperate Asia, East African mountains, southern Africa). – Temperate regions in Europe, Africa and Asia, eastern and southern Africa and southern Asia to New Guinea. Inflorescence capitulum with involucre. Epicalyx single, eight-ribbed. Flowers usually pentamerous. Stamens sometimes two or three. Fruit with calycine awns or bristles. n = 5, 7–9 (10). – The epicalyx morphology varies strongly among Dipsacoideae. The epicalyx is often provided with wings or spiny bristles. In species with reduced or without corona, the number of calyx bristles is often increased to form a pappus-like structure adapted to long-distance wind dispersal. Some groups are marked by additional epicalyx structures often associated with elaborate wing-like appendages. The ‘diaphragma’ is formed by a secondary meristem on the inner side of the epicalyx, and encloses the stalk of the calyx-like collar. The ‘epi-diaphragma’ is a small attachment present between the diaphragma and the ‘corona’, which is a membranous wing-like structure that forms the apex of the epicalyx.

Phylogeny (founded on Maximum parsimony and Bayesian inference trees) of Dipsacaceae s. str. based on DNA sequence data (Avino & al. 2009; Carlson & al. 2009; Mayer & Ehrendorfer 2013). Triplostegia is usually sister-group to the remaining Dipsacaceae (sometimes sister to Valerianaceae, or to the clade [Dipsacaceae+Valerianaceae]).

Distribution Temperate regions on the Northern Hemisphere south to the Himalayas and Mexico, with their highest diversity in China.

Fossils Winged fruits from North American Late Eocene have been described as Diplodipelta, and fossils assigned to ‘Dipelta’ (Linnaea) are also known from the Late Eocene of England. Pollen fossils were reported from the Eocene in Greenland and Central Europe.

Habit Bisexual, evergreen or deciduous shrubs or small trees, or creeping and partially woody.

Vegetative anatomy Phellogen ab initio pericyclic. Pericyclic fibrous envelope often well developed. 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 uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, or absent. Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces. Crystals?

Trichomes Hairs unicellular or multicellular?; glandular hairs often present.

Leaves Opposite, simple, entire, with supervolute (to curved) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids as platelets (in Dipelta and Kolkwitzia absent) Leaf margin serrate or entire.

Inflorescence Axillary, few-flowered cymose (flowers often paired), or flowers solitary axillary. Supernumerary bracts often present (cf. Caprifoliaceae).

Flowers Zygomorphic. Epigyny. Sepals four or five, with open aestivation, entirely or partially connate; median sepal adaxial. Petals four or five, with imbricate aestivation, connate into infundibuliform corolla. Abaxial petal with nectary as basal swelling with nectariferous unicellular hairs. Disc absent.

Androecium Stamens four or five, antesepalous, alternipetalous. Filaments hairy, free from each other, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (in ‘Abelia’ sometimes zonorate), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, psilate to spinulate.

Gynoecium Pistil composed of two (to four) connate carpels, one or two (adaxial-lateral) of which fertile. Ovary inferior, usually bilocular (sometimes trilocular or quadrilocular, with two locules compressed and sterile). Style single, simple. Stigma capitate, papillate?, Wet type? Pistillodium absent.

Ovules Placentation axile to apical? Ovule one or several? per carpel, anatropous, pendulous, apotropous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development cellular. Endosperm haustoria? Embryogenesis?

Fruit Usually a nut or drupe (rarely berry), often with calyx and/or bracts persistent and modified (cypseloid?; L. dipelta and L. yunnanensis have more or less fleshy two-seeded fruit with two accrescent bracts).

Seeds Aril absent. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, oily. Embryo straight, well developed, without chlorophyll. Cotyledons two. Germination phanerocotylar?

DNA Deletion in plastid gene clpP? Duplication of nuclear genes dipsCYC2 and dipsCYC3?

Phytochemistry Very insufficiently known. Group VI secoiridoids (e.g. secologanin) present. Flavonols (O-methylated?) dominating, flavones.

Rafinesque in Ann. Gén. Sci. Phys. Bruxelles 6: 88. Oct-Dec 1820 [’Morinidia’]

Distribution The Balkan Peninsula, southwestern Asia to Afghanistan, Central Asia, the Himalayas, East Asia.

Vegetative anatomy Phellogen ab initio pericyclic? Flank bridge present in stem between lateral vascular bundles. Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements libriform fibres with bordered? pits. Wood rays absent? Axial parenchyma? Sieve tube plastids S type. Nodes 5:5, quinquelacunar with five leaf traces. Calcium oxalate crystals as druses.

Trichomes Hairs usually unicellular; corolla with stalked multicellular glands.

Leaves Usually opposite (sometimes verticillate), simple, usually pinnately lobed (rarely entire), with involute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles?; petiole with vascular flank bridge connecting vascular strands leading into petioles. Leaf bases often sheathing and connate. Venation palmate. Stomata anomocytic. Cuticular wax crystalloids? Epidermis with or without mucilage cells, with secretory cells containing ethereal oils. Leaf margin usually serrate (often serrate-dentate; rarely entire).

Inflorescence Axillary, in cymose whorls along vertical stem; partial inflorescences consisting of sessile cymes. Supernumerary bracts absent. Epicalyx campanulate, 12-ribbed, consisting of two connate serrate-dentate floral prophylls (bracteoles, bracts?) enclosing flower.

Flowers Zygomorphic. Epigyny. Sepals two or four, with imbricate? aestivation, persistent, connate and bilabiate, with lips entire or bilobate. Petals five, with imbricate aestivation, connate into more or less tubular bilabiate corolla. Nectariferous disc intrastaminal, annular. Nectaries as unicellular hairs.

Androecium Stamens two fertile adaxial stamens and two staminodia (didynamous; adaxial stamen absent), haplostemonous, antesepalous, alternipetalous. Filaments long, thick, free from each other, adnate to corolla tube (epipetalous. Anthers dorsifixed or basifixed?, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial. Staminodia two (corresponding to abaxial two stamens) or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–4)-colporate (sometimes 3(–4)-pororate), shed as monads, bicellular at dispersal. Exine tectate, with acolumellate to columellate infratectum (with poorly developed columellae), imperforate?, spinulate? Pollen tube-like structures covered by intine formed prior to germination of pollen grains.

Gynoecium Pistil composed of two connate carpels, one of which degenerating (adaxial-lateral carpel fertile). Ovary inferior, unilocular. Style single, simple, narrow. Stigma capitate, type? Pistillodium absent.

Ovules Placentation apical. Ovule one per ovary, anatropous, pendulous, unitegmic, tenuinucellar. Integuments 14 to 18 cell layers thick, non-vascularized. Hypostase well developed, without cavity. Megagametophyte monosporous, Polygonum type. Antipodal cells proliferating. Chalazal cells producing multicellular structures. Endosperm development cellular. Endosperm haustoria absent. Embryogenesis solanad.

Fruit A single-seeded cypseloid nut with persistent remains of calyx. Pericarp thick, rugose.

Seeds Aril absent. Seed coat exotestal-epidermal. Testa thin, multiplicative, non-vascularized, consisting of outer epidermis. Exo- and endotestal cells used up during endosperm development. Perisperm not developed. Endosperm sparse, ruminate, oily. Embryo large, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar?

DNA Duplication of nuclear genes dipsCYC2 and dipsCYC3. Horizontal transfer of mitochondrial gene rps11? Elongation of plastid inverted repeat?

Phytochemistry Very insufficiently known. Alkaloids, saponins and polyphenols present. Ethereal oils present. Monoterpenoids? Iridoids, ellagic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found. Cathecole tannins?

Systematics Zabelia (12–13; Central Asia, Afghanistan, northwestern India, Nepal, China, the Korean Peninsula, Japan, the Russian Far East); Cryptothladia (4; C. chinensis, C. chlorantha, C. kokonorica, C. polyphylla; the Himalayas, western China), Morina (3; M. couteriana, M. longifolia, M. persica; Greece, Bulgaria, Romania, Turkey, Lebanon, Israel, Iraq, Iran, Central Asia, eastern Himalayas, western China).

Morinaceae are sister to [Dipsacaceae+Valerianaceae].

Zabelia, with psilate exine and endocingulum at inner nexine wall layer, is ususally sister to the remaining Morinaceae (but see Landrein & al. 2012). The generic limits and sister-group relationships among the majority of Morinaceae are still fairly confusing.

Cladogram of Morinaceae based on DNA sequence data (Bell 2004, Winkworth & al. 2008; Jacobs & al. 2010). Acanthocalyx is sister to [Dipsacaceae+Valerianaceae] and Zabelia sister to the clade [[Cryptothladia+Morina]+[Acanthocalyx+[Dipsacus clade+Valeriana clade]]] in some analyses by Landrein & al. (2012). However, in other most of their analyses Zabelia is part of a trichotomy also comprising Linnaeaceae and [Dipsacaceae+Valerianaceae] and Morinaceae sister-group to this trichotomy (Landrein & al. 2012).

Valerianales DC. ex Bercht. et J. Presl, Přir. Rostlin: 256. Jan-Apr 1820 [‘Valerianeae’]

Distribution Temperate regions on the Northern Hemisphere, Mexico, Central America, the West Indies, South America (c. 50% of the species occurring in the Andes), mountain areas in eastern and southern Africa, and South Asia to Sumatra and Java, with their largest diversity in the Mediterranean, West Asia and the Andes.

Habit Usually bisexual (sometimes polygamomonoecious or dioecious), usually perennial, biennial or annual herbs (rarely shrubs). Often evil-smelling.

Vegetative anatomy Phellogen ab initio pericyclic. Flank bridge in stem between lateral vascular bundles? Primary vascular tissue cylinder of vascular bundles. Vessel elements extremely small, usually with simple (in Patrinia scalariform) perforation plates; lateral pits alternate. Imperforate tracheary xylem elements libriform fibres and/or fibre tracheids (in Patrinia tracheids) with simple or bordered pits. Wood rays multiseriate or absent. Axial parenchyma usually scanty paratracheal (rarely vasicentric or confluent) or absent. Sieve tube plastids S type. Nodes usually 3:3–5, trilacunar with three to five leaf traces (rarely 1:1 or 5–7:5–7). Rhizomal and root cortex and cork tissue with idioblastic cells containing strongly smelling ethereal oils. Calcium oxalate crystals sometimes present.

Trichomes Hairs unicellular or multicellular, simple; glandular hairs often present.

Leaves Opposite, simple or pinnately compound, entire or pinnately lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole bases often sheathingly connate. Petiole vascular bundles?; vascular flank bridge connecting vascular strands leading into petioles. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Hydathodes sometimes present. Leaf margin and leaflet margins serrate or entire.

Inflorescence Terminal, umbel-like, corymbose, capitate or paniculate cymose. Supernumerary bracts absent. Epicalyx absent. Floral prophylls (bracteoles) present.

Flowers Zygomorphic or asymmetrical (in Patrinia and Nardostachys almost actinomorphic), usually small. Epigyny. Sepals usually minute and reduced (in Patrinia and Nardostachys five, well developed), persistent, connate, at anthesis often as whorl of up to 20 inrolled parts, later unrolled, expanded and bristle-like or plumose (pappus-like). Petals usually five (sometimes three), with imbricate aestivation, connate into infundibuliform or tubular (sometimes bilabiate) structure; abaxial petal often with nectariferous spur at base (corolla in Centranthus divided by vertical septum). Nectaries as unicellular hairs. Disc absent?

Androecium Stamen one (in Centranthus adaxial), two (adaxial, in ‘Fedia’), three (in ‘Fedia’ adaxial and central stamens absent), four (adaxial stamen absent; in Patrinia and Nardostachys) or five, haplostemonous, antesepalous, alternipetalous, asymmetrically inserted. Filaments usually free from each other (two stamens in ‘Fedia’ connate), adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile?, usually tetrasporangiate (in Valeriana subgenus Phyllactis disporangiate), introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial, with multinucleate cells. Staminodia absent.

Pollen grains: Microsporogenesis simultaneous. Pollen grains 3(–4)-colpate or 3(–4)-colpor(oid)ate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum (with branched columellae), perforate (to microperforate) spinulate or echinate.

Gynoecium Pistil composed of three connate carpels. Ovary inferior, trilocular (two locules reduced, sterile, adaxial-lateral carpel fertile; sometimes unilocular, pseudomonomerous). Style single, simple, narrow. Stigma capitate or bi- or trilobate, Dry type. Pistillodium absent.

Ovules Placentation apical. Ovule one per ovary, anatropous, pendulous, apotropous?, unitegmic, tenuinucellar. Integument ? cell layer thick, non-vascularized? Hypostase sometimes present. Megagametophyte monosporous, Polygonum type. Chalazal cells binucleate to quadrinucleate. Antipodal cells persistent. Endosperm development cellular. Endosperm haustoria? Embryogenesis asterad.

Fruit A single-seeded cypseloid nutlet, usually with persistent and accrescent calyx as pappus-like (winged, plumose, etc.) appendages, hooks or membranous margins (sometimes reduced).

Seeds Aril absent. Testa one-layered. Testal cells? Perisperm not developed. Endosperm absent or almost absent (sometimes sparse, starchy). Embryo large, straight, well differentiated, oily, sometimes with chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA Elongation of plastid inverted repeat. Duplication of nuclear genes dipsCYC2 and dipsCYC3. Horizontal transfer of mitochondrial gene rps11 to nuclear genome. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol), O-methylated flavonols, Route I iridoids and secoiridoids, valepotriates (triesters of Route I secoiridoids), Group V carbocyclic iridoids (valtrate, villoside, patrinoside), Group VI secoiridoids (morroniside), Group X secoiridoids (loganin, iridoid pyridine alkaloids), foetid ethereal monoterpenoids sesquiterpenoids, alkaloids, triterpene saponins, and valeric acid present. Quercetin, ellagic acid, proanthocyanidins, and cyanogenic compounds not found. Catechol tannins?

Use Ornamental plants, medicinal plants, perfumery (Nardostachys), vegetables (Valerianella).

Systematics Patrinia (c 17; Ural and western Asia to the Himalayas and East Asia); Nardostachys (1; N. jatamansi; the Himalayas, western China); Centranthus (9; southern Europe, the Mediterranean; in Valeriana?), ’Valeriana’ (c 270; temperate regions on the Northern Hemisphere, mountain areas in eastern and southern Africa, South Asia to Sumatra and Java, Mexico, Central America, the West Indies, South America with their highest diversity in the Andes; polyphyletic), Valerianella (c 70; temperate regions on the Northern Hemisphere south to North Africa).

Valerianaceae are sister-group to Dipsacaceae.

Patrinia is sister to the remaining Valerianaceae, and Nardostachys successive sister to the remainder. Sometimes Patrinia and Nardostachys form a joint sister-group to the remaining Valerianaceae. Patrinia contains villoside and patrinoside. Valerianella (including Fedia) and Valeriana celtica + V. hardwickii are successive sister-groups to Valeriana (including Centranthus) (Hidalgo & al. 2010).

Triplostegia is sometimes placed as sister-group to Valerianaceae (morphological or both morphological and molecular data). Both Triplostegia and Valerianaceae have valepotriate iridoids, a pollen aperture with halo and granulate colpus, and also a reduced endosperm, apart from similarities in epicalyx morphology.

Phylogeny of Valerianaceae based on DNA sequence data (Hidalgo & al. 2010; Bell & al. 2012). A similar topology was obtained by Bell & al. (2015), although with a larger number of taxa.

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Asplund E. 1920. Studien über die Entwicklungsgeschichte der Blüten einiger Valerianaceen. – Kungl. Sv. Vetensk.-Akad. Handl. 61(3): 1-65.

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DIPSIDAE D. C. Tank et M. J. Donoghue

DIPSACALES Juss. ex Bercht. et J. Presl

ADOXACEAE E. Mey.

CAPRIFOLIACEAE Juss.

DIERVILLACEAE (Raf.) Pyck

DIPSACACEAE Juss.

LINNAEACEAE (Raf.) Backlund

MORINACEAE Raf.

VALERIANACEAE Batsch