TRICOLPATAE M. J. Donoghue, J. A. Doyle et P. D. Cantino

Donoghue, Doyle et Cantino in Taxon 56: E26. Aug 2007

[Ranunculales+[Sabiaceae+Proteales+[Trochodendrales+[Didymelales+Gunneridae]]]]


RANUNCULALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 215. Jan-Apr 1820 [‘Ranunculaceae’]

Berberidopsida Brongn., Enum. Plant. Mus. Paris: xxv, 94. 12 Aug 1843 [’Berberineae’]; Ranunculopsida Brongn., Enum. Plant. Mus. Paris: xxvi, 96. 12 Aug 1843 [’Ranunculineae’]; Ranunculanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992; Ranunculidae Takht. ex Reveal in Novon 2: 235. 13 Oct 1992; Berberidanae Doweld, Tent. Syst. Plant. Vasc.: xxv, xxvi. 23 Dec 2001; Papaveranae Doweld, Tent. Syst. Plant. Vasc.: xxv. 23 Dec 2001

Fossils Teixeiraea lusitanica has been assigned to Ranunculales (and possibly to Menispermaceae). It is a fossilized male flower from the Late Aptian to the Early Albian of Portugal. The spirally arranged floral parts are free, the outermost ones being bracteate and grading into tepaloid parts. The 20 stamens have basifixed anthers and the tectate-perforate pollen grains have a columellate infratectum.

Habit Usually bisexual (sometimes monoecious or dioecious, rarely polygamomonoecious), shrubs, lianas, suffrutices, perennial, biennial or annual herbs (rarely trees).

Vegetative anatomy Phellogen ab initio superficially or deeply seated, or absent. Primary vascular tissue consisting of one or several cylinders of vascular bundles, or of scattered bundles. Secondary lateral growth normal, from cylindrical cambium, anomalous, from normal successive cambia, or absent. Fusiform cambial initials storied (not in Glaucidiaceae). Vessels confined to central portions of fascicular areas (vessel restriction patterns, with very few or no vessels touching rays). Vessel elements with simple or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, opposite or scalariform, bordered pits. 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 usually multiseriate (sometimes uniseriate), homocellular or heterocellular, or absent. Axial parenchyma apotracheal, diffuse or diffuse-in-aggregates or banded, or paratracheal, intervascular or scanty vasicentric, banded, or absent. Sieve tube plastids S or Ss type. Nuclei sometimes with dispersive P-protein. Nodes usually ≥3:≥3, trilacunar or multilacunar with three or more leaf traces (rarely 1:1, 1:4 or 1:5–11, unilacunar with one to many traces, or 2:2, bilacunar with two traces). Latex cells and laticifers with white or coloured latex sometimes present. Idioblasts with raphides sometimes present. Prismatic or rhomboidal calciumoxalate crystals often frequent; crystal sand sometimes present. Calcium oxalate crystals usually present in wood rays (not in Glaucidiaceae and Ranunculaceae).

Trichomes Hairs unicellular och multicellular, usually uniseriate (sometimes multiseriate, rarely branched), or absent; glandular hairs often present.

LeavesUsually alternate (spiral or distichous; sometimes opposite), simple or compound, entire or pinnately or palmately lobed, with supervolute, involute, conduplicate, plicate, subplicate, reclinate, curved or equitant ptyxis. Stipules intrapetiolar or absent; petiole base sometimes sheathing. Petiole vascular bundle transection arcuate or annular. Venation pinnate or palmate, actinodromous or acrodromous (rarely parallelodromous or flabellate). Stomata usually anomocytic (sometimes paracytic, staurocytic or cyclocytic) or absent. Cuticular wax crystalloids usually as irregularly shaped platelets or clustered tubuli (Berberistype), dominated by nonacosan-10-ol (without triterpenoids), or absent. Mesophyll sometimes with sclerenchymatous idioblasts and mucilage cells. Idioblasts with ethereal oils absent. Leaf margin serrate, serrate-dentate, spinose-serrate or glandular-serrate, with chloranthoid or platanoid teeth, crenate or entire.

Inflorescence Terminal or axillary, simple or compound, cymose, raceme-, head- or umbel-like, fascicle, panicle, thyrsoid, spike, raceme, or flowers solitary. Floral prophylls (bracteoles) single or pairwise (sometimes absent).

Flowers Usually actinomorphic (sometimes zygomorphic or bisymmetric). Hypogyny. Perianth sometimes trimerous. Tepals (three or) four to 15, spiral or whorled (rarely absent). Outer tepals two or more, with imbricate or valvate (rarely open) aestivation, sepaloid or petaloid, persistent or caducous, usually free. Inner tepals (staminodial in origin?) one to 13, with imbricate or valvate (sometimes crumpled), free, often with one or several nectaries at base (nectary sometimes absent). Disc absent.

Androecium Stamens (one to) three to more than 500, spiral or whorled. Filaments free or more or less connate, free from tepals. Anthers basifixed to somewhat dorsifixed, usually non-versatile, usually tetrasporangiate (rarely disporangiate), extrorse, latrorse or introrse, usually longicidal (dehiscing by longitudinal slits) or valvate (dehiscing by longitudinal valves). Tapetum usually secretory (rarely amoeboid-periplasmodial). Staminodia extrastaminal, petaloid and/or nectariferous, or absent; female flowers sometimes with staminodia.

Pollen grains Microsporogenesis usually simultaneous (rarely successive). Pollen grains di- or tricolpate to polycolpate or -porate, di- or tricolporate, zono- or pantoaperturate (sometimes syncolpate, rarely spiraperturate, clypeate or inaperturate), usually shed as monads (rarely dyads or tetrads), usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, microreticulate, reticulate or punctate, foveolate, striate, scabrate, spinulate, echinate, microechinate, verrucate or psilate.

Gynoecium Carpels one (monocarpellate or pseudomonomerous) to more than 100, spiral or whorled, usually free (sometimes more or less connate), or pistil composed of one to numerous carpels; carpel ascidiate, conduplicate or plicate, postgenitally usually entirely occluded, without canal. Ovary superior, unilocular (apocarpy or gynoecium monomerous) or trilocular to quinquelocular. Stylodia or style single, simple, or absent. Stigma of various types, lobate to capitate (sometimes decurrent), papillate or non-papillate, Dry or Wet type. Male flowers sometimes with pistillodium.

Ovules Placentation marginal or basal (when ovule single, or when ovary unilocular), laminal-lateral or marginal-lateral (when ovules several), or axile (when ovary plurilocular). Ovules one to numerous per carpel, usually anatropous (sometimes hemianatropous, rarely amphitropous or campylotropous), ascending, horizontal or pendulous, usually apotropous (rarely epitropous), usually bitegmic (sometimes unitegmic), usually crassinucellar (sometimes pseudocrassinucellar). Micropyle endostomal or bistomal, sometimes Z-shaped (zig-zag). Archespore unicellular to at least 15-celled. Nucellar cap usually present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type, or tetrasporous, Adoxa type, Fritillaria type or Peperomia type). Synergids sometimes with a filiform apparatus. Antipodal cells usually persistent, proliferating or non-proliferating. Endosperm development ab initio usually nuclear (rarely cellular). Endosperm haustoria chalazal or absent. Embryogenesis usually onagrad (sometimes solanad, rarely caryophyllad or chenopodiad).

Fruit An assemblage of follicles or achenes, a berry, or a loculicidal, septicidal or poricidal capsule with basipetalous or acropetalous dehiscence (sometimes a multifolliculus or a nut, rarely a lomentum with nut-like mericarps).

Seeds Aril usually absent. Seed coat usually exotestal (rarely endotegmic). Testa often multiplicative (sometimes thin). Exotesta palisade, with often thickened non-lignified cell walls, or seeds more or less pachychalazal, with thin testa. Mesotesta, endotesta and tegmen usually unspecialized. Exotegmen sometimes multiplicative. Perisperm not developed. Endosperm scarce to copious (rarely absent), oily, proteinaceous or starchy. Embryo small to large, undifferentiated to well differentiated at dispersal, without chlorophyll. Cotyledons (one or) two. Germination phanerocotylar or cryptocotylar.

Cytology n = 6–9, 11–13, 15, 19, 21

DNA Nuclear gene AP3 present and usually triplicated.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, rhamnocitrin; frequently additional oxygenation at carbons 6 or 8 of ring A in contrast to magnoliid clades), O-methylated flavonoids, flavones, isoprenylated flavonoids, cyanidin, delphinidin, dihydrochalcones, diterpenoids, tannins, cardioactive bufadienolides, Digitalis cardenolides, poisonous sesquiterpene lactones, caffeic acid, benzylisoquinoline and aporphine alkaloids (benzyltetrahydroisoquinoline and aporphine derivatives in dimeric form, bulbocapnine and other aporphines, morphinanes, pavines, isopavines, dehydrogenated benzophene anthridines, reduced benzophene anthridines, tyrosine-derived berberine, tetrahydroberberines, berberidine, protopines, rhoeadines, narceines, spirobenzyl isoquinolines, hydrastine, protopine, quaternary magnoflorine and its precursor corytuberine), hasubanane alkaloids (protostephanines, erythrinanes, cocculolidines, morphines, quettamine-morphine dimers, hasubanonines, acutumines, etc.), azafluoranthene alkaloids, tropoloisoquinoline alkaloids, protoberberine alkaloids, diterpene alkaloids (aconitine, methyl lycaconitine etc.), quinolizidine alkaloids (e.g. darvasamine), pyrrolizidine alkaloids as macrocyclic diesters, tyrosine-derived cyanogenic glycosides, lignan-β-glycosides, ranunculins (glucosides), triterpene saponins, steroidal saponins, meconic acid, chelidonic acid, fumaric acid, nitrophenyl ethan, phenylic cinnamide, furofuran lignans, mannitol, and glaupalol (a furanocoumarin) present. Ethereal oils, prodelphinidin, and flavonoids and tannins containing a trihydroxylated B-ring (gallic and ellagic acid derivatives) not found.

Systematics Ranunculales are sister-group to the remaining Tricolpatae.

Euptelea is sister to all other Ranunculales, according to most molecular analyses. The majority of Ranunculales are characterized by wide and tall multiseriate wood rays little altered during ontogeny, and relatively intact extensions of primary rays. These contrast with multiseriate rays dominating in other Tricolpatae, in which the large primary rays are rapidly broken into smaller segments during stem and root growth. In general, multiseriate rays in Ranunculales are composed of procumbent cells with the exception of one or two layers of upright sheathing cells.

The clade [[[Lardizabalaceae+Circaeasteraceae]+[Menispermaceae+[Berberidaceae+ [Hydrastidaceae+Glaucidiaceae]+Ranunculaceae]]]+[Papaveraceae s.lat.]] is characterized by the following potential synapomorphies (Stevens 2001 onwards): vessel elements with simple perforation plates, present in diagonal groups; presence of nucleated libriform fibres and vasicentric tracheids; leaves with palmate secondary venation; and Wet type stigma. Herbaceous growth and woody habit have evolved several times in the two main clades above Euptelea. Likewise, petaloid tepals have originated several times from staminodial stamens (e.g. in Berberidaceae, Lardizabalaceae, Ranunculaceae; Drinnan & al. 1994, etc.). In Ranunculaceae, the petaloid tepals have a single vascular strand, share parastichs with the stamens, are similar to stamens in their early development, and are often peltate.

Pteridophyllum and Papaveraceae have among others the following potential synapomorphies in common (Stevens 2001 onwards): herbaceous growth; roots diarch, i.e. lateral roots tetrastichous; perianth differentiated into outer sepaloid and inner petaloid tepal whorls; sepaloid tepals two, median; petaloid tepals four; carpels at least two collateral, occluded by secretion; placentation parietal; capsule septicidal, with persistent woody placenta; and endotesta well developed.

The second main clade has the topology [[Lardizabalaceae+Circaeasteraceae]+ [Menispermaceae+[Berberidaceae+Ranunculaceae]]] and the potential synapomorphies: wide wood rays; trimerous flowers; outer and inner tepals and stamens opposite each other; outer tepals with three or more vascular traces; and triplication of nuclear gene AP3. Both Lardizabalaceae and Circaeasteraceae have extrorse anthers and cellular endosperm development.

The clade [Menispermaceae+[Berberidaceae+[[Hydrastis+Glaucidium]+Ranunculaceae]]] has the benzylisoquinoline berberine and nuclear endosperm development. According to Stevens (2001 onwards), the clade [Berberidaceae+[[Hydrastis+Glaucidium]+Ranunculaceae]] has the potential synapomorphies: usually herbs with non-tuberous rhizome; bright yellow roots and rhizome due to presence of berberine; roots diarch; lateral roots tetrastichous; nodes often multilacunar; vascular bundles V-shaped, in herbaceous species often closed, scattered or arranged in concentric cylinders; broad leaf base; presence of petaloid staminodial nectaries; outer integument at least four cell layers thick; presence of nucellar cap; endosperm with reserves other than oils or proteins; and loss of mitochondrial intron coxII.i3. Hydrastis, Glaucidium and Ranunculaceae share the synapomorphies: uniseriate perianth; and numerous spiral stamens and carpels. Hydrastis and Glaucidium have vessel elements with simple and scalariform perforation plates, medullary vascular bundles, flattened vascular bundles; petiole vascular bundles annular in cross-section; medullary petiole vascular bundles; no palisade mesophyll; distichous leaves; flowers solitary and terminal; no nectaries; bilobate stigma; outer integument four to 13 and inner integument two to five cell layers thick; unmodified antipodal cells; and also abaxial dehiscence of follicle.

Cladogram of Ranunculales based on DNA sequence data (Soltis & al. 2011). Euptelea is usually sister to the remainder with high bootstrap support and, likewise, the clade [Papaveraceae+Pteridophyllum] is sister to the remaining Ranunculales, often with a bootstrap support of 100%. The other clades show high support varying between 75% and 100%. Circaeasteraceae are sister to Lardizabalaceae with a bootstrap support of 78%. According to Wang & al. (2009), Pteridophyllum (Pteridophyllaceae) is nested inside Papaveraceae as sister to Hypecoum. Until this hypothesis has been further confirmed, Pteridophyllum is treated as sister to Papaveraceae, according to, e.g., Kadereit & al. (1994, 1995).

BERBERIDACEAE Juss.

( Back to Ranunculales )

de Jussieu, Gen. Plant.: 286. 4 Aug 1789 [’Berberides’], nom. cons.

Podophyllaceae DC., Syst. Nat. 1: 126. 1-15 Nov 1817 [’Podophylleae’], nom. cons.; Berberidales Bercht. et J. Presl, Přir. Rostlin: 226. Jan-Apr 1820 [‘Berberideae’]; Leonticales Bercht. et J. Presl, Přir. Rostlin: 217. Jan-Apr 1820 [‘Leonticinae’]; Podophyllales Dumort., Anal. Fam. Plant.: 45. 1829 [’Podophyllarieae’]; Diphylleiaceae Schultz Sch., Nat. Syst. Pflanzenr.: 328. 30 Jan-10 Feb 1832; Nandinaceae Horan., Prim. Lin. Syst. Nat.: 90. 2 Nov 1834 [’Nandinaceae (Berberideae)’]; Epimediaceae Menge, Cat. Plant. Grudent. Gedan.: 122. 1839 [’Epimedineae’]; Leonticaceae Airy Shaw in Kew Bull. 18:263. 8 Dec 1965; Ranzaniaceae (Kumaz. et Terab.) Takht. in Bot. Žurn. 79(1): 96. Jan 1994; Nandinales Doweld in Byull. Mosk. Obshch. Ispyt. Prir., Biol. 105(5): 60. 9 Oct 2000

Genera/species 14 or 15/c 700

Distribution Eurasia from Western Europe eastwards to Malesia, North Africa, East African mountains, North and Central America, mountain areas in South America.

Fossils Fossil representatives of Berberis are known from the Oligocene onwards in the Northern Hemisphere.

Habit Bisexual, evergreen or deciduous shrubs or biennial to perennial rhizomatous or tuberous herbs (rarely trees). Roots and rhizome often bright yellow inside due to presence of berberine.

Vegetative anatomy Phellogen ab initio superficially or deeply sited (pericyclic). Primary medullary rays wide in Nandina. Primary vascular tissue consisting of one or more cylinders of bundles or scattered bundles. Secondary lateral growth absent or from normal cylindrical cambium. Xylem V-shaped. Vessel elements with usually simple (in Berberis sometimes also scalariform) perforation plates; lateral pits usually opposite or alternate (in Vancouveria scalariform or pseudoscalariform), bordered pits. Imperforate tracheary xylem elements usually libriform fibres (in Nandina also fibre tracheids; in Jeffersonia tracheids) with usually simple pits (in Nandina bordered pits also), septate or non-septate (also vasicentric tracheids). Wood rays multiseriate, homocellular or heterocellular. Axial parenchyma usually absent (rarely vasicentric scanty). Wood elements partially storied. Sieve tube elements Ss type. Nodes ≥3:≥3, trilacunar or multilacunar with three or more leaf traces. Secondary tissue often yellow due to presence of berberine. Prismatic calciumoxalate crystals often abundant. Wood rays often with rhomboidal crystals.

Trichomes Hairs unicellular or multicellular, uniseriate, or absent.

Leaves Alternate (sometimes in a basal rosette), simple or compound (pinnately or palmately compound; in Nandina and sometimes Leontice bipinnate; in Berberis trifoliolate or unifoliolate), entire or lobate, with conduplicate, reclinate, equitant or curved ptyxis; leaves of long shoots in Berberis usually modified into spines. Stipules rudimentary, intrapetiolar and caducous, or absent; leaf sheath absent. Petiole vascular bundle transection annular or arcuate. Venation pinnate or palmate. Stomata anomocytic. Cuticular wax crystalloids usually as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol (in ‘Podophyllum’ as solid rodlets sometimes aggregated to larger units). Idioblasts with ethereal oils absent. Leaf margin serrate (sometimes serrate-dentate or spinose-serrate), with chloranthoid teeth, or entire.

Inflorescence Terminal or axillary, spike, raceme or fasciculate etc. (in Nandina panicle), or flowers solitary.

Flowers Actinomorphic. Hypogyny. Tepals (eight to) twelve (to 15) (3+3+3, 3+3, 2+2+2, or 2+2; in Epimedium 4+4+4 or 4+5+5; absent in Achlys) in three to six (rarely seven) series (in Nandina slightly spiral), with imbricate aestivation, free, all tepals in Nandina sepaloid; outer tepals three to nine (in Nandina c. 20 to c. 50), in two series, sepaloid or petaloid (or bracts?), caducous; inner tepals six to twelve, in two to four series, outer series petaloid sepals?, inner two or three series petaloid staminodia? with nectaries (in Nandina nectar secreting staminodia; absent inAchlys, Diphylleia and ‘Podophyllum’). Four inner tepals in Epimedium with four nectariferous spurs. Disc absent.

Androecium Stamens (three to) six (to 19), usually as many (sometimes twice as many) as inner petals (in Achlys seven to 15; in Epimedium 2+2), in one whorl, alternisepalous, antepetalous, or in two or more whorls. Filaments free from each other, free? from or adnate? at base to tepals. Anthers basifixed, non-versatile, tetrasporangiate, usually extrorse (often with valves directed ab initio backwards, later usually inverted and turned up-side-down, with pollen grains against floral centre; in Dysosma and Nandina introrse), usually valvicidal (dehiscing by valves; in Dysosma, Nandina and ‘Podophyllum’ longicidal, dehiscing by longitudinal slits); connective sometimes slightly prolonged. Tapetum secretory or amoeboid-periplasmodial, with multinucleate cells. Staminodia three to nine, petaloid, extrastaminal (or absent?).

Pollen grains Microsporogenesis usually simultaneous (in Berberis and Ranzania? at least sometimes successive). Pollen grains usually tricolpate (rarely hexa- to dodecacolpate or spiraperturate; in Berberis sometimes irregularly polysyncolpate, spiraperturate or clypeate [with surface divided into platelets]; in Ranzania hexacolpate), usually shed as monads (in Ranzania etc. dyads; in some species of ‘Podophyllum’ tetrads), bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, usually perforate, microreticulate, striate-reticulate, psilate-punctate, punctate-striate, echinate, gemmate (in Diphylleia spinulate), in Achlys, Epimedium, Jeffersonia, and Vancouveria striate (with compound layer of striae).

Gynoecium Pistil composed of (seemingly?) one carpel (pseudomonomerous gynoecium developed from two or three connate carpels?); carpel secondarily ascidiate (primarily plicate, rarely ascoplicate), seemingly occluded postgenitally by secretion; carpels sometimes opening during seed development. Ovary superior, unilocular. Style short or absent. Stigma wide, often trilobate (rarely quadrilobate), papillate or non-papillate, Dry or Wet type. Carpellary walls in Caulophyllum etc. not enclosing ripening blue seeds. Pistillodium absent.

Ovules Placentation basal (when single ovule) or marginal-lateral (when several ovules). Ovules one to numerous per ovary (in Achlys one ovule; in Nandina two ovules, one of which degenerating), usually anatropous (sometimes hemianatropous or campylotropous), ascending, horizontal or pendulous, bitegmic, crassinucellar to pseudocrassinucellar. Micropyle bistomal, often Z-type (zig-zag). Outer integument five to eleven cell layers thick. Inner integument two to five cell layers thick. Parietal tissue one or two cell layers thick or absent. Nucellar cap six to eight cell layers thick, formed by periclinal divisions from apical cells of megasporangial epidermis. Megagametophyte usually monosporous, Polygonum type (in Caulophyllum at least sometimes tetrasporous, Peperomia type). Synergids sometimes with a filiform apparatus. Antipodal cells persistent, endopolyploid, with large nuclei, non-proliferating. Endosperm development ab initio usually nuclear (sometimes cellular). Endosperm haustoria usually chalazal (absent in some genera, e.g. in ‘Podophyllum’). Embryogenesis onagrad (or variations of this type).

Fruit Usually a berry or a follicle (in Achlys a nut; in Jeffersonia a capsule; in Gymnospermium a papery not completely closed envelope; fruit in Caulophyllum entirely strongly reduced, with fleshy seeds growing out through carpellary wall).

Seeds Elaiosome and aril usually absent (more or less rudimentary aril present in, e.g., Nandina, Epimedium, Jeffersonia and certain species of Berberis). Seed coat usually exotestal (in Nandina endotegmic). Testa often multiplicative. Exotesta palisade, usually well developed, in Berberis with thick-walled lignified cuboid cells. Mesotesta and endotesta unspecialized. Tegmen usually unspecialized (endotegmen in Nandina sclerotic). Perisperm not developed. Endosperm copious, oily or with hemicellulose. Embryo very small, straight, rudimentary, without chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Cytology n = 6 (Podophylloideae); n = 7 (Berberidoideae); n = 8, 10 (Nandinoideae)

DNA Plastid inverted repeat expanded by 11.5 kb into large single copy region in Berberis. Plastid gene rps7 lost in Podophyllum. Mitochondrial intron coxII.i3 lost. Triplication of nuclear gene AP3.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), isoprenylated flavonoids, cyanidin, delphinidin, tannin, frequent caffeic acid, benzylisoquinoline alkaloids (berberine, magnoflorine, protopine etc.), aporphine alkaloids, quinolizidine alkaloids (in Caulophyllum, Leontice and Gymnospermium; in Gymnospermium also darvasamine), cyanogenic compounds (in Nandina tyrosine-derived), and saponins present. Lignan-β-glycosides and similar compounds accumulated in Epimedium, ‘Podophyllum’ and Diphylleia (Dysosma?, Vancouveria?). Ellagic acid not found.

Use Ornamental plants, fruits (Berberis), medicinal plants, dyeing (yellow) substances.

Systematics Berberidaceae are sister-group to the clade [Ranunculaceae+[Hydrastis+Glaucidium]].

Nandina was sister to the remaining Berberidaceae in several molecular analyses (e.g. Kim & Jansen 1998). However, Wang & al. (2007) identified the Podophylloideae clade as sister-group to Berberidoideae in a strict sense, in which Nandina were nested in a clade sister to [Berberis+Ranzania]. The subdivision below follows the latter alternative, although the support for this topology is at most moderate.

Podophylloideae Eaton, Bot. Dict., ed. 4: 38. Apr-Mai 1836 [‘Podophylleae’]

6–8/c 90. Epimedium (c 55; the Mediterranean and North Africa to southwestern Asia, western Himalayas, northeastern Asia, Japan), Jeffersonia (2; J. dubia: Manchuria, northern China; J. diphylla: eastern North America), Achlys (3; Japan, western North America), Bongardia (1; B. chrysogonum; Greece and eastwards to Afghanistan), ‘Podophyllum’ (14; Himalayas to East Asia, eastern North America; paraphyletic; incl. Diphylleia and Dysosma?), Diphylleia (3; D. sinensis: western China, D. grayi: central and northern Japan, Sakhalin; D. cymosa: eastern North America; in Podophyllum?), Dysosma (7; East Asia; in Podophyllum?), Vancouveria (3; western North America). – Temperate regions on the Northern Hemisphere. Lowermost inflorescence branch arising from axil of reduced leaf. Outer tepals four to 18 (absent in Achlys). Inner tepals four to nine (absent in Achlys), with or without nectariferous spurs. Stamens in Achlys seven to 15, in ‘Podophyllum’ up to 19. Anthers longicidal (dehiscing by longitudinal slits). Microsporogenesis successive. Pollen grains sometimes shed as dyads or tetrads. Pollen wall development by centripetal furrowing. Exine usually striate (sometimes spinulate). Gynoecium arising from two carpellary primordia. Ovules one to numerous per carpel. Micropyle sometimes bistomal. Integuments lobate. Inner integument two or three cell layers thick. Megasporocytes in Diphylleia several. Parietal tissue one or two cell layers thick (sometimes absent). Fruit an achene, a berry or a follicle (also with transverse dehiscence). Aril rudimentary or absent. Testa multiplicative. n = 6. Lignan-β-glycosides and similar compounds often accumulated. – Podophylloideae may be sister-group to [Nandinoideae+Berberidoideae].

[Nandinoideae+Berberidoideae]

Tepals with a single trace. Gynoecium arising from two or three carpellary primordia. Stigma usually Wet type.

Nandinoideae Heintze, Cormofyt. Fylog.: 101. 1 Jun 1927

4/15. Nandina (1; Nandina domestica; central China?); Caulophyllum (3; northeastern Asia, eastern North America), Gymnospermium (8; Europe to Japan), Leontice (3; southeastern Europe to North Africa and Central Asia). – East Europe to East Asia. Primary medullary rays wide. In Nandina also fibre tracheids present. Petiole concave at base. Inflorescence paniculate, with lowermost branch arising from axil of expanded leaf. Tepals sepaloid, usually with three traces. Outer tepals in Nandina c. 20–50, spiral. Inner tepals (’petals’) six. Nectary in Nandina absent. Petaloid staminodium and stamen developing from single common primordium. Anthers in Nandina longicidal (dehiscing by longitudinal slits). Pollen grains in Nandina with massive endexine. Ovules one or two (to four) per carpel, (in Nandina two, one of which degenerating). Megasporangium in Nandina early absorbed. Fruit a berry or with evanescent or bladder-like pericarp. Funicle often swollen. Seed coat in Nandina endotegmic, with thin-walled testal cells. Exotesta and mesotesta crushed (Nandina). Endotestal cells in Nandina crystalliferous. Endotegmic cells in Nandina enlarged, lignified, thickened particularly on inner side, crystalliferous. Embryo minute. n = 8, 10. Protopine (a benzylisoquinoline alkaloid) and in Nandina tyrosine-derived cyanogenic compounds present.

Berberidoideae Eaton, Bot. Dict., ed. 4: 41. Apr-Mai 1836

2/>600. Berberis (>600; Europe, the Mediterranean, the Atlas Mountains, East African mountains, temperate Asia, America), Ranzania (1; R. japonica; Japan). – Temperate and mountainous regions, North Africa, South America, with their highest diversity in East Asia and eastern North America. Usually shrubs. Phellogen also pericyclic. Vessel elements usually with simple (in Berberis sometimes scalariform) perforation plates. Leaves imparipinnate or unifoliolate, with conduplicate to curved ptyxis. Stipules numerous. Petiole vascular bundles in Berberis arcuate. Venation sometimes pinnate. Leaf margin usually glandular- or spinose-dentate (sometimes entire). Lowermost inflorescence bransch arising from axil of reduced leaf. Outer tepals three to twelve. Nectar often secreted from paired nectaries at base of four or six inner petaloid staminodia. Stamens (four to) six (to more than 20). Anthers usually dehiscing by valves (rarely slits), often sensitive. Tapetum amoeboid-periplasmodial. Pollen grains 6–12-colpate or spiraperturate etc. Exine striate-reticulate, often undifferentiated. Placentation basal-lateral; placentae sometimes protruding-diffuse and vascularized. Ovules one to numerous per carpel. Megasporocytes sometimes several. Parietal tissue approx. two cell layers thick. Endosperm development nuclear or cellular. Fruit a berry. Embryo elongate, with relatively long radicula. n = 7.

Cladogram of Berberidaceae based on DNA sequence data (Kim & Jansen 1998).

Cladogram of Berberidaceae based on DNA sequence data (Wang & al. 2007)

CIRCAEASTERACEAE Kuntze ex Hutch.

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Hutchinson, Fam. Fl. Pl. 1: 98. 15 Jan 1926, nom. cons.

Kingdoniaceae A. S. Foster ex Airy Shaw in Kew Bull. 18: 262. 8 Dec 1965; Circaeasterales Takht., Divers. Classif. Fl. Pl.: 97. 24 Apr 1997

Genera/species 2/2

Distribution Himalayas, southwestern, western and northwestern China.

Fossils Unknown.

Habit Bisexual, annual or perennial (Kingdonia) or annual (Circaeaster) herbs.

Vegetative anatomy Mycorrhiza endotrophic (Kingdonia). Phellogen absent. Vessel elements usually with simple (in Kingdonia sometimes scalariform) perforation plates, bordered pits absent? Imperforate tracheary xylem elements?, non-septate? Wood rays absent. Axial parenchyma absent? Sieve tube plastids? Nodes 1:1 (Circaeaster) or 1:1–4 (Kingdonia), unilacunar with one or four leaf traces, respectively. Crystals?

Trichomes Hairs unicellular or multicellular, uniseriate, uncinate (on surface of achenes in Circaeaster), or absent.

Leaves Alternate (in Circaeaster in basal rosette; in Kingdonia usually only a single basal leaf, sometimes distichous), simple, entire or palmately lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate? Venation flabellate (open-dichotomously branched), with branches running into leaf teeth. Stomata anomocytic. Cuticular wax crystalloids as clustered tubuli of Berberis type, dominated by nonacosan-10-ol (Circaeaster). Leaf margin serrate.

Inflorescence Flowers inCircaeaster in terminal, compound thyrsoid inflorescences; flowers in Kingdonia terminal, solitary, long-pedunculate (scapose). Floral prophyll (bracteole) in Kingdonia adaxial, in Circaeaster absent.

FlowersAlmost actinomorphic, small. Hypogyny. Tepals with valvate aestivation, spiral, persistent, in Circaeastertwo or three scale-like sepaloid, in Kingdonia (four or) five to seven petaloid. Kingdoniawith eight to 13 intratepalous clavate glistening glands (nectar-secreting staminodia?). Nectary absent in Circaeaster. Disc absent.

Androecium Stamens inCircaeaster usually two (sometimes one or three), alternisepalous, in Kingdonia three to eight. Filaments free from each other and from tepals. Anthers in Circaeaster basifixed, non-versatile, disporangiate, introrse to latrorse, in Kingdonia tetrasporangiate, extrorse, valvicidal (dehiscing by longitudinal valves). Tapetum secretory. Staminodia in Circaeaster one sepaloid or absent, in Kingdonia eight to 13 extrastaminal, apically nectariferous and petaloid.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (sometimes tricolporate?), shed as monads, bicellular at dispersal. Exine (tectate to) semitectate, with columellate infratectum, striate-reticulate (with compound layer of fine striae).

Gynoecium Carpels usually two (sometimes one or three; Circaeaster) or three to nine (Kingdonia), short- to long-stalked, free; carpel ascidiate, occluded by secretion? Ovary superior, unilocular (monomerous or apocarpous). Stylodia short (Circaeaster) or subulate and recurved (Kingdonia). Stigma somewhat oblique, papillate (Circaeaster), Wet? type. Pistillodium absent.

Ovules Placentation subapical-marginal. Ovules two submarginal (Circaeaster, upper one degenerating) or one (Kingdonia) per carpel, orthotropous (Circaeaster) or hemianatropous (Kingdonia), pendulous, unitegmic, incompletely tenuinucellar, with meiocyte hypodermal at apex of megasporangium. Integument in Circaeaster approx. two cell layers thick (degenerating following fertilization), in Kingdonia two to five cell layers thick. Megagametophyte in Kingdonia monosporous, Polygonum type, in Circaeaster tetrasporous, Adoxa type (tetranucleate or octanucleate). Endosperm formation at least in Circaeaster cellular (in Kingdonia helobial?). Endosperm haustorium chalazal (Circaeaster). Embryogenesis chenopodiad.

Fruit An achene or assemblage of achenes.

Seeds Aril absent. Testa thin or absent (degenerating; in Circaeaster replaced by inner epidermis of pericarp and outer layer of endosperm). Perisperm not developed. Endosperm copious. Embryo small (Circaeaster) or fairly large (Kingdonia), straight, chlorophyll? Hypocotyl in Circaeaster strongly elongated. Cotyledons two, linear, persistent (Circaeaster). Germination phanerocotylar (Circaeaster). Leaves in Circaeaster at apex of elongated hypocotyl.

Cytology n = 9 (Kingdonia), n = 15 (Circaeaster)

DNA

Phytochemistry Virtually unknown. Flavonols? Cyanogenic compounds not found.

Use Unknown.

Systematics Circaeaster (1; C. agrestis; northwestern Himalaya from Kumaun in India through Nepal to southeastern Tibet and northwestern Yunnan, and the Kansu and Shensi mountains in northwestern China), Kingdonia (1; K. uniflora; western and northwestern China).

Circaeasteraceae are sister group to Lardizabalaceae.

EUPTELEACEAE K. Wilh.

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Wilhelm, Samenpflanzen: 17. Oct 1910, nom. cons.

Eupteleales Hu ex Reveal in Phytologia 74: 174. 25 Mar 1993; Eupteleanae Doweld, Tent. Syst. Plant. Vasc.: xxv. 23 Dec 2001; Eupteleineae Shipunov in A. Shipunov & J. L. Reveal, Phytotaxa 16: 63. 4 Feb 2011

Genera/species 1/2

Distribution Eastern Himalaya, Assam, southwestern and central China, Japan.

Fossils Fossilized leaves, pollen grains and fruits have been found in Paleocene to Miocene layers in several places in the Northern Hemisphere.

Habit Usually bisexual (sometimes unisexual male), deciduous trees.

Vegetative anatomy Phellogen ab initio cortical. Vessel elements with scalariform or partially reticulate perforation plates; lateral pits usually opposite to intermediary (sometimes alternate or scalariform), bordered pits. Vessel restriction patterns absent. Imperforate tracheary xylem elements fibre tracheids with usually bordered (sometimes simple) pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, diffuse-in-aggregates, or in tangential bands. Multiseriate phloem rays strongly sclerified. Sieve tube plastids S type, with approx. ten globular starch grains. Nodes 1:5–11, unilacunar with five to eleven leaf traces. Medulla and petioles with secretory cells, tanniniferous cells and cells with cluster crystals.

Trichomes Hairs present on young leaves, uniseriate, caducous, or absent.

Leaves Alternate (spiral), simple, entire, with subplicate-conduplicate ptyxis. Stipules and leaf sheath absent. Petiole base hollow; petiole vascular bundles? Venation pinnate, craspedodromous, with lateral veins ascending and running almost to leaf teeth, or palmate. Stomata anomocytic. Cuticular wax crystalloids absent or as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Mesophyll without sclerenchymatous idioblasts. Calciumoxalate druses present. Idioblasts with ethereal oils absent. Leaf margin glandular-serrate, with platanoid teeth (each gland with an apical cavity).

Inflorescence Axillary, with flowers one or few together in raceme- or umbel-like fasciculate inflorescence. Lowermost flowers often with one or two floral prophylls (bracteoles).

Flowers Bisymmetric, small, bent downwards. Hypogyny? Tepals absent. Nectary absent. Disc absent.

Androecium Stamens six to c. 20, in one whorl. Filaments short or somewhat elongated, filiform, free. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, valvicidal (dehiscing by longitudinal valves), horizontally somewhat widened; connective prolonged. Tapetum secretory, with uninucleate to quadrinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate or penta- to heptacolpate (rarely tetracolpate), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate to microreticulate, verrucate.

Gynoecium Carpels six to 18 (to 31), whorled, free (apocarpous); carpel plicate and ascidiate (intermediary), postgenitally closed, without canal, stipitate. Ovary superior?, unilocular. Style absent. Stigma decurrent, not reaching carpellary apex (due to asymmetrical growth of carpel), brush-like, papillate (with long unicellular papillae), Dry or slightly Wet type. Pistillodium absent.

Ovules Placentation lateral (submarginal). Ovules usually one to three (rarely four) per carpel, anatropous, pendulous, apotropous or epitropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument two to five cell layers thick. Inner integument (except its inner epidermis) crushed, two or three cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis caryophyllad or solanad.

Fruit A stalked discoid samara with the brush-like stigma persistent on one side.

Seeds Aril absent. Seed coat testal. Epidermis tanniniferous. Exotestal cells enlarged (wider than endo- and mesotestal cells). Mesotesta often sclerotic. Endotesta subpalisade, with lignified cell walls. Tegmen unspecialized. Perisperm not developed. Endosperm copious, oily and proteinaceous. Embryo small, straight, little differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology n = 14

DNA The plastid gene rpl22 is perhaps absent from Euptelea. Nuclear gene AP3 triplicated?

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, chalcones, dihydrochalcones, and triterpene saponins present. Myricetin, ellagic acid and cyanogenic compounds not found.

Use Medicinal plants, timber.

Systematics Euptelea (2; E. pleiosperma: eastern Himalaya, Assam, southwestern and central China; E. polyandra: Japan).

Euptelea is sister to the remaining Ranunculales.

GLAUCIDIACEAE Tamura

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Tamura in Bot. Mag. (Tokyo) 85: 40. Mar 1972

Glaucidiales Takht. ex Reveal in Novon 2: 238. 13 Oct 1992

Genera/species 1/1

Distribution Japan.

Fossils Unknown.

Habit Bisexual, perennial herb.

Vegetative anatomyPhellogen? Primary vascular tissue consisting of two irregularly concentric cylinders of vascular bundles: one outer cylinder with smaller bundles and one inner cylinder with larger bundles. Endodermis absent. Palisade mesophyll absent. Rhizome with normal secondary lateral growth. Xylem not V-shaped in cross-section. Vessel elements usually with simple (sometimes scalariform with few bars or reticulate) perforation plates; lateral pits scalariform or pseudoscalariform (in reality alternate). Vessel restriction patterns occurring. Imperforate tracheary xylem elements usually libriform fibres (sometimes tracheids) with bordered pits. Wood rays multiseriate, heterocellular? (with non-lignified cell walls). Axial parenchyma paratracheal, pervasive or intervascular. Sieve tube plastids S type. Nodes multilacunar with several? leaf traces. Crystals absent?

Trichomes Hairs unicellular?

Leaves Alternate (distichous), simple, palmately lobed, with conduplicate, supervolute to curved and plicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular and medullary. Venation palmate. Stomata anomocytic. Cuticle wax crystalloids as few small irregular platelets, dominated by nonacosan-10-ol. Leaf margin serrate.

Inflorescence Flowers terminal, usually solitary (rarely pairwise).

Flowers Actinomorphic. Hypogyny. Tepals four, petaloid, decussate, caducous, free. Nectary absent. Disc absent.

Androecium Stamens c. 350 to more than 500, spiral, fasciculate. Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse?, 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 (with reduced columellae), spinulate, punctate-perforate.

Gynoecium Carpels (one or) two (to four), antesepalous, conduplicate-plicate, somewhat connate at base. Ovary superior, unilocular (apocarpy; bilocular at base). Stylodia very short. Stigma bifid, decurrent?, type? Pistillodium absent.

Ovules Placentation marginal. Ovules c. 15 to c. 20 (to more than 30) per carpel, anatropous, bitegmic, usually tenuinucellar (rarely pseudocrassinucellar). Micropyle endostomal. Outer integument seven to ten (to 13) cell layers thick, vascularized. Inner integument three to five cell layers thick. Archespore usually ten- to 15-celled (megasporocytes). Primary parietal cell not formed. Nucellar cap c. 15 to c. 20 cell layers thick, massive, formed by periclinal divisions from apical cells of megasporangial epidermis. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral or persistent (degenerating immediately after fertilization or earlier), non-proliferating. Endosperm development nuclear. Endosperm haustoria? Embryogenesis unclassified type (resembles onagrad). Polyembryony frequent.

Fruit Ventricidal and dorsicidal follicles. Adaxial side of carpels expanding more than abaxial side during fruit development, causing stigma to become inserted on ‘lower’ (abaxial) side at fruit maturation.

Seeds Aril absent. Seeds flattened. Seed coat testal. Outer integument vascularized, developing into a wing-like structure on testa. Inner integument degenerating. Tegmen collapsed? Perisperm not developed. Endosperm copious, starchy? Embryo small, elongate, well differentiated, chlorophyll? Cotyledons two, foliaceous, with more or less connate petioles. Germination?

Cytology n = 10 – Chromosomes 1,5–2,5 µm long, reniform, T type (Thalictrum type).

DNA Mitochondrial intron coxII.i3 lost?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin, rhamnocitrin) and glaupalol (a furanocoumarin) present. Berberine and other alkaloids, and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Glaucidium (1; G. palmatum; high mountains on Hokkaido and northern and central Honshu in Japan).

Glaucidium is sister to Hydrastis (Hydrastidaceae) or, alternatively, sister to Ranunculaceae.

HYDRASTIDACEAE Martinov

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Martinov, Tekhno-Bot. Slovar: 318. 3 Aug 1820 [’Hydrasteae’]

Hydrastidales Takht., Divers. Classif. Fl. Pl.: 98. 24 Apr 1997

Genera/species 1/1

Distribution Central and eastern parts of temperate North America.

Fossils Unknown.

Habit Bisexual, perennial herb. Roots and rhizome inside bright yellow due to presence of berberine. Erect stem with swollen nodes.

Vegetative anatomyPhellogen? Upright stem (not rhizome) multilacunar, with cortical vascular bundles. Medullary bundles present. Primary vascular tissue amphicribral, consisting of two irregularly concentric cylinders of vascular bundles: an outer cylinder with smaller bundles and an inner cylinder with larger bundles. Endodermis absent. Palisade mesophyll absent. Secondary lateral growth absent. Xylem not V-shaped in cross-section. Vessel elements usually with simple (rarely scalariform with a single bar, or reticulate) perforation plates; lateral pits scalariform, bordered pits? Imperforate tracheary xylem elements libriform fibres with bordered pits. Wood rays multiseriate, heterocellular? Axial parenchyma paratracheal? Sieve tube plastids Ss type. Nodes multilacunar with several leaf traces; lower leaves usually with 9-lacunar nodes, upper leaves with 3–5-lacunar nodes. Crystals?

Trichomes Hairs absent?

Leaves Alternate (distichous, usually one basal leaf and two stem leaves), simple, entire or usually palmately lobed, with plicate ptyxis. Stipules and leaf sheath absent. Petiole base enclosing rhizome. Petiole vascular bundles eight to 24, radially arranged (transection annular and medullary: medullary petiole bundles present). Venation palmate. Palisade mesophyll absent. Stomata anomocytic. Cuticular wax crystalloids as few small irregular platelets, dominated by nonacosan-10-ol. Leaf margin serrate to biserrate.

Inflorescence Flowers terminal, solitary.

Flowers Actinomorphic. Hypogyny. Tepals (two or) three (or four), petaloid, small, with imbricate aestivation, early caducous. Nectary absent. Disc absent.

Androecium Stamens c. 40 to c. 75, spiral, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse?, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, striate-reticulate or striate (with complex layer of striae).

Gynoecium Carpels five to c. 15, spiral, conduplicate, ab initio free, later connate at base. Ovary superior, unilocular (apocarpy). Stylodium very short. Stigma bifid, with multicellular processes, type? Pistillodium absent.

Ovules Placentation marginal. Ovules one or two (sometimes three or four) per carpel, anatropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument four to eight (to 13) cell layers thick. Inner integument two to four (or five) cell layers thick. Integument two or more cell layers thick, non-vascularized. Archespore probably unicellular. Hypostase absent. Nucellar cap approx. eight to ten cell layers thick, formed by periclinal divisions from apical cells of megasporangial epidermis. Megagametophyte monosporous, Polygonum type. Antipodal cells non-modified, proliferating (dividing into five to nine long-lived cells). Endosperm development nuclear. Endosperm haustoria? Embryogenesis?

Fruit A multifolliculus consisting of five to c. 15 usually one-seeded fleshy berry-like (drupaceous?) follicles, dehiscing adaxially and abaxially (cf. Glaucidium with follicle both dorsicidal and ventricidal).

Seeds Aril absent. Seed coat exotestal-exotegmic. Exotesta palisade, with very elongated cells, multiplicative, accumulating dark-brown substance. Exotegmen lignified, multiplicative. Remaining testal and tegmic cell layers parenchymatous and collapsed. Perisperm not developed. Endosperm copious, starchy? Embryo very small, straight, without chlorophyll. Cotyledons two. Germination?

Cytology n = 13 – Chromosomes small, reniform, T type (Thalictrum type).

DNA Mitochondrial intron coxII.i3 lost?

Phytochemistry Flavonols and isoquinoline alkaloids (berberine, hydrastine etc.) present. Rhizome containing a ribitole-like substance and benzylisoquinoline alkaloid compounds with D-galactose. Cyanogenic compounds not found.

Use Medicinal plant.

Systematics Hydrastis (1; H. canadensis; northeastern United States, southeastern Canada).

Hydrastis may be sister to Glaucidium (Glaucidiaceae), the two species together forming a sister-group of Ranunculaceae. The cuticular wax crystalloids in Hydrastis are very similar to those occurring in Glaucidium. Hydrastis has a special organization of those vascular strands that supply the stamens (more or less fascicled, supplied from vascular bundles of the central cylinder) and the carpels (each carpel supplied by four vascular bundles).


LARDIZABALACEAE R. Br.

( Back to Ranunculales )

Brown in Trans. Linn. Soc. London 13: 212. 23 Mai-21 Jun 1821 [‘Lardizabaleae’], nom. cons.

Sargentodoxaceae Stapf ex Hutch., Fam. Fl. Pl. 1: 100. 15 Jan 1926 [‘Sargentadoxaceae’], nom. cons.; Decaisneaceae (Takht. ex H. N. Qin) Loconte in H. Loconte, L. M. Campbell et D. W. Stevenson, Plant Syst. Evol. 9: 105. Dec 1995; Lardizabalales Loconte in D. W. Taylor et L. J. Hickey, Fl. Pl. Orig. Evol. Phylog.: 274. 21 Dec 1995; Sinofranchetiaceae Doweld in Byull. Mosk. Obshch. Ispyt. Prir., Biol. 105(5): 60. 9 Oct 2000; Lardizabalineae Shipunov in A. Shipunov et J. L. Reveal, Phytotaxa 16: 63. 4 Feb 2011

Genera/species 8/c 40

Distribution Southern and eastern Himalayas, East Asia, Indochina, southern South America.

Fossils Seeds of Sargentodoxa are known from the Miocene of North America, and seeds of Akebia and Decaisnea have been found in Miocene layers in Germany. Kajanthus lusitanicus was described from a late Aptian to early Albian layer in western Portugal. It is a trimerous, radially symmetrical, bisexual flower with several perianth whorls, six stamens in two whorls, and three free carpels.

Habit Usually monoecious or dioecious (in Decaisnea polygamomonoecious), usually climbing or scrambling evergreen or deciduous shrubs or lianas (Decaisnea an upright shrub).

Vegetative anatomy Phellogen ab initio usually superficial (in Sargentodoxa in innermost layer of pericycle). Primary vascular tissue consisting of a cylinder of vascular bundles. Primary medullary strands wide, usually lignified. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits usually alternate (in Decaisnea scalariform or transitional), usually with bordered (in Decaisnea simple) pits. Imperforate tracheary elements tracheids or fibre tracheids, usually with bordered pits (in Holboellia and often also in Decaisnea simple pits), septate or non-septate (also vasicentric tracheids). Wood rays usually multiseriate (in Decaisnea usually uniseriate), homocellular or heterocellular. Axial parenchyma apotracheal diffuse, paratracheal scanty vasicentric or banded, or absent. Wood elements (particularly in secondary phloem) partially storied (sometimes irregularly). Sieve tube plastids Ss type, very large. Phloem in Sargentodoxa with tanniniferous secretory cells. Nodes 3:3, trilacunar with three leaf traces. Medulla with sclerenchyma cells (not in Decaisnea). Prismatic calciumoxalate crystals sometimes abundant. Wood rays sometimes with rhomboidal crystals (Stauntonia).

Trichomes Hairs multicellular, uniseriate, or absent.

Leaves Alternate (spiral), usually palmately compound (in Decaisnea pinnately compound; in Sargentodoxa sometimes simple), entire or lobate, with conduplicate ptyxis; petiolules usually pulvinate at base. Stipules usually absent (present in Lardizabala); leaf sheath absent. Petiole vascular bundle transection arcuate. Venation usually palmate (in Decaisnea pinnate). Stomata usually anomocytic (in some species of Stauntonia cyclocytic). Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Mesophyll with calciumoxalate druses or single prismatic crystals. Idioblasts with ethereal oils absent. Leaflet margins entire or serrate with chloranthoid teeth.

Inflorescence Axillary, panicle or raceme. Floral prophylls (bracteoles) present or absent.

Flowers Actinomorphic. Hypogyny. Outer tepals (three to) 3+3 (to eight), with imbricate – or outermost ones with valvate – aestivation, whorled, petaloid, free. Inner tepals (nectariferous staminodia?) usually 3+3, whorled, petaloid (sometimes absent). Nectaries present on apex of inner tepals (nectariferous staminodia?) or stamens, or absent. Disc absent.

Androecium Stamens (three to) 3+3 (to eight), antepetalous, alternisepalous, occasionally somewhat foliaceous. Filaments free or connate into tube, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse or latrorse, longicidal (dehiscing by longitudinal slits); connective often elongated into prolonged appendage; microsporangia often sunken into almost foliaceous wide connective. Tapetum secretory, with usually binucleate (sometimes trinucleate or quadrinucleate) cells. Female flowers usually with six staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate to tricolporate (sometimes dicolpate to dicolporate, rarely syncolpate), shed as monads, bicellular at dispersal. Exine tectate, with sparsely columellate or almost acolumellate infratectum, reticulate, microreticulate, (micro)perforate, punctate, foveolate, psilate, striate, or smooth.

Gynoecium Carpels usually three or six to nine (to twelve) in one to five whorls of three (in Sargentodoxa c. 50 to c. 90, spiral), free; carpel plicate, postgenitally partially occluded, with open secretory canal; extragynoecial compitum sometimes present. Ovary superior, unilocular (apocarpy). Stylodia very short or absent. Stigma peltate, often oblique, hairy, non-papillate, usually Wet type. Male flowers sometimes with pistillodium.

Ovules Placentation usually laminal or laminal-lateral (in Decaisnea submarginal, with ovules along ventral line of fusion). Ovules usually numerous (sometimes few) per carpel (in Sargentodoxa a single subapical pendulous ovule), usually anatropous (in Boquila and Lardizabala hemitropous), horizontal, bitegmic, crassinucellar. Micropyle endostomal. Outer integument three to five cell layers thick. Inner integument two or three cell layers thick. Parietal tissue approx. three cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells usually not persistent. Endosperm development ab initio cellular. Endosperm haustoria? Embryogenesis?

Fruit An assemblage of fleshy (with fleshy placenta) follicles (Akebia, species of Decaisnea) or a berry with leathery pericarp. Fruit in Decaisnea with laticifers.

Seeds Aril usually absent (present in Akebia). Seed coat exotestal. Testa usually multiplicative. Exotesta palisade. Mesotesta and endotesta unspecialized. Tegmen unspecialized. Perisperm not developed. Endosperm copious, oily (sometimes, e.g. in Sargentodoxa, also starchy or with hemicellulose). Embryo small, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology n = 11 (Sargentodoxa), n = 14 (Lardizabala), n = 15 (Decaisnea), n = 16 (Akebia)

DNA The nuclear gene AP3 is triplicated.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, phenols, and triterpene saponins present. Ellagic acid, alkaloids and cyanogenic compounds not found. Aluminium accumulated in some species of Holboellia and Stauntonia.

Use Ornamental plants, medicinal plants, fruits (Boquila, Lardizabala).

Systematics Lardizabalaceae are sister-group to Circaeasteraceae.

Sargentodoxoideae (Stapf ex Hutch.) Thorne et Reveal in Bot. Rev. (Lancaster) 73: 89. 29 Jun 2007

1/1. Sargentodoxa (1; S. cuneata; China, Laos, Vietnam). – Usually dioecious (with few bisexual flowers). Phellogen ab initio inner-pericyclic. Tanniniferous cells present. Leaves trifoliolate. Outer tepals four to nine. Inner tepals five to seven. Staminodia petaloid. Carpels c. 50 to c. 90, spiral, ascidiate. Ovule one per carpel, pendulous. Outer integument approx. four cell layers thick. Placenta fleshy in fruit. Testal surface unspecialized. n = 11. Triterpenoid saponins not found.

Lardizabaloideae Burnett, Outlines Bot.: 830. Feb 1835 [‘Lardizabalidae’]

6/c 40. Decaisnea (1; D. insignis; eastern Himalayas to central China), Sinofranchetia (1; S. chinensis; western and central China), Lardizabala (1; L. funaria; central and southern Chile between the Andes and the Pacific, southern Argentina), Boquila (1; B. trifoliolata; central and southern Chile between the Andes and the Pacific, southern Argentina), Akebia (3–4; China, the Korean Peninsula, Japan, Taiwan), Stauntonia (30–32; northeastern India, Himalayas, China, the Korean Peninsula, Japan, Taiwan). – Southern and eastern Himalayas, East Asia, southern South America. Usually monoecious (rarely dioecious or bisexual). Phellogen ab initio superficial. Vessel elements sometimes with scalariform perforation plates. Stomata sometimes cyclocytic. Leaves sometimes palmately compound or imparipinnate, with basal tooth or lobe. Venation sometimes pinnate. Outer tepals sometimes three. Inner tepals sometimes absent. Stamens sometimes three or eight. Filaments connate. Tapetum with up to quadrinucleate cells. Pollen grains sometimes colporoidate, sometimes tricellular at dispersal. Female flowers with staminodia. Carpels sometimes up to twelve. Stigma sometimes peltate. Placentation sometimes laminar. Ovules (few to) numerous per carpel, sometimes hemitropous. Outer integument three to five cell layers thick. Inner integument two or three cell layers thick. Parietal tissue approx. three cell layers thick. Antipodal cells in Decaisnea persistent. Endosperm development usually cellular (in Decaisnea nuclear). Fruit a berry or a fleshy follicle, often with fleshy placenta. Testa multiplicative. Exotestal cells lignified, elongated (in Decaisnea), or non-lignified and fibrous (in Akebia and Stauntonia). Hypodermal cells thickened. Endosperm starchy or with hemicellulose. n = 14–16, 17?, 18. Oleanone triterpenoid saponins present. Aluminium accumulation occurs in Stauntonia, a synapomorphy of this clade.

Cladogram of Lardizabalaceae based on DNA sequence data (Hoot & al. 1995).

MENISPERMACEAE Juss.

( Back to Ranunculales )

de Jussieu, Gen. Plant.: 284. 4 Aug 1789 [’Menisperma’], nom. cons.

Menispermales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 225. Jan-Apr 1820 [‘Menispermeae’]; Pseliaceae Raf., New Fl. N. Amer. 4: 8. med 1838 [’Pselides’]

Genera/species c 73/c 430

Distribution Tropical and subtropical regions in the Northern and Southern Hemispheres, and a few species in temperate eastern North America and temperate East Asia.

Fossils The oldest fossil consists of keeled endocarps of Prototinomiscium from the Turonian to the Maastrichtian of Central Europe. Anamirta pfeifferi is fossilized wood (probably of a liana) from the Maastrichtian Deccan Intertrappean Beds in India. A large number of endocarps, which can be assigned to Menispermaceae, are known from Cenozoic layers in Europe and North America.

Habit Dioecious, usually evergreen lianas, or scrambling and climbing perennial herbs (rarely shrubs or trees [Burasaia, Penianthus, Sphenocentrum]; one species of Stephania an erect herb).

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Secondary lateral growth usually anomalous (particularly in lianas), from successive cambia. Vessel elements with simple perforation plates; lateral pits alternate or scalariform, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids or libriform fibres with simple and/or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate, interfascicular, usually wide and very tall (rarely narrow), homocellular or heterocellular, often lignified. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, scalariform or in short bands connecting successive layers of vascular bundles. Wood elements often storied, especially in secondary phloem. Intraxylary (concentric) phloem usually present. Sieve tube plastids Ss type, very large. Nodes 3:3, trilacunar with three leaf traces. Stem and leaves with rows of secretory cells. Parenchyma with numerous asterosclereids and osteosclereids. Crystal sand present or absent. Raphid idioblasts (raphid cells) present or absent. Prismatic calciumoxalate crystals abundant. Wood rays and axial parenchyma often with rhomboidal crystals.

Trichomes Hairs unicellular or multicellular, uniseriate.

Leaves Alternate (spiral), usually simple (in Burasaia trifoliolate), entire or lobed (one species of Cocculus with phyllocladia), often peltate, with ? ptyxis. Stipules usually absent; leaf sheath absent. Petiole often with proximal and distal pulvinus. Petiole vascular bundle transection annular. Venation palmate, actinodromous or acrodromous, or pinnate. Stomata anomocytic, paracytic, staurocytic or often more or less cyclocytic (sometimes actinocytic), often surrounded by a rosette of subsidiary cells. Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Domatia usually as pockets (rarely hair tufts). Epidermal cells often with calciumoxalate crystals, sometimes with silica or trichome hydathodes. Mesophyll with sclerenchymatous idioblasts and mucilage cells. Idioblasts with ethereal oils absent. Laminar surface sometimes with ridges of laticifers. Leaf margin usually entire (sometimes serrate or lobed). Extrafloral nectaries present or absent.

Inflorescence Terminal or axillary, raceme- or headlike, or compound panicle (flowers rarely solitary or paired).

Flowers Usually actinomorphic (female flowers in Antizoma, Cyclea, Cissampelos and some species of Stephania slightly zygomorphic). Hypogyny. Tepals usually whorled (sometimes spiral); in female flowers sometimes fewer than in male flowers. Outer tepals (one to) six (to more than twelve), with imbricate or valvate aestivation, sepaloid, usually in whorls each one with usually three (rarely one), usually free (in Cyclea and Synclisia slightly connate). Inner tepals (absent or one to) six (to eight), with usually imbricate aestivation, petaloid, usually free (in Cyclea and some species of Disciphania connate; rarely enclosing stamens). Nectary absent. Disc absent.

Androecium Stamens three, six, twelve or more (in one species of Odontocarya one; in Hypserpa up to c. 40), majority antepetalous, spiral or whorled. Filaments free or more or less connate into synandrium, free from tepals. Anthers basifixed, non-versatile, usually tetrasporangiate (rarely disporangiate), with superposed thecae, usually introrse (rarely extrorse), usually longicidal (dehiscing by longitudinal [rarely transversal] slits). Tapetum secretory. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colpate, (2–)3(–4)-colporate (sometimes syncolporate), (2–)3(–4)-porate or (2–)3(–4)-pororate (rarely cryptoporate or inaperturate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate or microreticulate.

Gynoecium Carpels usually three, six or more (rarely one or two; in one species of Tiliacora and one species of Triclisia c. 20 to 32), free or slightly connate at base, often stipitate (on gynophore); carpel plicate (ascidiate?), postgenitally partially fused, with open secretory canal. Ovary superior, unilocular (apocarpy). Style very short or absent. Stigma entire, bifid or trifid, expanded above, hairy to non-papillate, usually Dry (occasionally Wet) type. Male flowers often with pistillodium.

Ovules Placentation ventral-marginal. Ovules usually two per carpel (one of which degenerating), anatropous or campylotropous (often amphitropous after fertilization), apotropous or epitropous, pendulous to horizontal (ascending?), unitegmic (derived from integumentary shifting) or bitegmic, crassinucellar. Micropyle endostomal or bistomal, Z-shaped (zig-zag). Outer integument two to five cell layers thick. Inner integument two cell layers thick (when one integument, then three or four cell layers thick). Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells multinucleate, often proliferating. Endosperm development ab initio nuclear (finally cellular). Endosperm haustoria? Embryogenesis onagrad.

Fruit An assemblage of single-seeded stipitate drupes on a globose, discoid, columnar or branched carpophore, often flattened or strongly curved (cf. the name Menispermum = ’moon-seed’). Exocarp thin or leathery. Mesocarp fleshy to fibrous, sometimes sclerified. Endocarp sclerified, hard, usually sculptured in different ways, usually with a condyle (placentary outgrowth or invagination; sometimes absent; condyle produced during ovary development due to intruding ovary wall on placenta, causing seed to curve).

Seeds Aril absent. Seed usually curved. Testa little differentiated (sometimes absent). Exotesta sometimes tabular, lignified. Mesotesta and endotesta unspecialized. Tegmen unspecialized. Perisperm not developed. Endosperm usually copious (sometimes partially or entirely ruminate; absent in most genera in cladogram 2), oily. Embryo small to large, straight or curved, chlorophyll? Cotyledons usually two (rarely one), flat or terete, often fleshy (genera in cladogram 2). Germination phanerocotylar or cryptocotylar.

Cytology n = (9-)11-13 – Polyploidy (tetraploids, hexaploids) occurring.

DNA The nuclear gene AP3 is triplicated.

Phytochemistry Flavonols (kaempferol), flavones, diterpenoids, sesquiterpenoids, tannins, benzylisoquinoline and aporphine alkaloids (benzyltetrahydroisoquinoline and aporphine derivatives in dimeric form, e.g. berberine, morphinane), hasubanane alkaloids (protostephanines, erythrinanes, cocculolidines, morphines, quettamine-morphine dimers, hasubanonines, acutumines, etc.), azafluoranthene alkaloids, tropoloisoquinoline alkaloids, tubocurarine chloride (a curare mixture of alkaloids), toxic sesquiterpene lactones, phenylic cinnamide, furofuran lignans, and frequent cyanogenic compounds present. Tyrosine-derived cyanogenic glycosides and caffeic acid rare. Ellagic acid and proanthocyanidins not found.

Use Ornamental plants, medicinal plants (Anamirta cocculus, Chondrodendron tomentosum, Jateorhiza palmata etc), fish- and arrow poisons (curare from Chondrodendron, Curarea, Sciadotenia etc., coccel kernels and picrotoxine from Anamirta), timber.

Systematics Menispermaceae are sister-group to the clade [Berberidaceae+[Ranunculaceae+ [Hydrastis+Glaucidium]]].

Burasaia was sister to the remaining Menispermaceae, according to analyses of morphological data by Jacques & Bertolino (2008). Ortiz & al. (2007), using ndhF sequence data, recovered Tinomiscium sister to the remainder. Hoot & al. (2009), using atpB and rbcL sequence data, identified a clade comprising Menispermum and Sinomenium as sister to the rest with low to moderate support, and Burasaia was nested deep inside Menispermaceae. On the other hand, adding ndhF data rendered Tinomiscium again sister to the rest and Menispermum was nested deep within Menispermaceae.

Tinosporoideae W. Wang et Z. D. Chen in W. Wang et al., Perspect. Plant Ecol., Evol. Syst. 11: 100. 2009

Tinosporeae Hook. f. et Thomson, Fl. Ind. 1: 176, 179. 1-19 Jul 1855

c 24/c 130. Anamirta (1; A. cocculus; India, Southeast Asia, Malesia), Arcangelisia (2; Southeast Asia, Malesia), Coscinium (2; Southeast Asia, Malesia), Calycocarpum (1; C. lyonii; eastern North America), Jateorhiza (2; tropical Africa), Disciphania (c 25; tropical America), Aspidocarya (1; A. uvifera; northeastern India to southwestern China), Parabaena (6; Southeast Asia, Malesia), Tinomiscium (1; T. petiolare; Southeast Asia, Malesia), Burasaia (5; Madagascar), Borismene (1; B. japurensis; tropical South America), Orthogynium (1; O. gomphloides; Madagascar?), Fibraurea (2; Assam, Southeast Asia, the Philippines, Borneo, Sulawesi), Penianthus (4; tropical West and Central Africa), Sphenocentrum (1; S. jollyanum; tropical West Africa), Dioscoreophyllum (3; tropical Africa), Kolobopetalum (4; tropical Africa), Rhigiocarya (2; tropical West and Central Africa), ’Tinospora’ (32; tropical Asia, tropical Australia; polyphyletic), Chasmanthera (2; tropical Africa), Leptoterantha (1; L. mayumbense; tropical Africa), Platytinospora (1; P. buchholzii; tropical West and Central Africa), Odontocarya (c 30; tropical America), Syntriandrium (1; S. preussii; tropical West and Central Africa). – Pantropical. Tangential cell walls of wood rays in Tinomiscium oblique to ray axis in cross-section. Leaf surface sometimes (e.g. in Fibraurea and Tinomiscium) with laticifers as fine ridges. Seed subglobose to reniform, ruminate. Cotyledons foliaceous. – Tinomiscium is provisionally included here in Tinosporoideae, pending information from additional plastid and nuclear genes.

Menispermoideae Arn. in R. Wight et G. A. W. Arnott, Prodr. Fl. Ind. Orient.: 11. 10 Oct 1834 [’Menispermeae’]

c 50/c 300. Menispermeae DC., Syst. Nat. 1: 510, 511. 1-15 Nov 1817 [’Menispermeae verae’]. Menispermum (2–4; East Asia, eastern North America), Sinomenium (1; S. acutum; central China, Japan). – Diploclisia clade Diploclisia (2; southern China, Southeast Asia, Malesia). – Anomospermeae Miers in Ann. Mag. Nat. Hist., ser. 2, 7: 36. Jan 1851. Pericampylus (2–3; Southeast Asia, Malesia), Sarcopetalum (1; S. harveyanum; southern New Guinea, eastern Queensland, eastern New South Wales, eastern Victoria), ‘Abuta’ (32; tropical South America; polyphyletic), Caryomene (4; tropical America), ’Anomospermum’ (6; tropical America; polyphyletic), ’Orthomene’ (4; tropical America; non-monophyletic), Elephantomene (1; E. eburnea; northeastern South America), Telitoxicum (6; tropical South America), Legnephora (5; New Guinea, eastern Queensland), Hypserpa (c 7; Southeast Asia, Malesia and eastwards to Polynesia), Parapachygone (1; P. longifolia; northeastern Queensland). – Tiliacoreae Miers in Ann. Mag. Nat. Hist., ser. 2, 7: 36. Jan 1851. Limacia (3; Burma to West Malesia); ‘Stephania’ (c 35; tropical regions in the Old World; paraphyletic), Antizoma (2; arid regions in southern Africa), Rhaptonema (4; Madagascar), ‘Cissampelos’ (c 20; tropical regions on both hemispheres; paraphyletic; incl. Cyclea?), Cyclea (c 30; China and eastwards to the Philippines; in Cissampelos?), Haematocarpus (2; eastern Himalayas, Southeast Asia to Sulawesi), ’Hyperbaena’ (c 20; tropical and subtropical America; paraphyletic), Pachygone (c 10; southern China, Southeast Asia, Malesia and eastwards to islands in the western Pacific), Limaciopsis (1; L. loangensis; tropical Africa), Cocculus (8; tropical and southern Africa, Madagascar, Socotra, tropical and subtropical Asia and eastwards to northern Australia, subtropical to temperate North America), Strychnopsis (1; S. thouarsii; Madagascar), Chondrodendron (10; Central America, tropical South America), Sciadotenia (c 20; tropical America), Curarea (4; tropical South America), Carronia (4; New Guinea, eastern Queensland, eastern New South Wales), Pycnarrhena (9; Southeast Asia, Malesia to tropical Australia), Triclisia (10; tropical Africa, Madagascar), Synclisia (1; S. scabrida; Central Africa), Ungulipetalum (1; U. filipendulum; Brazil), Tiliacora (22; tropical regions in the Old World), Beirnaertia (1; B. cabindensis; tropical Africa), ‘Albertisia’ (17; tropical and subtropical Africa; paraphyletic; incl. Anisocycla?), Anisocycla (6; tropical Africa, Madagascar; in Albertisia?). – Pantropical, few species in temperate regions. Style lateral to basal. Endocarp often sculptured. Seed curved, usually not ruminate. Endosperm sometimes absent. Embryo curved. Cotyledons fleshy, cylindrical. – Menispermeae were sister-group (with relatively weak bootstrap support) to the remaining Menispermoideae in Hoot & al. (2009), and Anomospermeae successive sister-group to the rest.

Unplaced Menispermaceae Chlaenandra (1; C. ovata; New Guinea), Dialytheca (1; D. gossweileri; Angola), Eleutharrhena (1; E. macrocarpa; Assam, southwestern China), Macrococculus (1; M. pomiferus; New Guinea), Pleogyne (1; P. australis; eastern Queensland), Sarcolophium (1; S. tuberosum; tropical Africa), Spirospermum (1; S. penduliflorum; Madagascar), Synandropus (1; S. membranaceus; northeastern Brazil), Syrrheonema (3; tropical West and Central Africa).

Cladogram 1 of Menispermaceae based on morphological data (Jacques & Bertolino 2008).

Cladogram 2 of Menispermaceae based on morphological data (Jacques & Bertolino 2008).

Bayesian inference tree of Menispermaceae based on atpB/rbcL sequence data (Hoot & al. 2009).

Bayesian inference tree of Menispermaceae based on atpB, rbcL and ndhF sequence data (Hoot & al. 2009).

Bayesian inference tree of Menispermaceae based on DNA sequence data (Wefferling & al. 2013; Sciadotenia added).

PAPAVERACEAE Juss.

( Back to Ranunculales )

de Jussieu, Gen. Plant.: 235. 4 Aug 1789, nom. cons.

Corydalaceae Vest, Anleit. Stud. Bot.: 266, 283. 1818 [‘Corydaloidea’, ‘Corydaloideae’], nom. illeg.; Chelidoniaceae Martinov, Tekhno-Bot. Slovar: 124. 3 Aug 1820 [’Chelidoneae’]; Fumariaceae Marquis, Esq. Règne Vég.: 50. 15-22 Jul 1820 [‘Fumarieae’], nom. cons.; Fumariales Bercht. et J. Presl, Přir. Rostlin: 216. Jan-Apr 1820 [‘Fumariae’]; Papaverales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 216. Jan-Apr 1820 [‘Papaveraceae’]; Papaveropsida Brongn., Enum. Plant. Mus. Paris: xxv, 93. 12 Aug 1843 [’Papaverineae’]; Eschscholziaceae Ser., Fl. Jard. 2: 106. Apr 1847; Platystemonaceae (Spach) Lilja, Skånes Fl., ed. 2: 862, 979. Apr-Dec 1870 [’Platystemoneae’]; Hypecoaceae (Dumort.) Willkomm et Lange, Prodr. Fl. Hispan. 3: 875. Apr-Mai 1880 [’Hypecoëae’]; Papaverineae Thorne et Reveal in Bot. Rev. (Lancaster) 73: 91. 29 Jun 2007

Genera/species 43/c 755

Distribution Temperate and subtropical parts if the Northern Hemisphere, Central America, northern South America, East African mountains, southern Africa, Macaronesia, with their largest diversity in the Mediterranean, West, Central and East Asia, and the southwestern United States.

Fossils Uncertain. No entirely convincing reports, although some Miocene seeds are similar to Corydalis.

Habit Bisexual, usually perennial, biennial or annual herbs (sometimes climbing; some species of Argemone, Bocconia, Dendromecon, Hunnemannia, and Romneya evergreen shrubs or small trees). Some species have tuberous rhizome or tuberous roots. Stem sometimes pachycaul.

Vegetative anatomy Mycorrhiza at least usually absent. Roots diarch and lateral roots tetrastichous. Phellogen? Primary vascular tissue consisting of one or several cylinders of vascular bundles; cross section of stem usually with a ring of loose collateral vascular bundles separated by wide multiseriate medullary rays. Cortex and medulla without vascular bundles. Secondary lateral growth usually absent (sometimes from normal cylindrical cambium). Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate (also vasicentric tracheids). Wood rays multiseriate, heterocellular. Axial parenchyma paratracheal scanty, vasicentric. Wood elements often storied. Sieve tube plastids Ss type. Nodes 1:1 or ≥3:≥3, unilacunar or trilacunar (rarely multilacunar) with one or more leaf traces. Latex cells and laticifers often articulated, often anastomosing, usually with white, yellow, orange or red latex, numerous in most genera of Papaveroideae. Idioblasts with watery secretions present in Fumarioideae. Crystals absent.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate (in Bocconia and Macleaya sometimes branched), or absent (multicellular glandular hairs present in Dicranostigma and Glaucium).

Leaves Usually alternate (spiral; rarely opposite or verticillate), simple or compound (usually once or twice pinnately compound), entire or lobed (usually pinnately lobed), with ptyxis of various types. Stipules and leaf sheath absent (petiole base occasionally somewhat sheathing). Petiole vascular bundle transection arcuate. Venation usually pinnate (sometimes palmate). Stomata anomocytic. Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Hydathodes present in some species. Idioblasts with ethereal oils absent. Leaf margin serrate (with chloranthoid teeth, often incised), lobed or entire.

Inflorescence Terminal or axillary, simple or compound, cymose or racemose of different shapes, or flowers solitary terminal.

Flowers Actinomorphic, bisymmetric or zygomorphic (transversely zygomorphic in most Fumarioideae). Hypanthium usually absent (present in Platystemon). Usually hypogyny (in Platystemon half epigyny). Outer (sepaloid) tepals usually two or three (rarely four), with imbricate or open aestivation, median, in one whorl, usually caducous, usually free (in Eomecon and Eschscholzia connate either at base or entirely into operculum), with or without basal spur or other type of appendage. Inner (petaloid) tepals usually 2+2 or 3+3, crumpled or with imbricate aestivation, whorled (in Sanguinaria six to twelve, often in several whorls; absent in Bocconia and Macleaya), usually free (in Fumarioideae more or less connate and adnate to stamens), usually caducous, without spur, in Fumarioideae one or two petals usually spurred. Nectaries in Papaveroideae absent, in Fumarioideae present at staminal bases. Disc absent.

Androecium Stamens usually 16 up to more than 200 (in Meconella four to six or 6+6; in Canbya six to twelve; in Hypecoum 2+2; in Romneya up to c. 700) in three to 15 whorls; androphore present or absent. Filaments filiform, clavate or wide (in some species of Dicentra petaloid), usually free (in Fumarioideae two free stamens and two groups of connate stamens opposite outer petaloid tepals, each group consisting of one median entire and two lateral half anthers), free from tepals. Anthers basifixed, non-versatile, usually tetrasporangiate (in Hypecoum sometimes disporangiate; in Fumarioideae lateral anther halves in each staminal group disporangiate and remaining anthers tetrasporangiate), usually extrorse (rarely latrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory or amoeboid-periplasmodial. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate to hexacolpate, syncolpate, polyporate or polyforate (sometimes polycolpate, pantoaperturate; in some species of Meconopsis inaperturate; in Dicentra, Hypecoum, and Platystemon dicolpate; in Fumarieae often polypantoporate, colliculate, with intine bulging from pores), shed as monads, usually bicellular at dispersal (in Papaver sometimes tricellular). Exine tectate or semitectate, with columellate infratectum (columellae in Fumarioideae small), reticulate, microreticulate, perforate, microperforate or punctate, psilate, verrucate or echinulate (sometimes spinulate).

Gynoecium Pistil composed of two (up to c. 25) paracarp and connate carpels (carpels in Platystemon etc. partially free; carpels in Fumarioideae two transverse); carpel plicate, seemingly occluded by secretion. Ovary usually superior (in Platystemon semi-inferior), usually unilocular (sometimes seemingly bilocular to 20-locular by intrusion of placentae), stipitate or sessile (gynophore present or absent). Style single, simple, short, with stylar canal (sometimes closed by hairs), or absent (stylodia in Hypecoum two, partially connate). Stigma carinal and/or commissural, capitate, or stigmas often connate and forming a disc-like roof on top of ovary, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually parietal (when ovary unilocular) or axile (when ovary multilocular) (placentae in Bocconia basal; in Romneya plurilocular ovary with parietal placentation; placentae sometimes protruding to diffuse). Ovules one to c. 100 per ovary (in Bocconia and one speces of Macleaya one ovule), anatropous, hemicampylotropous to campylotropous or hemiamphitropous, pendulous?, horizontal or ascending (with ventral raphe, collateral or superposed or above carpellary surface or biseriate), bitegmic, usually crassinucellar (in Papaver and Platystemon pseudocrassinucellar). Micropyle bistomal (sometimes Z-shaped, zig-zag). Outer integument (two to) four to ten cell layers thick. Inner integument two to four cell layers thick. Parietal tissue two to four cell layers thick. Nucellar cap approx. three cell layers thick. Megagametophyte usually monosporous, Polygonum type (in Platystemon tetrasporous, Fritillaria type). Synergids usually with a filiform apparatus. Antipodal cells usually three (in Papaver sometimes five), non-proliferating, persistent, endopolyploid (sometimes binucleate). Endosperm development ab initio nuclear. Suspensor haustorium present in Fumarioideae. Endosperm haustoria absent? Embryogenesis irregularly solanad (Papaveroideae) or caryophyllad (Fumarioideae).

Fruit A poricidal, septicidal (placenticidal) or loculicidal capsule with basipetalous or acropetalous dehiscence, sometimes siliquiform with replum (in Fumaria and one species of Macleaya a nut; in Platystemon and species of Hypecoum lomentum-like with two to c. 30 one-seeded nut-like mericarps).

Seeds Aril present or absent. Elaiosome present in numerous species (in, e.g., Chelidonium, Corydalis, Dendromecon); strophiole (tiny swellings of raphe) sometimes (in, e.g., Chelidonium) present. Seed coat testal-tegmic. Exotesta usually well developed, palisade. Endotesta often well developed, with calciumoxalate crystals and coarse fibrillar endoreticulum. Exotegmen often fibrous. Endotegmic cell walls thickened. Perisperm not developed. Endosperm copious, oily (rarely granular). Embryo small, straight or curved, well developed or poorly developed (seed in many Fumarioideae containing only proembryo, further growth being intraseminal), without chlorophyll. Cotyledons usually two (in some species of Dicentra and in tuberous species of Corydalis one). Germination phanerocotylar or cryptocotylar.

Cytology n = 6–11, (14, 19, 21) – Polyploidy frequently occurring (up to 2n = c. 140).

DNA Duplication of nuclear CYCLOIDEA genes

Phytochemistry Flavonols (kaempferol, quercetin), benzylisoquinoline alkaloids (e.g. 1-benzyltetrahydroisoquinoline alkaloids) and aporphine alkaloids (dehydrogenated benzophene anthridines, reduced benzophene anthridines in Chelidonieae, berberines, tetrahydroberberines, stylopine, chelidonine, protopines, rhoeadines, narceines, spirobenzylisoquinolines, etc. in Fumarioideae, aporphines, morphinanes, pavines, isopavines, reticuline, scoulerine, narcotine), meconic acid (in Meconopsis, Papaver and Roemeria), chelidonic acid (at least in Chelidonium and Stylophorum), fumaric acid (in Glaucium, Papaver and Fumarioideae), and nitrophenyl ethan present. Tyrosine-derived cyanogenic glycosides sometimes present. Ellagic acid, tannins and proanthocyanidins not found. Caffeic acid? The free amino acid δ-acetylornithine is probably the principal nitrogen transport compound in Fumarioideae.

Use Ornamental plants, baking (seeds from Papaver somniferum), seed oils for soap (Argemone, Glaucium, Papaver), medicinal plants, narcotics (opium from Papaver somniferum and P. bracteatum).

Systematics Papaveraceae are probably sister to Pteridophyllum (Pteridophyllaceae).

Papaveroideae Eaton, Bot. Dict., ed. 4: 38. Apr-Mai 1836 [‘Papaveraceae’]

23/c 235. Mainly temperate regions on the Northern Hemisphere. Usually herbs (rarely small trees). Nodes 1:1 or ≥3:≥3, unilacunar or trilacunar (rarely multilacunar) with one or more leaf traces. Latex milky white, yellow, orange or red. Leaves with ptyxis of various types. Colleters present. Flowers usually large. Outer tepals enclosing bud, caducous. Inner (petaloid) tepals four or six, wrinkled, usually early caducous (rarely absent). Nectary absent. Stamens (four to) numerous (possibly evolved from a small number), in multiples of two or three. Style present or absent. Stigmas often connate, Dry type. Placentation sometimes axile or almost axile. Ovules numerous per carpel, anatropous or campylotropous. Hypostase present. Outer integument (two to) four to ten cell layers thick. Inner integument two to four cell layers thick. Parietal tissue two to four cell layers thick. Nucellar cap approx. three cell layers thick. Antipodal cells uninucleate or multinucleate. Embryogenesis irregularly solanad. Capsule often also transversely dehiscing, sometimes siliquiform with replum (rarely a nut or schizocarp). Exotegmen often with thickened outer cell walls, unlignified; anticlinal cell walls sometimes sinuate. Endotegmen usually persistent. n = 5–10 (14, 19). Duplication of nuclear gene PAPACYL. – The gametophytic self-incompatibility system in Papaveraceae is non-RNase-based (differing from other Triaperturatae).

Eschscholzieae Baill., Hist. Plant. 3: 130, 142. 1871

3/13–16. Dendromecon (1–2; California, northwestern Mexico), Eschscholzia (10–12; western North America), Hunnemannia (2; eastern Mexico). – Western North America, northern and eastern Mexico. Stem with subepidermal collenchyma. Nodes usually 1:1. Hairs unicellular. Flowers in Eschscholtzia shortly perigynous. Pollen grains polycolpate. Outer integument in Dendromecon seven cell layers thick. Capsule explosively dehiscent from base. – Eschscholzieae are sister-group to [Papavereae+Chelidonieae].

Papavereae Dumort., Fl. Belg.: 130. 1827

11/c 170. Arctomecon (3; Mojave Desert in the United States), Argemone (23; North America, the West Indies, South America, the Hawaiian Islands), Meconopsis (c 50; Himalayas to western China), ‘Papaver’ (c 80; Europe, the Cape Verde Islands, the Mediterranean, southern Africa, temperate and subtropical Asia, western North America; paraphyletic?), Roemeria (3; the Mediterranean and eastwards to Afghanistan), Romneya (1–2; R. coulteri; southern California, northern Baja California), Canbya (2; western North America), Hesperomecon (1; H. linearis; southern Oregon, California), Meconella (3; western United States), Platystemon (1; P. californicus; western United States, Baja California). – Temperate regions on the Northern Hemisphere. Nodes often 1:1 or 3:3. Hairs often multicellular, multiseriate. Hypogyny. Pistil composed of (three or) four (diagonally arranged) to numerous (in Platystemon up to c. 25) usually connate (in Platystemon free) carpels. Style sometimes present. Epistase present. Megasporocytes sometimes several. Capsule often poricidal, often with replum. Benzylisoquinoline alkaloids in Papaver synthesized in sieve tubes. – Papavereae are sister to Chelidonieae.

Chelidonieae Dumort., Fl. Belg.: 130. 1827

9/47–49. Sanguinaria (1; S. canadensis; eastern North America), Bocconia (10; tropical and subtropical regions in America), Chelidonium (1; C. majus; Europe, temperate and subarctic Asia), Dicranostigma (3–5; Himalayas, western China), Eomecon (1; E. chionantha; eastern China), Glaucium (23; Europe, the Mediterranean, southwestern to Central Asia), Hylomecon (3; temperate East Asia), Macleaya (2; temperate East Asia), Stylophorum (3; East Asia, eastern United States). – Europe, Asia, eastern North America, Central and South America, the West Indies. Latex yellow, orange, or red. Nodes 3–5(–9):3–5(–9). Hairs multicellular, uniseriate at apex. Perianth in Sanguinaria dimerous. Pollen grains also polyporate. Pistil sometimes composed of three connate carpels. Gynophores present in Bocconia. Ovules sometimes one per carpel (in Bocconia basal). Fruit elongate. Aril often present. δ-acetylornithine present. – Chelidonieae are sister to Papavereae.

Fumarioideae Eaton, Bot. Dict., ed. 4: 46. Apr-Mai 1836 [‘Fumariaceae’]

20/c 520. Mainly temperate regions in the Northern Hemisphere, southern Africa. Nodes usually 1:1, unilacunar with one leaf trace (rarely multilacunar with up to five traces). Stem with subepidermal collenchyma. Latex absent; watery juice present in often non-articulated idioblasts (probably reduced laticifers). Flowers transversely zygomorphic or bisymmetric. Tepals in multiples of two. Outer (sepaloid) tepals minute, not enclosing inner tepals. Inner (petaloid) tepals four, often spurred. Stamens four or six, free or connate in groups. Nectaries inserted at staminal bases. Pollen grains sometimes tricolporoidate. Exine often spinulate. Secondary pollen presentation frequent. Style present, often long. Stigma very variable. Placentation sometimes axile. Ovules one to numerous per carpel, campylotropous. Outer integument two to four cell layers thick. Inner integument approx. two cell layers thick. Parietal tissue approx. four cell layers thick. Nucellar cap usually absent. Suspensor haustorium present. Embryogenesis caryophyllad. Fruit sometimes lomentum-like or a nutlet. Seed curved. Exotesta often palisade. Endotesta without fibrillar reticulum. Exotegmen not fibrous. Embryo sometimes elongate. n = (6–)8 (or more). Coularine, δ-acetylornithine, and sometimes berberine present.

Hypecoeae Dumort., Fl. Belg.: 130. 1827

1/c 20. Hypecoum (c 20; the Mediterranean and eastwards to Central Asia, western and northern China). – Outer tepals two, sepaloid, anterio-posterior, with open aestivation. Inner tepals petaloid; outer ones not spurred; inner ones trilobate. Stamens 2+2. Anthers latrorse, with monothecal median anthers fused in pairs. Secondary pollen presentation: pollen deposited in bud within pockets inside innermost tepals, which close prior to stigmatic maturation, and open at touch of pollinators, these then becoming dusted with pollen. Pollen grains dicolpate. Stylodia two, partially connate. Inner integument approx. three cell layers thick. Fruit usually an elongate schizocarp detaching into one-seeded units (resembling a lomentum), sometimes siliqua-like, acropetalous, and with replum and persistent placentae. Seed coat with rectangular crystals. Suspensor consisting of two massive cells. n = 9. – Pteridophyllum was identified as sister to Hypecoum with fairly high bootstrap support in Wang & al. (2009), using both morphological and a number of molecular data. On the other hand, there are arguments from many other analyses that Pteridophyllum is instead sister to the entire Papaveraceae.

Fumarieae Dumort., Anal. Fam. Plant.: 51. 1829

19/c 500. Lamprocapnos (1; L. spectabilis; Siberia, northern China, the Korean Peninsula, Japan), Ehrendorferia (2; California), Ichtyoselmis (1; I. macrantha; southern China, northern Burma), Dicentra (10; temperate Asia and North America), Adlumia (1; A. fungosa; eastern North America), Capnoides (1; C. sempervirens; northeastern North America), Corydalis (c 400; temperate regions on the Northern Hemisphere, tropical African mountains), Dactylicapnos (10; Himalayas to Southeast Asia), Cysticapnos (4; southern Africa), Discocapnos (1; D. mundtii; Western Cape), Trigonocapnos (1; T. lichtensteinii; southern Africa), Cryptocapnos (1; C. chasmophytica; Afghanistan), Fumaria (c 50; Europe, the Mediterranean and eastwards to Central Asia and Himalayas, mountains in tropical East Africa), Rupicapnos (7; southern Spain, northwestern Africa), Pseudofumaria (2; southern Alps, northwestern Balkan Peninsula), Platycapnos (3; Macaronesia, western Mediterranean), Fumariola (1; F. turkestanica; Central Asia), Ceratocapnos (3; western Europe, the Mediterranean), Sarcocapnos (4; western Mediterranean). – Mainly temperate regions in the Northern Hemisphere (with their highest diversity in Himalaya and southwestern China), southern and eastern Africa. Nodes sometimes with up to five traces. Inflorescence often racemose. Flowers transversely zygomorphic or bisymmetric. Outer (sepaloid) tepals minute. One or two outer petaloid tepals spurred; inner petaloid tepals connate at apex, not lobate. Stamens in two groups of three: two median dithecal and four lateral monothecal anthers (some species of Dicentra with six separate stamens). Secondary pollen presentation: pollen deposited on stigma. Stigma Wet type, sometimes flattened, with marginal lobes. Ovule often one per carpel. Fruit often a nut. Elaiosome often present. Exotesta usually pigmented. Endotesta sometimes palisade, usually not crystalliferous. Suspensor cells resembling a bunch of grapes. Embryo sometimes undifferentiated.

Phylogeny of Papaveraceae and Pteridophyllum (Papaveroideae: strict consensus tree based on morphology and DNA sequence data, Hoot & al. 1997; Fumarioideae: 50% majority rule bootstrap tree based on DNA sequence data, Lidén & al. 1997; Discocapnos to Sarcocapnos: Bayesian consensus tree of ITS and cpDNA sequences, Pérez-Gutiérrez & al. 2012).

PTERIDOPHYLLACEAE (Murb.) Nakai ex Reveal et Hoogland

( Back to Ranunculales )

Reveal et Hoogland in Bull. Mus. Natl. Hist. Nat. (Paris), sér. 4, sect. B, Adansonia, 13: 91. 4 Oct 1991

Genera/species 1/1

Distribution Japan.

Fossils Unknown.

Habit Bisexual, perennial herbs. Blackening when drying.

Vegetative anatomy Mycorrhiza absent. Roots diarch. Phellogen? Secondary lateral growth absent. Vessel elements with simple perforation plates; lateral pits? Imperforate tracheary xylem elements libriform fibres with simple pits. Wood rays absent? Axial parenchyma paratracheal. Sieve tube plastids Ss type. Nodes? Watery juice in non-articulate idioblasts, probably representing reduced laticifers? Crystals?

Trichomes Hairs multicellular, pointed.

Leaves Alternate (spiral, in basal rosette), pinnately compound (with ten to c. 20 pairs of leaflets) to pinnetely lobed, with ? ptyxis. Leaf base surrounded by several large round membranous cataphylls. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? (as platelets?). Mesophyll without sclerenchymatous idioblasts. Leaflet margins serrate-dentate (glandular-serrate?).

Inflorescence Terminal, one- to four-flowered, raceme-like (scapiflorous with separate one- to four-flowered cyme).

Flowers Bisymmetric or slightly zygomorphic. Pedicel thin. Hypogyny. Outer tepals two, with open aestivation, median, petaloid, caducous, free. Inner tepals 2+2, petaloid, outer ones with imbricate aestivation, caducous, free. Nectary? Disc absent.

Androecium Stamens four (lateral two stamens absent; rarely six), in two alternipetalous diagonally arranged groups. Filaments narrow, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolpate (rarely di- or tetracolpate), shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, perforate, echinulate.

Gynoecium Pistil composed of two connate transverse carpels; carpel occluded by secretion. Ovary superior, unilocular. Style single, simple, long. Stigmatic lobes commissural; stigmatic area bifid, densely papillate, Dry? type. Pistillodium absent.

Ovules Placentation parietal. Ovules usually one (rarely two) per carpel, anatropous (or amphitropous) to almost campylotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Antipodal cells persistent? Endosperm development nuclear. Endosperm haustoria? Embryogenesis?

Fruit A many-seeded, siliqua-like, septicidal (placenticidal) capsule with persistent placental strands, acropetally dehiscing.

Seeds Aril absent. Seed coat endotestal. Exotesta more or less collapsing. Endotesta well developed, with coarse network of cellulose fibrils. Tegmen thin. Perisperm not developed. Endosperm copious (starchy?). Embryo small, chlorophyll? Cotyledons two. Germination?

Cytology n = 9

DNA

Phytochemistry Very insufficiently known. Protopine present. Flavonols? Berberine?

Use Ornamental plant.

Systematics Pteridophyllum (1; P. racemosum; Honshu and Hokkaido? in Japan).

Pteridophyllum is probably sister to Papaveraceae. However, according to molecular-morphological analyses by Wang & al. (2009), Pteridophyllum is sister to Hypecoum and should be included in Fumarioideae-Hypecoeae.

RANUNCULACEAE Juss.

( Back to Ranunculales )

de Jussieu, Gen. Plant.: 231. 4 Aug 1789, nom. cons.

Thalictraceae Raf., Anal. Nat.: 176. Apr-Jul 1815 [‘Thalictria’]; Anemonaceae Vest, Anleit. Stud. Bot.: 264, 275. 1818 [’Anemonoideae’]; Helleboraceae Vest, Anleit. Stud. Bot.: 264, 274. 1818 [‘Helleboroideae’]; Aconitaceae Bercht. et J. Presl, Přir. Rostlin: 216. Jan-Apr 1820 [‘Aconiteae’]; Calthaceae Martinov, Tekhno-Bot. Slovar: 92. 3 Aug 1820 [‘Calthoideae’]; Clematidaceae Martinov, Tekhno-Bot. Slovar: 134. 3 Aug 1820 [‘Clematites’]; Actaeaceae Bercht. et J. Presl, Přir. Rostlin 1 (16*-53): 2, 140. 1823; Xanthorhizaceae Bercht. et J. Presl, Přir. Rostlin 1(16*-53): 2, 145. 1823 [‘Xanthorhizeae’]; Cimicifugaceae (Arn.) Bromhead in Mag. Nat. Hist., n. s., 4: 336, 338. Jul 1840 [’Cimicifugeae’]; Ranunculineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7:2, 10. 1846 [’Ranunculeae’]; Nigellaceae J. Agardh, Theoria Syst. Plant.: 76. Apr-Sep 1858; Aquilegiaceae Lilja, Skånes Fl., ed. 2: 375, 861, 979. Apr-Dec 1870; Delphiniaceae Brenner, Florist. Handb.: 52. 1886; Helleborales Nakai in J. Jap. Bot. 24: 10. Dec 1949

Genera/species 47–49/2.150–2.250

Distribution Cosmopolitan, with their largest diversity in temperate regions in the Northern and Southern Hemispheres.

Fossil Numerous fossilized fruits and seeds and a few records of fossil leaves are known from Cenozoic sediments.

Habit Usually bisexual (rarely dioecious), usually perennial, biennial or annual herbs (in Xanthorrhiza suffrutices; in Clematis lianas). Some species of Ranunculus and Caltha are aquatic. Main root often early withering and replaced by adventitious roots; some species with tuberous roots. Roots and rhizomes in Xanthorhiza and Coptis and bark in Xanthorhiza intensively yellow-coloured due to presence of berberine (a major alkaloid).

Vegetative anatomyPhellogen ab initio deeply seated. Medullary, primary, rays (in ligneous representatives) wide, persistent. Primary vascular tissue consisting of one or several cylinders of vascular bundles, or as scattered bundles, atactostele. Secondary lateral growth usually absent (sometimes from normal cylindrical cambium). Xylem usually V-shaped in cross-section, surrounding phloem. Vessel elements with simple and/or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple pits, non-septate (vasicentric tracheids usually absent, although present in many species of Clematis, especially in temperate regions). Wood rays multiseriate, heterocellular, or absent. Axial parenchyma apotracheal intervascular or scanty vasicentric (Clematis) (paratracheal parenchyma absent or pervasive). Wood elements sometimes (Clematis) partially storied. Phloem surrounded by xylem. Sieve tube plastids Ss type. Nodes usually 3:3, trilacunar with three leaf traces, or ≥5:≥5, multilacunar with at least five traces (rarely 1:1, unilacunar with one trace, or 2:2, bilacunar with two traces). Calciumoxalate crystals rhomboidal or absent.

Trichomes Hairs unicellular och multicellular, uniseriate; glandular hairs often present.

Leaves Usually alternate (spiral or sometimes distichous; in Clematis opposite), usually compound (sometimes simple, sometimes two to several times pinnately compound), usually lobed (sometimes entire); lamina and/or lobes with supervolute, involute, curved-involute, or conduplicate ptyxis. Stipules intrapetiolar or absent; petiole base often sheathing. Petioles in Clematis modified into tactile tendrils. Petiole vascular bundle transection annular or arcuate. Venation usually palmate (rarely parallelodromous; in some species of Ranunculus flabellate, with open dichotomously branched veins). Stomata usually anomocytic (sometimes paracytic), sometimes on adaxial side of leaf only, or absent. Cuticular wax crystalloids usually as irregularly shaped platelets (sometimes as clustered tubuli of Berberis type), chemically dominated by nonacosan-10-ol, or absent. Hydathodes present in some representatives. Idioblasts with ethereal oils absent. Leaf margin usually glandular serrate, with chloranthoid teeth (sometimes crenate or entire). Extrafloral nectaries present or absent.

Inflorescence Terminal or axillary, usually cymose, simple or compound, of different types (rarely racemes), or flowers solitary terminal. Bracts in Anemone often sepaloid.

Flowers Usually actinomorphic (in Aconitum and Delphinium zygomorphic). Receptacle usually elongate (in Myosurus strongly prolonged). Hypogyny. Tepals spiral or whorled. Outer tepals usually five (sometimes two to four, six or more), with imbricate or valvate aestivation, sepaloid or petaloid, persistent or early caducous, usually free. Inner tepals absent or one to 13, often interpreted as staminodia, usually petaloid, with imbricate or valvate aestivation, free, often (e.g. in Ranunculus) with one or several nectaries at base (in Ranunculus giganteus up to c. 30) as nectar-producing pockets or spurs (in Aquilegia five spurs; nectariferous spurs in Delphinium pairwise). Nectariferous hairs present on carpels in Caltha. Disc absent. Androgynophore present in some species of Ranunculus.

Androecium Stamens usually (secondarily?) abundant (rarely few; in Laccopetalum up to more than 2.000), usually (secondarily?) spiral (rarely whorled). Filaments free from each other and from tepals. Anthers basifixed to somewhat dorsifixed, usually non-versatile, tetrasporangiate, extrorse, latrorse or introrse, usually longicidal (dehiscing by longitudinal slits, rarely by longitudinal valves). Tapetum usually secretory (in Actaea sometimes amoeboid-periplasmodial), with binucleate to quadrinucleate cells. Staminodia extrastaminal, petaloid and/or nectariferous, or absent.

Pollen grains Microsporogenesis usually simultaneous (in Thalictrum successive). Pollen grains tri-, hexa- or polycolpate or -porate (zono- or pantoaperturate or syncolpate; rarely spiraperturate; in ’Souliea’ inaperturate), shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine usually tectate (rarely semitectate), with columellate infratectum, usually perforate and scabrate or micro-echinate, sometimes spinulate (larger columellae projecting through tectum) or punctate (in Helleborus reticulate; in Trollius striate).

Gynoecium Carpels one to more than 100 (in Ranunculus giganteus to more than 10.000), spiral or whorled, usually free (sometimes, e.g. in Nigella, more or less connate, compitum absent); carpel conduplicate-plicate, postgenitally usually entirely fused, without canal (sometimes ascidiate). Ovary superior, unilocular (apocarpy or gynoecium monomerous) or trilocular to quinquelocular. Stylodia short to long, or absent. Stigmas of different types, papillate or non-papillate, Dry type. Pistillodium absent.

Ovules Placentation marginal or basal (when ovary unilocular) or axile (when ovary plurilocular). Ovules one to numerous per carpel, usually anatropous (sometimes hemianatropous), ascending, horizontal or pendulous, apotropous, usually bitegmic (sometimes unitegmic, derived from integumentary shifting, especially when ovule single), crassinucellar or pseudocrassinucellar (especially when unitegmic; in Anemone-Clematis clade). Micropyle usually endostomal (sometimes bistomal). Outer integument (two to) four to six (to ten) cell layers thick. Inner integument (one or) two or three cell layers thick. Parietal tissue one or two cell layers thick or absent. Nucellar cap present, two to six cell layers thick (mainly in taxa with a single ovule per carpel), or absent. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type; in Thalictrum etc. sometimes tetrasporous, Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells usually persistent, often large, proliferating (sometimes multinucleate). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually onagrad (sometimes solanad, rarely caryophyllad).

Fruit Usually an assemblage of follicles or achenes (sometimes a capsule with more or less fused walls, or single; in some species of Actaea a berry).

Seeds Aril absent. Seed coat exotestal. Exotesta palisade, with often thickened non-lignified cell walls, or seeds pachychalazal, with a thin testa. Mesotesta and endotesta unspecialized. Tegmen unspecialized. Perisperm not developed. Endosperm scarce to copious (rarely absent), usually starchy. Embryo very small to large, undifferentiated to well differentiated at dispersal, without chlorophyll. Cotyledons usually two (in some species of Anemone, Eranthis and Ranunculus one), often connate (cotyledonary tube often present; embryo without organs at seed dispersal, further growth intraseminal). Germination phanerocotylar or cryptocotylar. Radicula often ephemeral.

Cytology x = 8, 9 (Coptoideae), x = 7 (Thalictroideae), x = 6–9 (Ranunculoideae) – Polyploidy and aneuploidy frequent; one group of genera with small reniform T type (Thalictrum type) chromosomes and a second group of genera with long two-armed and often curved R type (Ranunculus type) chromosomes.

DNA Plastid gene infA lost/defunct (Caltha with pseudogene, Clematis, Ranunculus). Plastid inverted repeat in Anemone and Clematis expanded by 4,1 kb into large single copy region. Two inversions (39 and 24 kb, respectively) in plastid genome of Adonis Subg. Adonis; one inversion of 42 kb in plastid genome of Adonis Subg. Adonanthe. Four inversions in plastid genome of Anemone and Clematis. Two (or four [two parallel]) inversions in plastid genome of Clematis and at least three species of Anemone. Plastid gene rps16 lost in Adonis Subg. Adonis. One intron absent from plastid gene rps12 of Anemone canadensis, A. dichotoma and A. richardsonii. Mitochondrial intron coxII.i3 lost. Nuclear gene AP3 triplicated.

Phytochemistry Flavonols, (kaempferol, quercetin; frequently additional oxygenation at carbons 6 or 8 of ring A in contrast to magnoliid clades), cyanidin, caffeic acid, cardioactive bufadienolides (in Helleborus), Digitalis cardenolides (in Adonis), diterpenoid alkaloids (e.g. strongly poisonous aconitine and methyl lycaconitine, in Aconitum and Delphinium), damascenine (a protoalkaloid), benzylisoquinoline alkaloids (berberine, quaternary magnoflorine and its precursor corytuberine in Coptoideae and Thalictroideae), quinolizidine alkaloids (in Actaea etc.), pyrrolizidine alkaloids as macrocyclic diesters (in Caltha), tyrosine-derived cyanogenic compounds (abundant in Thalictroideae), ranunculines (in Ranunculeae, Anemoneae and Helleborus in Ranunculoideae), triterpene saponins (in Anemone, Ranunculus etc.), tetracyclic triterpene saponins with lanostane sapogenins (in Actaea, Beesia and some species of Thalictrum), and steroidal saponins (in Helleborus etc.), chelidonic acid present. Ellagic acid and tannins not found. Mannitol accumulation occurring in Aconitum and Delphinium. Oxalate accumulation sometimes occurring.

Use Ornamental plants, spices (seeds of Nigella sativa), medicinal plants, dyeing sources (Xanthorhiza, Coptis).

Systematics Ranunculaceae may be sister to the clade [Glaucidiaceae+Hydrastidaceae].

Coptoideae Tamura in Sci. Rep. Coll. N. Coll. Osaka Univ. 17: 52. 1 Feb 1968

2/c 16. Coptis (c 15; temperate regions on the Northern Hemisphere), Xanthorhiza (1; X. simplicissima; eastern North America). – Temperate regions on the Northern Hemisphere. Perennial herbs or suffrutices. Nectaries five to ten, petaloid, thick, stipitate. Carpels stipitate. n = (8) 9. Chromosomes small and rod-like, T type (Thalictrum type). Benzylisoquinoline alkaloids present in Coptis. – Coptoideae may be sister to [Thalictroideae+Ranunculoideae], although the support is low.

Thalictroideae Raf., Anal. Nat.: 176. Apr-Jul 1815 [‘Thalictrinia’]

9–10/c 450. Thalictrum (250–330; temperate regions on the Northern Hemisphere, tropical and southern Africa, New Guinea, tropical America), Leptopyrum (1; L. fumarioides; western Siberia to East Asia), Paraquilegia (5; western Iran to Himalayas and western China; incl. Paropyrum?), Paropyrum (1; P. anemonoides; Himalayas, western China; in Paraquilegia?), Urophysa (2; China), Aquilegia (c 80; temperate regions on the Northern Hemisphere), Semiaquilegia (6; East Asia), Dichocarpum (c 20; Himalayas, East Asia), Enemion (6; northeastern Asia, North America), Isopyrum (4; Central Europe, temperate Asia). – Temperate regions on the Northern Hemisphere, tropical and southern Africa, New Guinea, tropical America. Hairs capitate. Leaflets with curved-involute ptyxis, papillate. Stipule-like processes present in Thalictrum. Nectaries petaloid, stipitate. Intrastaminal staminodia sometimes present. Integument single, seven or eight cell layers thick. x = 7. Chromosomes small and reniform, T type (Thalictrum type). Tyrosine-derived cyanogenic compounds often present. Benzylisoquinoline alkaloids present in Isopyrum. – Thalictroideae may be sister to Ranunculoideae.

Ranunculoideae Arn., Botany: 94. 9 Mar 1832 [’Ranunculineae’]

36–37/1.700–1.800. Calathodes (3; Himalayas, China, Taiwan), Adonis (c 25; Europe, temperate Asia), Trollius (c 30; temperate regions on the Northern Hemisphere), Callianthemum (14; Central Europe and eastwards to Central Asia), Caltha (12; temperate regions on both hemispheres; section Psychrophila: the Andes to Ecuador, Tierra del Fuego, Falkland Islands, southeastern Australia, Tasmania, New Zealand), Helleborus (c 20; West, Central and South Europe, the Mediterranean and eastwards to Caucasus and northern Syria, one species, H. thibetanus, in Tibet and western China); Delphinium (c 320; temperate regions on the Northern Hemisphere), Staphisagria (3; the Mediterranean), Aconitum (>250; temperate regions on the Northern Hemisphere), Gymnaconitum (1; G. gymnandrum; Tibet, western China), Nigella (c 25; Europe, the Mediterranean, temperate Asia); Actaea (27; temperate regions on the Northern Hemisphere), Anemonopsis (1; A. macrophylla; central Honshu in Japan), Beesia (2; western and southwestern China, northern Burma), Asteropyrum (2; China), Eranthis (8–9; Europe, temperate Asia). – Ranunculeae DC., Syst. Nat. 1: 130, 228. 1-15 Nov 1817. Kumlienia (1; K. hystricula; Sierra Nevada in California), Callianthemoides (1; C. semiverticillatus; the Andes in southern Chile and southern Argentina), Hamadryas (6; southern Chile, Tierra del Fuego), Peltocalathos (1; P. baurii; southern Africa), Beckwithia (4; temperate and polar regions on the Northern Hemisphere), Cyrtorrhyncha (1; C. ranunculina; western North America), Oxygraphis (5; temperate Asia), Halerpestes (c 10; temperate Asia, North America), Arcteranthis (1; A. cooleyae; Alaska, northwestern Canada), Trautvetteria (1–2; T. caroliniensis; East Asia, western and southern North America), Coptidium (2; cold-temperate and arctic regions on the Northern Hemisphere), Ficaria (5; Europe, the Mediterranean and eastwards to Central Asia), Myosurus (6; temperate regions on both hemisphere), Ceratocephala (2–6; nearly cosmopolitan), Krapfia (8; the Andes), Laccopetalum (1; L. giganteum; the Andes in Peru), Ranunculus (c 600; polar, temperate and subtropical regions, tropical mountains), Paroxygraphis (1; P. sikkimensis; eastern Himalayas). – Anemoneae DC., Syst. Nat. 1: 129, 168. 1-15 Nov 1817. Anemone (c 150; temperate regions on the Northern Hemisphere, East African mountains, southerns Africa, Sumatra, Tasmania, New Zealand, North America and southwards to the Andes in Chile), Clematis (c 325; temperate regions on the Northern Hemisphere, tropical African mountains, Madagascar, Pacific islands, South America). – Cosmopolitan. Hairs clavate. Leaflets usually with involute (sometimes supervolute and/or curved) ptyxis. Petiole vascular bundle transection sometimes arcuate; wing bundles and medullary bundles sometimes present. Flowers sometimes zygomorphic. Ovule in Ranunculeaeand Anemoneaeone per carpel, unitegmic, pseudocrassinucellar, and integument six to twelve cell layers thick. Micropyle sometimes bistomal. Parietal tissue one or two cell layers thick or absent. Nucellar cap usually present. Endosperm development nuclear. Testa sometimes vascularized. Exotesta sometimes short-palisade. Endotesta sometimes developed. Embryo sometimes undifferentiated. x = (6–)8(–9). Chromosomes large, elongated, two-armed and often curved, R type (Ranunculustype). Lactone-forming glycosides (ranunculin, etc.). Cardenolides and bufadienolides sometimes present. Berberine absent. Benzylisoquinoline alkaloids sparse or absent.

Cladogram (simplified) of Ranunculaceae based on DNA sequence data (Ro & al. 1997; Wang & al. 2005; Wang & Chen 2007, Wang & al. 2010). The positions of Callianthemum, Caltha, Helleborus and Nigella, in particular, are very uncertain, and several other clades also have low support.

Cladogram of Ranunculeae based on DNA sequence data (Emadzade & al. 2010).


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