ROSALES Perleb Main Tree, Synapomorphies

Habit Usually bisexual (sometimes monoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), evergreen or deciduous trees, shrubs or suffrutices (sometimes lianas), or perennial (sometimes annual) herbs.

Vegetative anatomy Ectomycorrhiza sometimes present. Roots diarch (lateral roots tetrastichous) or tetrarch. Root nodules with nitrogen-fixing bacteria (Frankia) often present. Frankia infection sometimes taking place through intercellular penetration. Phellogen subepidermal, cortical or outer-cortical. Vessel elements usually with simple (sometimes reticulate or scalariform) perforation plates; lateral pits alternate or opposite, simple or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, usually non-septate (also vasicentric tracheids). Wood rays uniseriate to multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, scalariform, reticulate, confluent, unilateral, vasicentric or banded (sometimes absent). Fibres often present in secondary phloem. Sieve tube plastids Ss or S₀ type (rarely Pc type); sieve tubes at least sometimes with non-dispersive protein bodies. Nodes 1:1, unilacunar with one leaf trace, or 3:3, trilacunar with three traces (sometimes ≥5:≥5, penta- to multilacunar with five or more traces). Heartwood often with gum-like substances. Laticiferous cavities or ducts sometimes present. Calciumoxalate usually as solitary prismatic crystals (sometimes also druses or acicular crystals, rarely raphides) or absent. Prismatic crystals usually present in wood ray cells.

Trichomes Hairs unicellular or multicellular, simple or branched, furcate, stellate, lepidote, or peltate; glandular hairs unicellular or multicellular (sometimes peltate-lepidote or pearl glands); prickles or stinging hairs sometimes present.

Leaves Usually alternate (spiral or distichous; rarely opposite or verticillate), pinnately or palmately compound, or simple and entire or lobed, with conduplicate, supervolute, involute, plicate, curved or flat ptyxis. Stipules lateral, axillary, inserted on branch or adnate to petiole (rarely interpetiolar), small to foliaceous, usually persistent (sometimes caducous, rarely absent); leaf sheath absent. Petiole vascular bundle transection arcuate, U-shaped or annular. Venation usually pinnate (sometimes palmate, rarely single-veined), craspedodromous or camptodromous (sometimes acrodromous). Stomata usually anomocytic (sometimes cyclocytic, rarely anisocytic, helicocytic or paracytic). Cuticular wax crystalloids as tubuli or rosettes of platelets (Fabales type), or absent. Domatia as pits, pockets or hair tufts, or absent. Epidermis with or without mucilaginous idioblasts. Tanniniferous or mucilaginous cells often present. Cystoliths sometimes present. Laticiferous cavities or ducts sometimes present. Leaf margins and leaflet margins usually serrate or biserrate (sometimes crenate or entire, rarely glandular serrate); teeth often rosoid or urticoid.

Inflorescence Terminal or axillary, cymes, panicle, thyrse, raceme-, spike-, catkin-, head-, spadix- or umbel-like, corymb, raceme, umbellate, spicate or fasciculate (sometimes pseudanthia), or flowers solitary. Bracts and floral prophylls (bracteoles) sometimes absent.

Flowers Usually actinomorphic (occasionally zygomorphic). Hypanthium usually present (rarely absent). Hypogyny, epigyny or half epigyny. Sepals (one to) four or five (to ten), with valvate, induplicate-valvate or imbricate (rarely open) aestivation, persistent, free, or absent. Petals (three or) four or five (to ten), usually with imbricate (sometimes induplicate-valvate) aestivation, often clawed, caducous, free, or absent. Nectaries usually on adaxial side of hypanthium or on staminal bases, or absent. Nectariferous disc intrastaminal, annular or divided, inserted around hypanthial orifice, or absent.

Androecium Stamens one to more than 20. Filaments free or connate in lower part, free from tepals, usually inserted on hypanthium. Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse (sometimes latrorse or extrorse), longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores). Tapetum secretory. Staminodia usually absent (sometimes extrastaminal, petaloid; female flowers often with staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate or (1–)2–6(–15)-porate (sometimes stephanoporate), shed as monads, bicellular at dispersal, often starchy. Exine tectate or semitectate, with columellate or granular infratectum, perforate, reticulate, microreticulate, striate, rugulate, fossulate, scabrate, spinulate, echinulate, verrucate, pilate, or psilate.

Gynoecium Pistil composed of usually two to five more or less connate carpels, or carpels one (sometimes pseudomonomerous) or numerous (sometimes spiral), free or more or less connate or adnate to hypanthium (rarely somewhat stipitate, on gynophore). Ovary superior, inferior or semi-inferior, unilocular (often apocarpy) to quinquelocular (to octalocular). Style single, simple, or stylodia few to numerous, usually terminal (sometimes lateral or subbasal), free or more or less connate, or absent. Stigmas capitate, clavate, punctate, widened, decurrent or bilobate (rarely fimbriate or penicillate), papillate or non-papillate, Dry or Wet type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation axile, basal, subbasal, apical or subapical. Ovules usually one or two (sometimes several, rarely numerous) per carpel, usually anatropous (sometimes hemianatropous, campylotropous, or orthotropous), usually ascending or pendulous (rarely erect), apotropous or epitropous, bitegmic or unitegmic, crassinucellar. Micropyle bistomal or endostomal (sometimes exostomal, rarely almost absent). Obturator (sometimes funicular) sometimes present. Archespore unicellular or multicellular. Nucellar cap often present. Nucellar beak sometimes present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, 8-nucleate, Allium type, or tetrasporous, Drusa or Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes proliferating. Endosperm development usually nuclear (rarely helobial). Endosperm haustorium chalazal or absent. Embryogenesis usually asterad (sometimes onagrad, rarely solanad).

Fruit An assemblage of follicles, drupelets or achenes, a drupe, nut, samara, pome, loculicidal or septicidal(-septifragal) capsule (sometimes a cyconium or other types of fleshy syncarp, or a cynarrhodium, rarely schizocarp or anthocarp), often with persistent hypanthium and/or calyx.

Seeds Funicular aril sometimes present. Elaiosome rarely present. Seed coat usually mesotestal (sometimes exotestal, occasionally reduced). Testa sometimes multiplicative. Exotesta often palisade (sometimes tabular); exotestal cells often malpighiaceous with thickened walls (often spiral or with reticulate thickenings), sometimes periclinally elongate. Mesotesta usually sclerotic, sometimes aerenchymatous. Endotesta sometimes tanniniferous. Exotegmen crushed. Endotegmic cells somewhat thickened, sometimes tanniniferous, sometimes cuboid. Perisperm usually not developed. Endosperm copious, sparse or absent, oily. Embryo straight or curved (rarely spirally twisted), with or without chlorophyll. Cotyledons usually two (rarely one). Germination phanerocotylar or cryptocotylar.

DNA Plastid gene infA lost/defunct. Mitochondrial intron coxII.i3 lost. Cis-spliced (Rosaceae) or trans-spliced intron in mitochondrial gene nad1 (Rosales except Rosaceae).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin) and their glycosides, isoflavonoids, afzelechin, flavone-C-glycosides, flavanone glycosides, biflavonoids (biflavonyls), cyanidin, epigallocatechin-3-gallate, dihydrochalcones, dammaranes, cucurbitacins, oleanolic acid derivatives, methylated and non-methylated ellagic acids, gallic acid, hydrolyzable ellagitannins (casuaricitin, geraniins, pedunculagin, stachyurins, tellimagrandin I, tellimagrandin II), non-hydrolyzable tannins, proanthocyanidins (prodelphinidins), chlorogenic acid, caffeic acid derivatives, hydroxycinnamate derivatives, indole alkaloids, isoquinoline alkaloids (e.g. liriodenine and benzylisoquinoline alkaloids), peptide alkaloids, phenylalanine-, tyrosine- or leucine-derived cyanogenic glucosides (amygdalin, cardiospermin, dhurrin, heterodendrin, prunasin), triterpene saponins, anthraquinones, naphthoquinones, arbutin, benzophenones, xanthones, polyacetate-derived arthroquinones, sesquiterpene lactones, coumarins, furanocoumarins, p-coumaric acid, acetophenones, L-quebrachitol, cannabinoids (tetrahydrocannabinol etc.), simple indole bases, humulones, lupulones, chelidonic acid, sinapic acid, ursolic acid, eleostearic acid, barbeyol, naphthalene and lupeolin derivatives, and lignans present.

Rosales except Rosaceae and Cynomoriaceae are characterized by trans-spliced intron in mitochondrial nad1 gene.

The clade [Rhamnaceae+[Elaeagnaceae+[Barbeyaceae+Dirachmaceae]]] has the following potential synapomorphies (Stevens 2001 onwards): petiole bundle transection arcuate; stamens as many as petals, alternitepalous/antepetalous; placentation basal; ovule erect; parietal tissue five or six cell layers thick; capsule septicidal; seed coat multiplicative; exotesta palisade, with thick-walled cells; and cotyledons large. Barbeyaceae and Dirachmaceae share the features: anther connective prolonged; and outer integument three to five cell layers thick. Rhamnaceae and Dirachma differ from the Ulmaceae-Urticaceae clade by pits of wood ray parenchyma resembling lateral pits of vessel elements.

The clade [Ulmaceae+[Cannabaceae+[Moraceae+Urticaceae]]] is characterized by the following potential synapomorphies (Stevens 2001 onwards): presence of watery exudate; hairs unicellular and multicellular-glandular; cambium storied; presence of libriform fibres; phloem stratified; sieve tubes with non-dispersive protein bodies; presence of globose and usually calciumcarbonate cystoliths; epidermal and hair cell walls silicified and calcified; presence of at least one prominent prophyllar bud; stipules cauline; presence of urticoid teeth (secondary leaf veins proceeding straight to non-glandular teeth, higher-order veins convergent on those teeth); flowers small; absence of corolla and nectary; stamens as many as tepals, antetepalous; pollen grains porate; infratectum granular; carpels two; only abaxial carpel fertile; stigmas sessile, diverging, with pollen receptive area extending down adaxial surface and confluent; placentation apical; ovule epitropous; fruit a drupe; testa perforated; endosperm sparse; polyembryony frequent; x = 14; centromeres median and subterminal; presence of flavonols and their glycosides, including myricetin; and absence of ellagic acid.

The clade [Cannabaceae+[Moraceae+Urticaceae]] has the potential synapomorphies (Stevens 2001 onwards): flowers unisexual; sieve tube plastids sometimes Ss type (with starch grains); unicellular hairs usually micropapillate; stipules cauline-intrapetiolar; venation palmate; embryo curved; and presence of flavone-C-glycosides. Finally, Moraceae and Urticaceae share the following potential synapomorphies: presence of laticifers and latex (usually white, milky); venation sometimes pinnate; stamens inflexed in bud; presence of pistillodium; and absence of polyembryony.

Rendle in D. Oliver (ed. D. Prain), Fl. Trop. Afr. 6(2): 14. Mar 1916, nom. cons.

Barbeyales Takht. et Reveal in Phytologia 74: 172. 25 Mar 1993; Barbeyanae Takht. ex Reveal et Doweld in Novon 9: 549. 30 Dec 1999

Habit Dioecious, evergreen trees with silvery indumentum on abaxial side of lamina.

Vegetative anatomy Phellogen? Primary medullary strands narrow, alternating with wide strands. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays usually uniseriate or biseriate, heterocellular. Axial parenchyma paratracheal scanty, or absent. Tyloses abundant in vessels. Secondary phloem with numerous peripheral fibres and sclereids. Sieve tube plastids S type, with up to ten starch grains; sieve tubes with compound perforations. Nodes 1:1, unilacunar with one leaf trace. Mucilage cells? Laticifers and latex absent. Cystoliths absent. Wood ray cells without prismatic crystals.

Leaves Opposite, simple, entire, with supervolute-curved ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate, camptodromous. Stomata anomocytic or laterocytic. Cuticular wax crystalloids? Mucilaginous idioblasts absent. Leaf margin entire.

Inflorescence Axillary, fasciculate or umbel-like, cymose. Bracts and floral prophylls (bracteoles) absent.

Flowers Actinomorphic. Hypanthium absent. Hypogyny. Sepals three or four, in male flowers with valvate aestivation, in female flowers with imbricate aestivation, persistent, in female flowers free, in male flowers somewhat connate at base. Petals absent. Nectary absent. Disc absent.

Androecium Stamens six to nine (to twelve). Filaments short, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits); connective apically prolonged. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpor(oid)ate, shed as monads, ?-cellular at dispersal. Exine tectate to semitectate, with granular to intermediate infratectum, perforate or finely reticulate, rugulate, spinulate or verrucate.

Gynoecium Carpels one or two (or three), free (apocarpy) or connate at base. Ovary superior, usually unilocular (monomerous; sometimes bilocular or trilocular), sometimes slightly stipitate. Stylodia one or two (or three), free or connate at base. Stigmas clavate, decurrent, papillate, type? Pistillodium absent.

Ovules Placentation subapical. Ovule one per carpel, anatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument five or six cell layers thick. Hypostase present. Nucellar cap approx. five cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Seeds Aril absent? Testa indistinct, unspecialized, non-multiplicative. Exotesta perforated, aerenchymatous, not palisade, with sinuous anticlinal cell walls. Endotesta?, tanniniferous. Exotegmen? Endotegmic cells tanniniferous, with sinuous anticlinal walls. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, fleshy. Germination?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin), ellagic acid, and barbeyol (phenolic indane) present. Alkaloids possibly present? Myricetin and glycoflavonols not found.

Humulaceae Bercht. et J. Presl, Přir. Rostlin: 258. Jan-Apr 1820; Lupulaceae Schultz Sch., Nat. Syst. Pflanzenr.: 370. 30 Jan-10 Feb 1832 [’Lupulinae’]; Celtidaceae Endl., Ench. Bot.: 171. 15-21 Aug 1841 [’Celtideae’]

Distribution Tropical and subtropical regions, temperate regions on the Northern Hemisphere.

Fossils Endocarps assigned to Aphananthe cretacea and Gironniera gonnensis have been found in the Maastrichtian of Germany. Eoceltis dilcheri, a fossil male flower of proposed affinity to Celtidoideae, has been described from the mid-Eocene of Texas. The pollen grains are triporate and scabrate. Celtoidanthus pseudorobustus is another male flower of questioned affinity with numerous stamens and triporate pollen grains from the Miocene of Germany. Cenozoic fossil Cannabaceae comprise leaves, pollen grains and endocarps from the Northern Hemisphere.

Habit Usually monoecious, polygamomonoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees or shrubs (some species of Celtis are lianas), perennial or annual herbs (Humulus is twining and climbing by means of specialized hairs).

Vegetative anatomy Ectomycorrhiza sometimes (Gironniera) present. Root nodules with nitrogen-fixing bacteria (Parasponia only known genus outside Fabaceae with other types of nitrogen-fixing bacteria, Rhizobium, than actinobacteria; rhizobia persisting inside infection threads). Phellogen ab initio superficial. Medulla in Celtis septated by diaphragms. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, usually simple pits. Imperforate tracheary elements tracheids or libriform fibres with usually simple (sometimes bordered) pits, septate or non-septate (in, e.g., Ampelocera septate). Wood rays uniseriate to multiseriate, homocellular or heterocellular. Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, unilateral, or banded (sometimes apotracheal diffuse), or absent. Tyloses often frequent (sometimes sclerotic). Secondary phloem stratified into hard fibrous and soft parenchymatous non-fibrous layers. Sieve tube plastids Ss type or S₀ type; sieve tube elements with extruded nucleoli and non-dispersive protein bodies. Nodes 3:3, trilacunar with three leaf traces (in Humulus split laterals or commissural bundle). Secretory ducts usually without latex (coloured latex rare?). Mucilage cells absent. Cystoliths (of calciumcarbonate?) globose, usually with pegs (cellulose expansions of cell walls, covered with crystals). Epidermal cell walls often with calciumcarbonate or silica. Prismatic calciumoxalate crystals abundant; druses or crystal sand present in some species.

Trichomes Hairs rigid or soft, unicellular (usually micropapillate) or multicellular, usually simple, often with inflated base (sometimes furcate); unicellular or multicellular glandular hairs (also peltate-lepidote) often abundant.

Leaves Usually alternate (usually distichous, sometimes spiral; in Humulus and Lozanella opposite), usually simple (in, e.g., Cannabis palmately compound), usually entire (rarely palmately lobed), with conduplicate-plicate (sometimes laterally; sometimes conduplicate or supervolute) ptyxis. Stipules usually lateral and axillary, intrapetiolar (in Lozanella interpetiolar; in Parasponia intrapetiolar), sometimes more or less connate, persistent or caducous; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate or palmate, usually brochidodromous or eucamptodromous (in Aphananthe and Parasponia craspedodromous; in Trema acrodromous), usually with three main-veins from base; secondary veins usually not reaching into teeth (secondary and tertiary veins in Cannabis and Humulus proceeding into lobe tips and teeth: urticoid teeth). Stomata anomocytic. Cuticular wax crystalloids? Domatia? Epidermis and mesophyll often with mucilaginous idioblasts. Leaf margin serrate or entire. Leaves often with unicellular hairs with inflated base, often with glands with bitter-tasting substances (lupulin etc. in Humulus) or glandular hairs with aromatic resins (tetrahydrocannabinols in Cannabis).

Inflorescence Axillary, cymose (often thyrsoid, sometimes spicate; female inflorescence in Humulus cone-like; flowers solitary in Chaetachme). Prophylls usually basal (in Humulus absent).

Flowers Actinomorphic, small to minute. Hypogyny. Sepals (four or) five (to seven), usually with imbricate (in Trema induplicate-valvate) aestivation, whorled, sometimes persistent, usually connate at base (rarely free). Petals absent. Nectary absent. Disc absent.

Androecium Stamens usually (four or) five (to seven; in Ampelocera four to 16), antesepalous. Filaments usually filiform, sometimes inflexed in bud, free, adnate to sepal bases. Anthers dorsifixed, versatile, tetrasporangiate, usually introrse (sometimes extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to quadrinucleate cells. Female flowers sometimes (especially in Celtis) with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3–5(–6)-porate, shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, perforate scabrate, spinulate, verrucate or echinulate.

Gynoecium Pistil composed of two partially connate carpels. Ovary superior, unilocular. Stylodia two, sometimes connate at base. Stigmas decurrent, ventral (in Lozanella bilobate), papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation subapical. Ovules one per ovary, anatropous to orthotropous or campylotropous, pendulous, epitropous, bitegmic, crassinucellar (to tenuinucellar?). Micropyle usually endostomal? (in Celtis bistomal). Outer integument two to four (to eight) cell layers thick. Inner integument two or three cell layers thick. Hypostase present. Parietal tissue approx. six cell layers thick. Nucellar cap approx. two cell layers thick. Megagametophyte monosporous, Polygonum type, with haustorium. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis onagrad (at least in Cannabis and Humulus).

Fruit Usually a drupe (in Pteroceltis a samara; in Cannabis and Humulus a nutlet enclosed by calyx).

Seeds Aril absent. Exotestal cells periclinally elongate, with arm-shaped processes, with unthickened walls (anticlinal walls in Humulus sinuous). Endotesta? Tegmen? Perisperm not developed. Endosperm usually sparse or absent (in Lozanella and Parasponia carnose), oily. Embryo curved (in Humulus spirally twisted), usually without chlorophyll (in Humulus with chlorophyll). Cotyledons two, folded, inrolled. Germination phanerocotylar.

Cytology n = 8–11, 13–15 (20) (30) (42) (c. 60) – Polyploidy occurring. Centromeres medial/submedial, simple. Cannabis and Humulus with X-autosome balance system (sex chromosomes, X or Y).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin; in Aphananthe, Ampelocera, Gironniera, Cannabis, and Humulus) and their glycosides, flavone-C-glycosides, prodelphinidins, sesquiterpene lactones, alkaloids, triterpene saponins, tyrosine-derived cyanogenic compounds, quebrachitol, cannabinoids (in Cannabis), humulones (bitter-tasting α-lupulic acids, in Humulus), and lupulones (bitter-tasting β-lupulic acids, in Humulus) present. Ellagic acid, sesquiterpenes and lignans not found. Raffinose and stachyose present in phloem exudates.

Use Ornamental plants, fibre plants (ropes and paper from phloem of male plants of Cannabis sativa), beer spices (Humulus lupulus), seed oils, medicinal plants and narcotics (Cannabis sativa), timber, carpentries.

Habit Dioecious, evergreen shrubs or small trees. Long and short shoots well differentiated.

Vegetative anatomy Phellogen ab initio subepidermal. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Vestured pits absent. Imperforate tracheary xylem elements tracheids with more or less simple pits, non-septate (also vasicentric tracheids). Wood rays usually uniseriate or biseriate, heterocellular. Axial parenchyma apotracheal banded, or paratracheal scanty vasicentric. Tyloses or gum-like substances frequent in heartwood. Secondary phloem stratified into hard fibrous and soft parenchymatous non-fibrous layers. Sieve tube plastids S type? Nodes? Prismatic calciumoxalate crystals abundant; druses present in cortical and medullary parenchyma cells.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules intrapetiolar, linear-triangular (subulate), persistent; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata anomocytic or cyclocytic. Cuticular wax crystalloids? Epidermis with mucilaginous idioblasts. Leaf margin coarsely serrate to crenate.

Flowers Actinomorphic, large. Hypanthium-like structure present between sepals and petals. Pedicel with ’epicalyx’ (’hypocalyx’) consisting of four to eight bracts. Half epigyny. Sepals five or six (Dirachma somalensis) or eight (Dirachma socotrana), with valvate aestivation, caducous above hypanthium-like structure (consisting of perianth and sometimes also androecium), free. Petals five or six (Dirachma somalensis) or eight (Dirachma socotrana), with contorted aestivation, with ventral appendages at base, free. Nectariferous glands present at petal bases (petal nectaries) or on subbasal (ventral) appendages, caducous. Disc absent.

Androecium Stamens five or six (Dirachma somalensis) or eight (Dirachma socotrana), obhaplostemonous, alternisepalous, antepetalous. Filaments free, adnate at base to petals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing from apex by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

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

Gynoecium Pistil composed of five or six (Dirachma somalensis) or eight (Dirachma socotrana) connate antesepalous carpels. Ovary semi-inferior, quinquelocular or sexalocular (Dirachma somalensis) or octalocular (Dirachma socotrana), and lobate. Style single, simple. Stigmas five or six (Dirachma somalensis) or eight (Dirachma socotrana), clavate to cylindrical or punctate, type? Pistillodium absent.

Ovules Placentation basal to axile. Ovule one per carpel, anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument six to ten cell layers thick. Inner integument two or three cell layers thick. Hypostase present? Nucellar cap minute or absent. Megagametophyte monosporous, probably Polygonum type. Endosperm development ab initio nuclear, later cellular. Endosperm haustorium chalazal. Embryogenesis?

Fruit A septicidal and septifragal capsule with columella, dehiscing adaxially from base to apex (a schizocarp with five or six [Dirachma somalensis] or eight [Dirachma socotrana] ventricidal follicular mericarps).

Seeds Seed flattened, with small funicular aril. Seed coat exotestal-endotegmic. Testa multiplicative, with median antiraphal vascular bundle. Exotesta palisade with thick anticlinal cell walls, tanniniferous. Endotesta and exotegmen crushed. Endotegmic cells thick-walled, tanniniferous. Perisperm not developed. Endosperm scarse or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, large. Germination?

Elaeagnales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 234. Jan-Apr 1820 [‘Elaeagneae’]; Hippophaëaceae G. Mey., Chloris Han.: 456, 460. Jul-Aug 1836 [’Hippophaëae’]

Distribution Temperate and subtropical regions on the Northern Hemisphere, Southeast Asia, Queensland.

Fossils Pollen grains similar to Elaeagnaceae have been reported from Cenozoic and even Late Cretaceous layers in Asia and Europe (e.g. Slowakipollis from the Oligocene of eastern Europe). Pollen of Shepherdia was recorded from the Miocene of Oregon and Idaho. Fossil leaves of Elaeagnus were reported from Miocene layers on the Qinghai-Tibet Plateau.

Habit In Elaeagnus monoecious, androdioecious or gynodioecious, in Hippophae and Shepherdia dioecious, usually deciduous (sometimes evergreen) trees or shrubs (rarely climbing), often spiny and with silvery leaves. Some species are xerophytes.

Vegetative anatomy Ectomycorrhiza often present. Root nodules with nitrogen-fixing bacteria (Frankia etc.) usually present. Cluster roots present in Hippophae. Phellogen ab initio superficial. Cambium storied. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids or fibre tracheids (libriform fibres?) with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or absent (Shepherdia). Wood elements often more or less storied. Tyloses frequent. Phloem usually tangentially stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids S₀ type; sieve tube nuclei with non-dispersive protein bodies. Nodes 1:1, unilacunar with one leaf trace. Heartwood often with gum-like substances. Calciumoxalate as crystal sand often present. Wood ray cells without prismatic crystals.

Leaves Usually alternate (spiral; rarely opposite), simple, entire, often coriaceous, with conduplicate-flat ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Mucilaginous idioblasts absent. Leaf margin entire.

Inflorescence Axillary, usually raceme, spike or fascicle (flowers rarely solitary).

Flowers Actinomorphic, usually small. Receptacle flat (often in male flowers) or tubular hypanthium (in female and bisexual flowers). Hypogyny. Sepals (two to) four (to six), usually with valvate (rarely open) aestivation, sometimes petaloid, connate in lower part into tubular hypanthium. Petals absent. Receptacular nectaries present. Disc consisting of separate parts.

Androecium Stamens (two to) four (to six) (Elaeagnus, Hippophae) or 4+4 (rarely 6+6) (Shepherdia), haplostemonous or obdiplostemonous, antesepalous and/or alternisepalous. Filaments free, adnate to upper part of hypanthium. Anthers basifixed or dorsifixed, usually non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate, shed as monads, bicellular or tricellular at dispersal. Exine tectate, with columellate infratectum, perforate, punctate or somewhat rugulate.

Gynoecium Pistil composed of one carpel (pseudomonomery?). Ovary superior, unilocular. Style (stylulus) single, simple, long. Stigma decurrent or capitate, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation basal. Ovule one per ovary, anatropous, ascending, bitegmic, crassinucellar. Funicle short. Micropyle endostomal? Outer integument five to 16 cell layers thick. Inner integument three or four cell layers thick. Obturator funicular. Archespore multicellular. Megagametophyte usually monosporous, Polygonum type (in Shepherdia disporous, 8-nucleate, Allium type). Endosperm development ab initio nuclear. Endosperm haustorium inElaeagnus chalazal. Embryogenesis?

Fruit An achene surrounded by accrescent and fleshy hypanthium (baccate or drupaceous anthocarp). Pericarp thin.

Seeds Aril absent. Testa very thick, hard. Exotesta often palisade, with malpighiaceous cells with thick, at least partially sinuous anticlinal cell walls. Mesotesta aerenchymatous, with thick-walled cells. Endotesta? Tegmen? Perisperm not developed. Endosperm sparse, sometimes starchy, or absent. Embryo straight, well differentiated, oily and with aleuron (starch absent), without chlorophyll. Cotyledons two, fleshy, plano-convex. Germination phanerocotylar.

Cytology n = 6, 10, 11, 13, 14 (x = 12 in Hippophae, x =14 in Elaeagnus, x = 11, 13 in Shepherdia)

Phytochemistry Flavonols (kaempferol, quercetin) and other O-methylflavonoids, pentacyclic triterpenes, ellagic and gallic acids, hydrolyzable ellagitannins, condensed tannins, simple indole bases, indole alkaloids, saponins, p-coumaric acid, L-quebrachitol, and sinapic acid present. Dihydroflavonols? Myricetin, cyanidin, proanthocyanidins (prodelphinidins), caffeic acid, and cyanogenic compounds not found.

Artocarpaceae Bercht. et J. Presl, Přir. Rostlin: 260. Jan-Apr 1820 [‘Artocarpeae’]; Ficaceae Bercht. et J. Presl, Přir. Rostlin: 260. Jan-Apr 1820 [‘Ficeae’]; Dorsteniaceae Chevall., Fl. Gén. Env. Paris 2: 376. 1827; Ficales Dumort., Anal. Fam. Plant.: 15. 1829 [‘Ficarieae’]; Artocarpales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 14. Sep-Oct 1835 [’Artocarpeae’]; Morales Endl. in C. F. P. von Martius, Consp. Regn. Veg.: 13. Sep-Oct 1835 [‘Moreae’]

Distribution Tropical and subtropical regions, few species in temperate regions.

Fossils Apart from numerous Cenozoic fossils of Moraceae, there are leaves and reproductive organs of presumed Artocarpus from the Late Cretaceous of Greenland.

Habit Monoecious or dioecious (rarely gynodioecious), usually evergreen (sometimes deciduous) trees, shrubs, lianas, suffrutices or perennial herbs (Dorstenia and other tuberous geophytes or succulents; in Fatoua annual herbs). Often with large plank buttresses. Certain species of Ficus are epiphytes, some of which, the ’stranglers’, anchor in the soil by means of adventitious roots and then ‘strangle’ the host tree by the growth of these roots causing the death of the host.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes outer-cortical). Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes opposite), usually simple (sometimes bordered) pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres (often very long) usually with simple (sometimes bordered) pits, septate (in Castilleae) or non-septate (also vasicentric tracheids). Wood rays usually multiseriate (sometimes uniseriate), usually heterocellular (sometimes homocellular). Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, scalariform, reticulate, unilateral, or banded (rarely apotracheal banded). Wood elements storied. Tyloses frequent (sometimes sclerotic). Secondary phloem often stratified into hard fibrous and soft parenchymatous non-fibrous layers. Sieve tube plastids Ss type or S₀ type; sieve tube elements with extruded nucleoli and non-dispersive protein bodies. Nodes usually 3:3, trilacunar with three leaf traces (in Ficus 5:5, pentalacunar with five traces). Laticifers usually with abundant white (sometimes colourless) latex (absent in Fatoua; in Fatoua and Malaisia coloured juice); growth and branching of laticiferous cells not accompanied by cell wall formation. Secretory mucilaginous cavities usually present. Heartwood sometimes with gum-like substances. Cystoliths abundant, globose (calciumcarbonate?), usually with pegs? (cellulose expansions of cell wall, covered with crystals). Epidermal cell walls sometimes with calciumcarbonate or silica. Prismatic calciumoxalate crystals abundant; druses present in some species.

Trichomes Hairs unicellular (usually micropapillate) or multicellular, simple, often capitate (with unicellular stalk and uni- or multicellular head), sometimes uncinate, sometimes calcified or silicified; pearl glands often frequent.

Leaves Usually alternate (spiral or distichous; rarely opposite or verticillate), usually simple (rarely pinnately or palmately compound), entire or pinnately or palmately lobed, often coriaceous, with various ptyxis. Stipules often large, intrapetiolar or interpetiolar, sometimes sheathing (in Ficus open in leaf axils) or reduced, persistent or deciduous (sometimes connate; stipules absent in Fatoua); leaf sheath absent. Petiole vascular bundle transection? Venation pinnate or palmate; secondary and tertiary veins proceeding into non-glandular teeth (urticoid teeth)? Extrafloral nectaries present in Ficus on abaxial or adaxial side of lamina. Stomata usually anomocytic (rarely anisocytic or cyclocytic). Cuticular wax crystalloids? Domatia usually absent (sometimes as pockets or hair tufts). Mesophyll often with sclerenchymatic idioblasts with tannins, gum, mucilage or crystals of calciumcarbonate, calciumoxalate or silica. Hydathodes sometimes present. Leaf margin serrate, crenate or entire (rarely glandular-serrate).

Inflorescence Terminal or axillary, simple cymose (often raceme-, spike-, umbel- or headlike) or racemose, or compound open or almost closed, often spicate male inflorescence and globose female inflorescence, often pseudanthium (flowers rarely solitary). Receptacle often accrescent and fleshy in fruit, in some genera urceolate and dome-shaped with flowers on adaxial side (Ficus), or open and discoid (Dorstenia etc.). Inflorescence in Ficus concave, almost closed syconium, with apical pore, ostiole. Prophylls usually basal.

Flowers Actinomorphic, small to minute. Usually hypogyny (sometimes half epigyny). Sepals (one to) four or five (to ten), with valvate or imbricate aestivation, persistent or caducous, free or (often in female flowers) connate at base (sometimes adnate to sepals of adjacent flowers), or absent. Petals absent. Nectary absent (possibly present in some genera). Disc absent.

Androecium Stamens (one to) four (to eight), antesepalous, usually as many as sepals. Filaments usually inflexed in bud (in Moreae recurved with lightning rapidity at anthesis; rarely straight), usually free (sometimes connate), usually free from (sometimes adnate to?) sepals. Anthers basifixed or dorsifixed, often versatile, usually tetrasporangiate (rarely disporangiate), introrse or extrorse, longicidal (dehiscing by longitudinal slits, sometimes explosively dehiscent). Tapetum secretory. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (1–)2–3(–5)-porate (rarely ≥6-pantoporate), shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, smooth or scabrate.

Gynoecium Pistil composed of two (or three) connate carpels, one or two of which usually degenerating (in Broussonetia modified into pseudostyle). Ovary usually superior (sometimes semi-inferior), unilocular (or bilocular). Style single, simple, or stylodia two, free or connate at base. Stigmas decurrent, papillate or non-papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation usually apical or subapical (sometimes almost lateral, rarely basal). Ovule one per ovary, anatropous or campylotropous, usually pendulous (rarely erect and basal), epitropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument three or four cell layers thick. Inner integument approx. three cell layers thick. Hypostase present or absent. Nucellar cap approx. five cell layers thick. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, 8-nucleate, Allium type). Antipodal cells usually not proliferating (in at least Dorstenia proliferating). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually asterad.

Fruit Usually a drupe with dehiscing exocarp (rarely an indehiscent drupe or achene) or enclosed by fleshy perianth and/or sunken into accrescent fleshy floral receptacle and/or fleshy inflorescence axis; single fruits often connate into fleshy syncarp (in, e.g., Ficus, Morus and Artocarpus; sepals, bracts and receptacle in Artocarpus and Parartocarpus not distinctly separate; syncarp in Artocarpus as ripe weighing up to c. 50 kg; syncarp in Ficus a syconium with nutlets enclosed within fleshy inflorescence axis).

Seeds Aril absent. Testa indistinct, poorly differentiated, usually vascularized (in Prainea with several thickened cell layers). Exotesta tanniniferous. Tegmen? Perisperm not developed. Endosperm usually sparse, often oily, or absent. Embryo usually large, straight to curved, usually without chlorophyll. Cotyledons single (one degenerating) or two, flattened or folded. Germination phanerocotylar or cryptocotylar.

Cytology n = 12–16, 18, 20, 21, 24, 28, 32 (large variation in Dorstenia) – Polyploidy occurring; centromeres terminal and median.

Phytochemistry Flavonols (kaempferin, quercetin) and their glycosides, flavone-C-glycosides and other flavonoids, isoflavonoids, cyanidin, alkaloids, triterpene saponins, cyanogenic compounds, hydroxycinnamate derivatives, benzophenones, xanthones, coumarins, and furanocoumarins (in Ficus) present. Ellagic acid and tannins not found. Latex containing gums, proteins (e.g. papain-like enzyme), waxes, triterpenoid resins, polyphenols, cardenolids, etc. Raffinose and stachyose present in phloem exudates.

Use Ornamental plants, fruits (Ficus carica, Artocarpus altilis, Artocarpus heterophyllus, Morus nigra), textile (tapa) and paper (Antiaris, Broussonetia papyrifera), nutrient substrate of silk moth larvae (Bombyx mori on Morus alba), narcotics (Brosimum acutifolium), medicinal plants, arrow poison, rubber (Castilla elastica, Ficus elastica), timber.

Systematics Moraceae are sister-group to Urticaceae.

The clade [Artocarpeae+Moreae] is sister to the remaining Moraceae.

[Artocarpeae+Moreae]

Female flowers with peltate bracts.

Artocarpeae Lam. et DC., Syn. Plant. Fl. Gall.: 183. 30 Jun 1806

3/65–70. Batocarpus (3; B. amazonicus, B. costaricensis, B. orinocensis; Costa Rica to Bolivia; paraphyletic?), Clarisia (3; C. biflora, C. ilicifolia, C. racemosa; southern Mexico to Bolivia; in Batocarpus?), Artocarpus (c 60; tropical Asia east to islands in western Pacific). – Tropical Asia, tropical America. Male flowers dimerous. Stamen usually single (sometimes three). Filaments straight.

Moreae Dumort., Anal. Fam. Plant.: 17. 1829

6/50–55. Sorocea (14; southern Mexico, Central America, tropical South America), Bagassa (1; B. guianensis; northeastern South America), Trophis (9; Madagascar, Malesia to New Caledonia, southern Mexico, Central America, the West Indies, tropical South America), Morus (10–15; tropical and southern Africa, Southwest, East and Southeast Asia, eastern United States, Texas, Mexico, Central America, tropical America), Milicia (2; M. excelsa, M. regia; tropical Africa), Streblus (14; tropical Africa, Madagascar, Southeast Asia, eastern Queensland, eastern New South Wales, the Solomon Islands, Norfolk Island, New Zealand, Micronesia, Polynesia). – Pantropical, few species in temperate regions. – Sorocea may be sister to the remaining Moreae.

[Maclureae+[[Parartocarpus+Hullettia]+[Dorstenieae+[Castilleae+Ficeae]]]]

Maclureae W. L. Clement et Weiblen in Syst. Bot. 34: 545. 2009

1/11–12. Maclura (11–12; tropical and subtropical regions on both hemispheres). – Dioecy. Axillary spines often present. Inflorescence often with glands with yellow dye. Bracts of male flowers not peltate. Bracts of female flowers sometimes peltate. Stigmas usually two, often unequal in length.

Parartocarpus (4; P. bracteatus, P. triandrus, P. venenosus, P. woodii; Malesia) and Hullettia (2; H. dumosa, H. griffithiana; southern Burma, Peninsular Thailand, the Malay Peninsula, Sumatra) form a clade sister to [Dorstenieae+[Castilleae+Ficeae]] in the analyses by Gardner & al. (2017).

[Dorstenieae+[Castilleae+Ficeae]]

Monoecy (inflorescences bisexual). Radial laticiferous ducts present. Pistillodium conical.

Dorstenieae Dumort., Anal. Fam. Plant.: 16. 1829

12/145–150. Fatoua (2; F. pilosa, F. villosa; Madagascar, East and Southeast Asia, northern and easternmost Australia, New Caledonia), Broussonetia (8; Madagascar, tropical and subtropical Asia), Malaisia (1; M. aculeata; Malesia), Bleekrodea (4; B. insignis, B. madagascariensis, B. malayana, B. tonkinensis; Madagascar, the Malay Peninsula, Borneo), Sloetia (1; S. elongata; West Malesia, the Philippines, Sulawesi), Dorstenia (c 105; tropical regions on both hemispheres), Utsetela (2; U. gabonensis, U. neglecta; Gabon, Congo), Brosimum (15–18; southern Mexico, Central America, tropical South America), Trilepisium (1; T. madagascariense; tropical and southern Africa, Madagascar, the Mascarene Islands), Treculia (3; T. africana, T. lamiana, T. obovoidea; tropical West and Central Africa, Madagascar), Bosqueiopsis (1; B. gilletii; tropical Africa)?, Scyphosyce (2; S. manniana, S. pandurata; tropical West and Central Africa)? – Tropical and subtropical regions on both hemispheres, with their highest diversity in the Old World. Male flowers sometimes trimerous, often with connate sepals. Female flowers sometimes dimerous, sometimes sunken into receptacle. – Fatoua is sister to the remaining Dorstenieae.

[Castilleae+Ficeae]

Inflorescence axis urceolate, with insects (thrips in Castilleae; fig wasps, Agaonidae in Ficeae) reproducing among flowers and simultaneously pollinating these. Filaments straight. Bracts of female flowers not peltate. Sepals free.

Castilleae C. C. Berg in Acta Bot. Neerl. 26: 78. 15 Feb 1977

11/65–70. Sparattosyce (1; S. dioica; New Caledonia), Antiaropsis (2; A. decipiens, A. uniflora; New Guinea), Poulsenia (1; P. armata; southern Mexico to Bolivia), Antiaris (1; A. toxicaria; tropical regions in the Old World), Mesogyne (1; M. insignis; tropical Africa), Naucleopsis (16–20; tropical America; non-monophyletic?), ’Perebea’ (9–10; tropical America; polyphyletic), Maquira (4; M. calophylla, M. coriacea, M. guianensis, M. sclerophylla; tropical South America), Helicostylis (17; tropical America), Castilla (3; C. elastica, C. tunu, C. ulei; Central America, tropical South America), Pseudolmedia (10–12; southern Mexico, Central America, the West Indies, tropical South America). – Tropical and subtropical regions on both hemispheres, with their largest diversity in tropical America. Wood fibres septate. Cystoliths absent. Inflorescence axis involucrate, discoid to urceolate. Bracts absent in male flowers. Bracts usually present in female flowers. Pistillodium absent. – The clade [Antiaropsis+Sparattosyce] (Antiaropsineae) is sister to the remaining Castilleae.

Ficeae Dumort., Fl. Belg.: 24. 1827 [‘Ficineae’]

1/800–850. Ficus (800–850; tropical and subtropical regions on both hemispheres, few species in warm-temperate regions). – Latex usually present. Leaves sometimes opposite. Inflorescence syconium. Flowers trimerous. Bracts of male flowers not peltate. Stamens two.

Phylogeny (Baysian inference, simplified) of Moraceae based on DNA sequence data (Clement & Weiblen 2009).

Frangulaceae DC. in de Lamarck et A. P. de Candolle, Fl. Franç., ed. 3, 4(2): 619. 17 Sep 1805; Rhamnales Link, Handbuch 2: 118. 4-11 Jul 1829 [‘Rhamneae’]; Gouaniaceae Raf., Fl. Tellur. 2: 73. Jan-Mar 1837 [’Guanidia’]; Frangulopsida Endl., Gen. Plant.: 1081. Apr 1840; Rhamnopsida Brongn., Enum. Plant. Mus. Paris: xxxi, 121. 12 Aug 1843 [’Rhamnoideae’]; Phylicaceae J. Agardh, Theoria Syst. Plant.: 186. Apr-Sep 1858; Frangulales Wirtg., Anleit. Pflanzenk. 2: 36. 17-18 Sep 1860 [‘Frangulaceae’]; Ziziphaceae Adans. ex Post et Kuntze, Lex. Gen. Phan.: 665, 714. 20-30 Nov 1903 [’Zizyphaceae’]; Rhamnanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992

Distribution Cosmopolitan except polar areas, with their largest diversity in tropical and subtropical regions.

Fossils Coahuilanthus belinda from the Late Campanian of Mexico is represented by pentamerous bisexual flowers with antepetalous stamens and a prominent nectariferous disc. Floral fossils from Eocene strata of the Claiborne Formation (North America) and assigned to Solanites pusillus are apparently of Rhamnaceae origin (see Millan & Crepet 2014). Fossil leaves, pollen grains and fruits of Rhamnaceae are known from Oligocene strata onwards (e.g. fruits of Paliurus from North America and Europe).

Habit Usually bisexual (rarely monoecious, dioecious or androdioecious), evergreen or deciduous trees, shrubs or lianas (sometimes with tendrils or hooks; Crumenaria decumbens is a perennial herb), often with spines, often xeromorphic. Some species have phyllocladia and reduced leaves.

Vegetative anatomy Ectomycorrhiza present at least in Pomaderris and Rhamnus. Root nodules with nitrogen-fixing actinobacteria (e.g. Frankia) present in, i.a., Ceanothus and numerous species of Colletieae (Discaria, Colletia, Kentrothamnus, Retanilla and Trevoa). Phellogen ab initio superficial. Primary vascular tissue cylinder, without separate vascular bundles. Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits usually alternate (sometimes opposite), often confluent, simple or bordered pits. Vestured and non-vestured pits present. Imperforate tracheary xylem elements tracheids or libriform fibres with simple or bordered pits, usually non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, or banded (rarely apotracheal diffuse), or absent. Secondary phloem often stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Heartwood often with brown gum-like deposits. Secretory lysigenous mucilaginous cavities present or absent. Silica bodies present in wood ray cells of some species. Prismatic to diamond-shaped or acicular (sometimes styloids or crystal sand) calciumoxalate crystals abundant.

Trichomes Hairs unicellular or multicellular, usually simple (in Sageretia bifid, in Pomaderreae also or only stellate); glandular hairs?

Leaves Alternate (spiral or distichous) or opposite, simple, entire, sometimes strongly reduced, with conduplicate (-plicate) or involute ptyxis. Stipules small, persistent or caducous (sometimes intra- or interpetiolarly connate or modified into spines; petiolar in Colletia; absent in most species of Phylica); leaf sheath absent. Colleters present. Petiole vascular bundle transection arcuate or U-shaped. Venation usually pinnate (rarely palmate or leaves one-veined); secondary veins often parallel and prominent; tertiary veins scalariform. Stomata usually anomocytic (sometimes paracytic or anisocytic), sometimes sunken into cavities or furrows. Cuticular wax crystalloids as rosettes of platelets (Fabales type), cuticular wax as crust or smooth layer. Domatia usually absent (sometimes as pits, pockets or hair tufts). Pellucid dots present in lamina in Karwinskia. Epidermis often with lysigenous mucilaginous cavities. Mesophyll usually with mucilaginous idioblasts, often with calciumoxalate as druses or single prismatic crystals (in Gouanieae acicular styloids). Leaf margin serrate or entire. Extrafloral nectaries reported as rarely present as glands at leaf base in Gouania.

Inflorescence Usually axillary (sometimes terminal), umbellate, corymbose, raceme- or spike-like or panicle, thyrse or fascicle (flowers rarely solitary terminal).

Flowers Actinomorphic, small. Hypanthium present (sometimes long, tubular) or absent. Hypogyny to epigyny. Sepals (three or) four or five (or six), with valvate aestivation, with adaxial median ridge, persistent or caducous, usually free (rarely connate). Petals (three or) four or five (or six), with cucullate (induplicate-valvate) aestivation, often shortly clawed, often enclosing stamens, free (rarely absent). Nectariferous disc intrastaminal, entire or lobed, often adnate to hypanthium (receptacular nectaries; rarely absent).

Androecium Stamens (three or) four or five (or six), alternisepalous, antepetalous. Filaments thin, free, free from tepals or adnate to petal bases. Anthers very small, dorsifixed, often versatile, tetrasporangiate, usually introrse, longicidal (dehiscing by longitudinal slits or valves). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely tetracolporate), shed as monads, bicellular at dispersal. Exine usually tectate, with columellate (or granular) infratectum, microreticulate or perforate, striate, rugulate, fossulate, verrucate, pilate or psilate.

Gynoecium Pistil composed of two to five connate antesepalous carpels; median carpel adaxial. Ovary superior to inferior, bilocular to quinquelocular (sometimes unilocular by degeneration of carpels). Style single, simple, or stylodia several, more or less free, usually with two to four (rarely one, five or no) canals. Stigmas terminal, papillate, Dry type. Pistillodium absent. Common hypanthial and stylar indumentum sometimes forming secondary pollen display.

Ovules Placentation basal. Ovules usually one (in Karwinskia two median) per carpel, anatropous, ascending, apotropous or epitropous, bitegmic, crassinucellar. Micropyle usually exostomal (sometimes bistomal). Outer integument four to ten cell layers thick. Inner integument three or four cell layers thick. Obturator funicular. Hypostase sometimes present (in Ziziphus epistase). Parietal tissue four to seven cell layers thick. Archespore unicellular or multicellular. Nucellar cap (formed by apical cells of megasporangial epidermis) five or six cell layers thick; nucellar beak sometimes present; megasporangium often protruding through micropyle. Megagametophyte usually monosporous, Polygonum type (rarely disporous, 8-nucleate, Allium type). Synergids usually with a filiform apparatus. Antipodal cells degenerating. Endosperm development ab initio nuclear. Chalazal endosperm haustorium possibly present? Embryogenesis usually asterad (in Ziziphus solanad).

Fruit A drupe or a septicidal and/or loculicidal capsule (sometimes explosively dehiscent; rarely a nut; in Paliurus and Ventilago a samara; in most Gouanieae a schizocarp with two to five mericarps and often persistent sepals). Pericarp often distinctly two-layered (inner layer often lignified and finally spiralized), with layers often separating during maturation. Endocarp sclerenchymatous.

Seeds Funicular aril present in ziziphoid clade. Elaiosome present in some species. Seed often laterally flattened. Seed coat exotestal. Testa, usually multiplicative, often with median antiraphal vascular bundle. Exotesta often palisade, with malpighian cells with thickened walls. Mesotesta sometimes with sclerotic cells. Endotesta? Tegmen crushed. Endotegmen consisting of cuboid cells, with scalariform thickenings, usually not lignified (sometimes poorly lignified). Perisperm not developed. Endosperm sparse, usually oily (sometimes starchy; in Reynosia ruminate), or absent. Embryo large, usually straight (rarely curved), well differentiated, oily, with chlorophyll. Cotyledons two, large. Polyembryony common (in Ziziphus synergid polyembryony). Germination phanerocotylar or cryptocotylar.

DNA Trans-spliced intron present in mitochondrial gene nad1. Mitochondrial coxI intron present in numerous genera.

Phytochemistry Flavonols (kaempferol, quercetin), biflavonyls (biflavonoids), cyanidin, dammaranes, oleanolic acid derivatives, gallic acid, condensed tannins, proanthocyanidins (prodelphinidins), benzylisoquinoline alkaloids, liriodenine, peptide alkaloids, anthraquinone glycosides (in Rhamnus), saponins, naphthoquinones, arbutin, polyacetate derived arthroquinones, chelidonic acid, and naphthalene and lupeolin derivatives present. Myricetin, ellagic acids and cyanogenic compounds not found.

Use Ornamental plants, medicinal plants, fruits, timber, carpentries, dyeing substancies.

Systematics Rhamnaceae may be sister-group to the clade [Elaeagnaceae+[Barbeyaceae+Dirachmaceae]] (Zhang & al. 2011).

Three main clades can be recognized in Rhamnaceae (Richardson & al. 2000a): (1) rhamnoids (Ventilagineae, Maesopsideae and Rhamneae) with epigyny; (2) ampeloziziphoids (Bathiorhamnus, Ampeloziziphus and Doerpfeldia) with palmate foliar venation; and (3) ziziphoids (including Gouanieae, Paliureae, Colletieae, Phyliceae and Pomaderreae). Fay & al. (2001) received different results in their analyses. The classification below is based mainly on Richardson & al. (2000b).

A plausible topology (Richardson & al. 2000a, b) followed here is: [[Ventilagineae+[Maesopsideae+Rhamneae]]+[[Bathiorhamneae+[Ampelozizipheae+Doerpfeldieae]]+[Gouanieae+[Paliureae+[Lasiodiscus+[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]]]]]]

[Ventilagineae+[Maesopsideae+Rhamneae]]

Ventilagineae Benth. et Hook. f., Gen. Plant. 1: 372. 7 Aug 1862

2/20–45. Ventilago (9–35; tropical regions in the Old World, with one species, V. africana, in continental Africa and one species in Madagascar), Smythea (c 10; the Seychelles, Southeast Asia to Fiji and the Caroline Islands). – Tropical regions in the Old World.

[Maesopsideae+Rhamneae]

Maesopsideae Engl. et Weberb. in Engler et Prantl, Nat. Pflanzenfam. Nachtr.: 229. 4 Oct 1897

1/1. Maesopsis (1; M. eminii; tropical Africa).

Rhamneae Horan., Char. Ess. Fam.: 138. 17 Jun 1847

13/265–270. Rhamnus (c 120; temperate to tropical regions on the Northern Hemisphere south to South Africa and Brazil), Scutia (4; S. arenicola, S. buxifolia, S. myrtina, S. spicata; tropical regions in the Old World south to southern Africa, tropical South America), Sageretia (c 35; Somalia and southwestern Asia to Japan and Taiwan, tropical and subtropical Asia), Berchemiella (2; B. berchemiifolia: Japan; B. wilsonii: Hubei in China), Rhamnella (c 10; northern Pakistan to China, the Korean Peninsula and Japan, New Guinea, eastern Queensland, Fiji and Tonga), Dallachya (1; D. vitiensis; New Guinea, islands in western Pacific), Berchemia (c 20; East Africa to East Asia, New Caledonia, western North America), Rhamnidium (12; Central America, the West Indies, tropical South America), Karwinskia (16; southwestern United States and the West Indies to Bolivia), Condalia (18–20; southwestern United States, Mexico, Central America, tropical and subtropical South America), Auerodendron (10; the West Indies), Reynosia (c 15; Florida, Central America, the West Indies), Krugiodendron (2; K. acuminatum, K. ferreum; Mexico, Costa Rica, the West Indies). –

[[Bathiorhamneae+[Ampelozizipheae+Doerpfeldieae]]+[Gouanieae+[Paliureae+[Lasiodiscus+[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]]]]]

[Bathiorhamneae+[Ampelozizipheae+Doerpfeldieae]]

Bathiorhamneae J. E. Richardson in Kew Bull. 55: 335. 15 Aug 2000

1/7. Bathiorhamnus (7; B. capuronii, B. cryptophorus, B. dentatus, B. louvelii, B. macrocarpus, B. reticulatus, B. vohemarensis; Madagascar).

[Ampelozizipheae+Doerpfeldieae]

Ampelozizipheae J. E. Richardson in Kew Bull. 55: 335. 15 Aug 2000

1/2. Ampelozizyphus (2; A. amazonicus, A. guaquirensis; northern tropical South America).

Doerpfeldieae J. E. Richardson in Kew Bull. 55: 335. 15 Aug 2000

1/1. Doerpfeldia (1; D. cubensis; Cuba).

[Gouanieae+[Paliureae+[Lasiodiscus+[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]]]]

Gouanieae Rchb., Handb. Nat. Pfl.-Syst.: 222. 1-7 Oct 1837 [‘Gouaniaceae’]

5/c 63. Gouania (c 50; tropical and subtropical regions on both hemispheres), Helinus (5; H. brevipes, H. integrifolius, H. lanceolatus, H. mystacinus, H. spartioides; tropical and southern Africa, Madagascar, northwestern India), Reissekia (1; R. smilacina; Brazil), Alvimiantha (1; A. tricamerata; Brazil), Crumenaria (6; C. decumbens, C. dffusa, C. erecta, C. glaziovii, C. lilloi, C. steyermarkii; Central America, Colombia to Argentina). – Tropical and subtropical regions on both hemispheres.

[Paliureae+[Lasiodiscus+[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]]]

Paliureae Reissek ex Endl., Gen. Plant.: 1095. Apr 1840

3/110–160? Hovenia (3; H. acerba, H. dulcis, H. trichocarpa; the Himalayas, northern Burma, China, the Korean Peninsula, Japan), Paliurus (5; P. hemsleyanus, P. hirsutus, P. orientalis, P. ramosissimus, P. spina-christi; the Mediterranean to Japan; in Ziziphus?), ‘Ziziphus’ (100–150?; tropical and subtropical regions on both hemispheres; paraphyletic; incl. Paliurus?). – Tropical and subtropical regions on both hemispheres.

[Lasiodiscus+[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]]

Lasiodiscus clade

1/12. Lasiodiscus (12; tropical Africa, Madagascar).

[Emmenosperma+[Alphitonia+Granitites+Colubrina]+[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]]

Emmenosperma clade

1/5. Emmenosperma (5; E. alphitonoides, E. cunninghamii, E. micropetalum, E. pancherianum, E. papuanum; New Guinea, northern and eastern Australia, New Caledonia, Fiji).

Colubrina clade

3/c 52. Jaffrea (2; J. erubescens, J. xerocarpa; New Caledonia), Alphitonia (c 15; Malesia to New Guinea, northern and eastern Australia, islands in western Pacific, the Hawaiian Islands), Granitites (1; G. intangendus; southwestern Western Australia), Colubrina (34; tropical and subtropical regions on both hemispheres, with their highest diversity in tropical America). – Tropical and subtropical regions on both hemispheres.

[[Schistocarpaea+Colletieae]+[Phyliceae+[Ceanothus+Pomaderreae]]]

[Schistocarpaea+Colletieae]

Schistocarpaea clade

1/1. Schistocarpaea (1; S. johnsonii; northeastern Queensland).

Colletieae Reissek ex Endl., Gen. Plant.: 1099. Apr 1840

6/29. Colletia (5; C. hystrix, C. paradoxa, C. spartioides, C. spinosissima, C. ulicina; southern South America), Discaria (12; southeastern Queensland, eastern New South Wales, eastern Victoria, Tasmania, New Zealand, South America), Adolphia (2; A. californica, A. infesta; southwestern Unites States, northwestern Mexico), Kentrothamnus (1; K. weddellianus; Bolivia, Argentina), Retanilla (4; R. ephedra, R. patagonica, R. stricta, R. trinervia; Peru, Chile, western Argentina), Trevoa (5; T. campanulata, T. closiana, T. glauca, T. quinquenervia, T. spinifer; the Andes). – Southeastern Australia, Tasmania, New Zealand, southwestern Unites States, northwestern Mexico, South America.

[Phyliceae+[Ceanothus+Pomaderreae]]

Phyliceae Reissek ex Endl., Gen. Plant: 1100. Apr 1840

3–4/190–195. Phylica (c 190; southern Africa, Madagascar, the Mascarene Islands, Amsterdam Island, Tristan da Cunha, Gouth Island; incl. Nesiota?), Nesiota (1; N. elliptica; St. Helena; in Phylica?), Trichocephalus (1; T. stipularis; Western Cape), Noltea (1; N. africana; Western and Eastern Cape, KwaZulu-Natal). – Southern Africa, Madagascar, the Mascarene Islands, Amsterdam Island, St. Helena, Tristan da Cunha, Gouth Island.

[Ceanothus+Pomaderreae]

Ceanothus clade

1/c 55. Ceanothus (c 55; North America, Mexico, with their highest diversity in California and northwestern Mexico).

Pomaderreae Reissek ex Endl., Gen. Plant.: 1101. Apr 1840

10/210–220. Pomaderris (c 75; Southeast Asia, southern and eastern Australia, Tasmania, New Zealand), Siegfriedia (1; S. darwinioides; southwestern Western Australia; in Pomaderris?), Trymalium (14; southwestern Western Australia, southeastern South Australia), Spyridium (c 30; southern Australia, Tasmania), Stenanthemum (30–35?; southern and central Australia, Tasmania, with their highest diversity in southwestern Western Australia; non-monophyletic?), Cryptandra (55–60; Australia, Tasmania, with their largest diversity in southwestern Western Australia and southeastern Australia), Blackallia (1; B. nudiflora; southwestern Western Australia), Serichonus (1; S. gracilipes; southwestern Western Australia), Papistylus (2; P. grandiflorus, P. intropubens; southwestern Western Australia), Polianthion (4; P. bilocularis, P. collinum, P. minutiflorum, P. wichurae; southwestern Western Australia, southeastern Queensland). – Australia, Tasmania, New Zealand, with their highest diversity in southwestern Western Australia.

Cladogram (successive weighting) of Rhamnaceae based on DNA sequence data (Richardson & al. 2000a). According to Fay & al. (2001), [Alphitonia+Granitites] is sister-group to [Stenanthemum+Cryptandra+ [Siegfriedia+[Trymalium+[Spyridium+Pomaderris]]]] and Colubrina is sister to all other Rhamnaceae except [Bathiorhamnus+[Ampeloziziphus+Doerpfeldia]].

Spiraeaceae Bertuch, Taf. Allg. Naturgesch. Gewächs-Reich: Enum. 10, Syn. Tab. 3. 1801 [’Spiraeae’]; Poteriaceae Raf., Anal. Nat.: 173. Apr-Jul 1815 [’Poteria’]; Fragariaceae Nestl., Monogr. Potentilla: 14. Jun 1816; Pyraceae Vest, Anleit. Stud. Bot.: 268, 286. 1818 [’Pyroideae’]; Alchemillaceae Martinov, Tekhno-Bot. Slovar: 17. 3 Aug 1820 [’Alchemilleae’]; Amygdalaceae Marquis, Esq. Règne Vég.: 49. 15-22 Jul 1820 [’Amygdaleae’], nom. cons.; Potentillaceae Bercht. et J. Presl, Přir. Rostlin: 231. Jan-Apr 1820 [‘Potentilleae’]; Prunaceae Martinov, Tekhno-Bot. Slovar: 511. 3 Aug 1820 [‘Pruniferae’]; Prunales Bercht. et J. Presl, Přir. Rostlin: 230. Jan-Apr 1820 [‘Prunaceae’]; Sanguisorbaceae Bercht. et J. Presl, Přir. Rostlin: 231. Jan-Apr 1820 [‘Sanguisorbeae’]; Tormentillaceae Martinov, Tekhno-Bot. Slovar: 636. 3 Aug 1820 [’Tormentillae’]; Agrimoniaceae Gray, Nat. Arr. Brit. Pl. 2: 395, 574. 10 Jan 1822; Dryadaceae Gray, Nat. Arr. Brit. Pl. 2: 395, 577. 10 Jan 1822 [’Dryadeae’]; Ulmariaceae Gray, Nat. Arr. Brit. Pl. 2: 395, 588. 10 Jan 1822 [’Ulmariae’]; Amygdalales Link, Handbuch 2: 72. 4-11 Jul 1829 [’Amygdaleae’]; Dryadales Link, Handbuch 2: 98. 4-11 Jul 1829 [’Dryadeae’]; Sanguisorbales Link, Handbuch 2: 113. 4-11 Jul 1829 [‘Sanguisorbeae’]; Spiraeales Link, Handbuch 2: 94. 4-11 Jul 1829 [‘Spiraeaceae’]; Mespilaceae Schultz Sch., Nat. Syst. Pflanzenr.: 509. 30 Jan-10 Feb 1832 [’Mespileae’]; Rosineae Rchb., Deutsch. Bot. Herb.-Buch: lxvi. Jul 1841; Neilliaceae Miq., Fl. Ned. Ind. 1(1): 390. 20 Dec 1855; Cydoniaceae Schnizl., Anal. Nat. Ordn. Gew.: 14. 1856; Cercocarpaceae J. Agardh, Theoria Syst. Plant.: 287. Apr-Sep 1858 [’Cercocarpeae’]; Coleogynaceae J. Agardh, Theoria Syst. Plant.: 171. Apr-Sep 1858 [’Coleogyneae’]; Lindleyaceae J. Agardh, Theoria Syst. Plant.: 166. Apr-Sep 1858; Rhodotypaceae J. Agardh, Theoria Syst. Plant.: 172. Apr-Sep 1858 [’Rhodotypeae’]; Chamaemoraceae Lilja, Skånes Fl., ed. 2: 349, 980. Apr-Dec 1870 [’Chamaemoreae’]; Sorbaceae Brenner, Florist. Handb.: 94. 1886; Malaceae Small, Fl. S.E. U.S.: 529. 22 Jul 1903, nom. cons.

Distribution Cosmopolitan except continental Antarctica, with their largest diversity in temperate regions on the Northern Hemisphere.

Fossils Paleorosa similkameenensis is a flower (similar to those in Pyrodae) from the mid-Eocene of Canada and numerous other fossil Rosaceae have been recorded from the Eocene onwards in Europe and North America.

Habit Usually bisexual (sometimes monoecious, dioecious or polygamous), evergreen or deciduous trees, shrubs or suffrutices, or usually perennial (rarely annual) herbs. Some species are xerophytes.

Vegetative anatomy Ectomycorrhiza often present. Root nodules with nitrogen-fixing actinobacteria (e.g. Frankia) present at least in Dryadoideae. Phellogen usually deeply seated (in Exochorda subepidermal). Polydermis commonly present. Endodermis often prominent. Medulla without lysigenous mucilage canals, in some species septate by diaphragms. Vessel elements usually with simple (sometimes also reticulate or scalariform) perforation plates; lateral pits alternate or opposite, bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids or fibre tracheids (rarely libriform fibres) with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate to multiseriate, homocellular or heterocellular. Axial parenchyma usually apotracheal diffuse or diffuse-in-aggregates (sometimes paratracheal scanty or banded, rarely unilateral or vasicentric, or absent). Tyloses absent. Sieve tube plastids Ss type; cell nuclei with non-dispersive protein bodies. Nodes usually 3:3, trilacunar with three leaf traces (sometimes unilacunar or penta- to multilacunar with one or several traces, respectively). Heartwood often with gum-like substances. Calciumoxalate usually as solitary prismatic crystals (in Pruneae also druses) or absent (in Pyrinae prismatic crystals in axial parenchyma).

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, sometimes stellate; multicellular glandular hairs often frequent; prickles present in several genera.

Leaves Usually alternate (spiral; in Lyonothamnus, Coleogyne and Rhodotypos opposite), simple or pinnately or palmately compound, entire or lobed, sometimes coriaceous, usually with conduplicate ptyxis. Stipules inserted on branch or adnate to petiole, small to foliaceous, usually persistent (absent in Oemleria; in some species of Rosa with extrafloral nectaries); leaf sheath short? (with free margins) or absent. Petiole vascular bundle transection arcuate or annular. Venation usually pinnate (sometimes palmate), craspedodromous or camptodromous. Stomata anomocytic. Cuticular wax crystalloids often as rodlets or ribs (sometimes as granules or clustered tubuli of Berberis type, chemically dominated by nonacosan-10-ol). Domatia as pits, pockets or hair tufts, or absent. Epidermis with or without mucilaginous idioblasts. Cystoliths absent. Hydathodes sometimes present. Leaf margin and leaflet margins usually serrate (sometimes crenate or entire). Extrafloral nectariferous glands present on distal part of petiole or on base of lamina (sometimes on stipules), especially in Rosa and Prunus.

Inflorescence Terminal or axillary, usually cymose (sometimes racemose) of various shapes (cyme, panicle, raceme, corymb, umbel, fascicle, etc.), or flowers solitary axillary or terminal.

Flowers Usually actinomorphic (rarely zygomorphic). Hypanthium usually cupular, campanulate, tubular or discoid (rarely absent). Epicalyx, consisting of sepalous stipules of calyx lobes inserted on top of hypanthium, present in certain Rosoideae. Hypogyny, epigyny or half epigyny. Sepals (three or) four or five (to ten), with valvate or imbricate aestivation, persistent, free. Petals (three or) four or five (to ten), usually with imbricate aestivation, often clawed, caducous, free (rarely absent). Nectaries usually on adaxial side of hypanthium or on staminal bases. Nectariferous disc intrastaminal, annular or divided, inserted around hypanthial orifice (sometimes absent).

Androecium Stamens usually numerous (often 10+5+5, outer stamens antesepalous; rarely one [Aphanes clade of Alchemilla] or three, or five to ten), inflexed in bud, centripetally developing. Filaments free or connate below, free from tepals, usually inserted on hypanthium. Anthers dorsifixed, usually versatile, tetrasporangiate, usually introrse (in, e.g., Potentilla latrorse), longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores). Tapetum secretory. Staminodia usually absent (sometimes extrastaminal, petaloid).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpor(oid)ate (rarely triporate or, in Sanguisorbinae, pantoporate), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or striate.

Gynoecium Carpels one to numerous (sometimes spiral), often antesepalous or antepetalous, secondarily free or more or less connate or adnate to hypanthium (rarely somewhat stipitate, on gynophore). Ovary superior, inferior or semi-inferior, unilocular (apocarpy, monomery, sometimes epigyny + apocarpy), or bilocular to quinquelocular (syncarpy). Style single, simple, or stylodia few to numerous, usually terminal (in Rosoideae sometimes lateral or subbasal), free or more or less connate, or absent. Stigmas capitate, punctate, widened, decurrent or bifid (rarely fimbriate or penicillate), papillate or non-papillate, Dry or Wet type. Pistillodium?

Ovules Placentation usually basal or subbasal (rarely apical; when syncarpy then axile). Ovules usually one or two (sometimes several, rarely numerous) per carpel, usually anatropous (sometimes hemianatropous or campylotropous), usually ascending, apotropous (sometimes pendulous, epitropous), usually bitegmic (sometimes unitegmic), crassinucellar. Micropyle usually endostomal (rarely bistomal or almost absent). Outer integument five to 14 cell layers thick. Inner integument three to six cell layers thick. Obturator (in Pyrodae funicular) present in some genera. Parietal tissue two to four cell layers thick. Archespore multicellular. Nucellar cap approx. four cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes proliferating. Endosperm development usually ab initio nuclear (in Lyonothamnus helobial). Endosperm haustorium chalazal (in Prunus). Embryogenesis usually asterad.

Fruit An assemblage of follicles, an achene, an achenetum (multiple achenes from a single flower), a coccetum (multiple carpels from a single flower, separating at maturity and each dehiscing along a pair of sutures), a drupe, a drupetum (multiple drupelets from a single flower), a nuculanium (similar to drupe or drupetum but mesocarp not fleshy), a polyprenous drupe, a pome (carpels adnate to hypanthium, with fleshy layer consisting of hypanthial and sometimes carpellary tissue, often with numerous sclereids, and with unlignified endocarp); in Lindleya a capsule; in Rosa a cynarrhodium (multiple achenes enclosed by an urceolate carnose hypanthium); in Fragaria multiple achenes on surface of fleshy swollen floral receptacle; in Rubus an assemblage of partially or entirely connate drupelets.

Seeds Aril absent. Seed coat usually mesotestal (sometimes reduced and indistinct). Testa occasionally winged. Exotestal cells unspecified, periclinally elongate, with thickened radial walls, or palisade or tabular; cell walls spiral or with reticulate thickenings; outer wall often eventually mucilaginous. Mesotesta often sclerotic. Endotesta? Exotegmen? Endotegmic cells somewhat thickened. Perisperm usually not developed. Endosperm usually sparse or absent (in, e.g., Physocarpus and Rhodotypos copious). Embryo straight or curved, usually without chlorophyll. Cotyledons two, fleshy or flattened. Germination phanerocotylar or cryptocotylar.

Cytology x = 9 (Rosoideae, Dryadoideae, Spiraeoideae pro parte); x = 7 (8) (Rosoideae, Spiraeoideae pro parte); x = (9, 15–)16–17 (Pyrinae); n = 27 (Lyonothamnus) – A special type of maternal inheritance present in the Rosa canina-group (2n = 28 + 7). Agamospermy (diplospory, apogamety, apospory, pseudogamy etc.) occurring in several genera (e.g. Alchemilla, Crataegus, Potentilla, Rubus, and Sorbus).

DNA Cis-spliced intron present in mitochondrial gene nad1. Ancient duplication of GBSSI gene (gene coding for granule-bound starch synthase) resulting in two copies in all diploid Rosaceae with x = 7, 8 or 9, and another duplication (due to paleopolyploidization) resulting in two additional copies in diploid taxa with x = 15 or 17 (i.e. GBSSI-1A, GBSSI-1B, GBSSI-2A, andGBSSI-2B in Pyreae).

Phytochemistry Flavonols (kaempferol, quercetin), afzelechin, flavanone glycosides (especially in Prunus), biflavonoids, cyanidin, epigallocatechin-3-gallate, phlorizin and other dihydrochalcones, dammaranes, triterpenes, cucurbitacins, methylated and non-methylated ellagic acids, gallic acid, hydrolyzable ellagitannins (casuaricitin, geraniins, pedunculagin, stachyurins, tellimagrandin I, tellimagrandin II), proanthocyanidins (prodelphinidins), chlorogenic acid, phenylalanine-, tyrosine- or leucine-derived cyanogenic glucosides (amygdalin, cardiospermin, dhurrin, heterodendrin, prunasin), triterpene saponins, arbutin, acetophenones, eleostearic acid, condensed tannins (hydrolyzable tannins rare), indole alkaloids, and ursolic acid present. Sorbitol often replacing saccharose as transport carbohydrate.

Use Ornamental plants, fruits, perfumes (Rosa), medicinal plants, timber, carpentries.

Systematics Rosaceae are sister-group to the remaining Rosales.

Rosoideae are sister to [Dryadoideae+Spiraeoideae].

Rosoideae Arn., Botany: 107. 9 Mar 1832 [’Roseae’]

28–31/2.080–>2.840. Subcosmopolitan, with their highest diversity in temperate regions on the Northern Hemisphere. Usually perennial (rarely annual) herbs or shrubs (rarely trees). Wood rays often narrow. Leaves alternate, usually compound (sometimes simple). Cuticular wax crystalloids as narrow riblets and triangular rodlets. Hypanthium absent or more or less developed. Epicalyx (possibly modified stipules associated with sepals) present in Potentilleae. Receptacle sometimes enlarged. Hypanthium free from the separate ovaries. Carpels usually numerous. Ovule one per carpel, usually anatropous (rarely orthotropous), epitropous, unitegmic. Fruit an achenetum or drupetum. x = 7 (8). Ellagic acid and 2-pyrone-4,6-dicarboxylic acid present. Cyanogenic glycosides and sorbitol absent.

Filipendula clade

1/12. Filipendula (12; Europe, temperate Asia, northeastern North America). – Herbs. Receptacle enlarged. Ovules two per carpel. – Filipendula is sister to the remaining Rosoideae.

Rosodae T. Eriksson, Smedmark et M. S. Kerr in D. Potter et al. in Plant Syst. Evol. 266: 36. 2007

27–30/2.070–>2.830. Subcosmopolitan, with their highest diversity in temperate regions on the Northern Hemisphere.

Roseae Lam. et DC., Syn. Plant. Fl. Gall.: 331. 30 Jun 1806 [‘Rosae’]

1/100–150. Rosa (100–150; temperate regions on the Northern Hemisphere, tropical mountains in Ethiopia, the Philippines and Mexico). – Prickly arching shrubs. Hypanthium carnose, urceolate. Ovules two per carpel, collateral. Integuments approx. eight cell layers thick.

Rubeae Dumort., Anal. Fam. Plant.: 39. 1829

1/>250(–700). Rubus (>250(–700). sexual species; cosmopolitan except polar regions, with their highest diversity in temperate regions on the Northern Hemisphere). – Prickly scrambling shrubs. Receptacle enlarged. Ovules two per carpel. Integuments approx six cell layers thick. Fruit a drupetum.

Sanguisorbeae DC., Prodr. 2: 588. Nov (med.) 1825

12/300–310. Agrimoniinae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 502. 1846 [‘Agrimonieae’]. Agrimonia (12–15; temperate regions on the Northern Hemisphere south to Central and southern Africa, West Malesia, Hispaniola and Brazil), Leucosidea (1; L. sericea; South Africa, Lesotho, Zimbabwe), Aremonia (1; A. agrimonioides; southern and southeastern Europe), Spenceria (1; S. ramalana; western China), Hagenia (1; H. abyssinica; Central African mountains, Sudan, Ethiopia to Zimbabwe). – Sanguisorbinae Torr. et A. Gray, Fl. N. Amer. 1: 428. Jun 1840 [‘Sanguisorbeae’]. Cliffortia (c 120; southern Africa to Angola and Kenya, with their highest diversity in the Cape Provinces), 'Acaena' (c 100; temperate and subtropical regions on the Southern Hemisphere including Antarctic Islands and Macquarie Island, one species, A. pinnatifida, in California, one possibly extinct species, A. exigua, on Puu Kukui on Maui in the Hawaiian Islands; paraphyletic; incl. Margyricarpus and Tetraglochin?), Margyricarpus (c 8; South America), Tetraglochin (8; Peru to southern Argentina and central Chile), Polylepis (c 20; the Andes), Poterium (13; Europe, Madeira, the Canary Islands, the Mediterranean, North Africa, temperate Asia), Poteridium(1; P. annuum; western United States, northwestern Mexico), Sanguisorba (c 15; temperate regions on the Northern Hemisphere). – Subcosmopolitan, few in tropical Asia and tropical America. Carpels one to five. Integuments six to eight cell layers thick.

Potentilleae Sweet, Brit. Fl. Gard. 2: 124. 1 Sep 1825 [‘Potentillae’]

11–13/>1.360. Potentillinae J. Presl, Wšobecný Rostl. 1: 491. 1846 [‘Potentilleae’]. Potentilla (c 250; temperate and arctic regions on the Northern Hemisphere, few species in temperate regions on the Southern Hemisphere). – Fragariinae Torr. et A. Gray, Fl. N. Amer. 1: 435. Jun 1840 [’Fragarieae’]. Argentina (c 65; Eurasia to New Guinea), Alchemilla (>1.000; temperate regions on the Northern Hemisphere, tropical mountains in the Old World, southeastern New South Wales, eastern Victoria, South America), Comarum (1; C. palustre; temperate regions on the Northern Hemisphere), Farinopsis (1; F. salesoviana; the Himalayas, Central Asia, Siberia, Mongolia, China), Sibbaldia (2–4; S. procumbens, S. retusa; arctic and alpine regions on the Northern Hemisphere), Sibbaldianthe (2; S. adpressa, S. bifurca; eastern Europe to Siberia and western China; in Sibbaldia?), ‘Sibbaldiopsis’ cuneifolia (Pakistan, Kashmir, northern India, Nepal, Bhutan, Sikkim, Xizang in China; in Sibbaldia?); Dasiphora (3; D. davurica, D. fruticosa, D. parvifolia; temperate regions on the Northern Hemisphere), Chamaerhodos (7; C. altaica, C. canescens, C. corymbosa, C. erecta, C. grandiflora, C. sabulosa, C. trifida; Central and East Asia, western North America), Drymocallis (3; D. arguta, D. glandulosa, D. rupestris; Europe, the Mediterranean, temperate Asia), Chamaecallis (1; C. perpusilloides; Afghanistan, the Himalayas, southwestern China, Burma); Fragaria (c 20; temperate regions on both hemispheres). – Temperate, arctic and alpine regions on the Northern Hemisphere, tropical mountains, temperate regions on the Southern Hemisphere. Epicalyx sometimes present. Receptacle enlarged. Anther thecae in Fragariinae confluent. Style in Potentillinae often lateral/gynobasic. Integuments approx. four cell layers thick. Parietal tissue in Fragariinae approx. two cell layers thick and nucellar cap approx. seven cell layers thick.

Colurieae Rydb., N. Amer. Fl. 22. 240. 12 Jun 1908

2–3/c 60. Geum (c 50; temperate regions on both hemispheres), Oncostylus (7; Tasmania, New Zealand and adjacent islands, subantarctic South America; in Geum?), Fallugia (1; F. paradoxa; southwestern United States, northwestern Mexico). – Temperate regions on both hemispheres, tropical mountains, Chile. Ovule in Geum apotropous.

[Dryadoideae+Spiraeoideae]

Sorbitol present as transport carbohydrate. Cyanogenic glycosides present.

Dryadoideae Sweet, Brit. Fl. Gard. 1: ad t. 43. Apr 1830 [‘Dryadeae’]

4/21. Dryas (2; D. integrifolia: eastern Siberia, Greenland, northern and western North America; D. octopetala: arctic and alpine regions from eastern Greenland through Europe, the Caucasus and northern Asia to western North America), Cercocarpus (9; western United States, northwestern Mexico), Chamaebatia (2; C. australis: southern California, northern Baja California in northwestern Mexico; C. foliolosa: mountains in California), Purshia (8; southwestern Canada, western United States, northwestern Mexico). – Temperate, arctic and alpine regions on the Northern Hemisphere. Shrubs, shrublets or small trees. Ectomycorrhiza sometimes present (at least in Cercocarpus). Symbiosis with root-dwelling nitrogen fixing Frankia. Leaves compound in Chamaebatia, otherwise simple. Stipules present. Hypanthium free from ovary. Carpels one to numerous. Pistil single (Cercocarpus, Chamaebatia, Purshia) or four to many (Dryas, Purshia). Carpel in Cercocarpus and Chamaebatia single. Placentation in Cercocarpus and Chamaebatia basal. Ovules usually orthotropous (in Dryas anatropous, apotropous). Obturator absent in Cercocarpus and Chamaebatia. Fruits achenes or achenetum with hairy styles. x = 9. Polyploidy not known. Cyanogenic glycosides present. Sorbitol present in trace amounts.

Spiraeoideae Arn., Botany: 107. 9 Mar 1832 [‘Spiraeeae’]

45/1.040–>1.330. Usually shrubs or trees. Leaves usually alternate, simple. Stipules usually present. Cuticular wax crystalloids as tubules or platelets. Hypanthium usually free from ovary/ovaries. Carpels one to five, antepetalous, usually free. Stigma usually Wet type. Ovules two or more per carpel. Papillate funicular obturator present. Fruit a follicetum, an achene, an achenetum, a coccetum, a drupe, a drupetum, a polyprenous drupe, a nuculanium, or a pome (in Pyrinae). x = 8, 9, 15, 17. Flavones, cyanogenic glycosides and sorbitol (often abundant) present. Ellagic acid not found.

Lyonothamneae Brouillet in J. Bot. Res. Inst. Texas 2: 385. 24 Jul 2008

1/1. Lyonothamnus (1; L. floribundus; Channel Islands off southern California). – Leaves opposite, compound. Stipules deciduous. Half epigyny. Placentation apical. Ovules four to six per carpel, apical, epitropous. Cyanogenic glycosides not found. – Relictual endemic. Three fossil species of Lyonothamnus have been described and the group had a significantly larger distribution during the Neogene (Erwin & Shorn 2000).

Amygdaleae DC., Prodr. 2: 529. Nov (med.) 1825

1/200–430. Prunus (200–430; temperate regions on both hemispheres, tropical mountains). – Usually bisexual (dioecious in ‘Maddenia clade’) Ectomycorrhiza often present. Phellogen superficial. Tracheids absent. Leaves with vertically or laterally conduplicate ptyxis. Extrafloral nectaries present on petiole or abaxial side of lamina. Petals absent in ‘Maddenia clade’. Carpel single. Ovules two per carpel, hemianatropous, unitegmic. Outer integument six to eight cell layers thick. Inner integument three to six cell layers thick. Obturator developed from ovary wall. Fruit a drupe. Seed coat usually pachychalazal. x = 8.

Neillieae Maxim. in Trudy Imp. S.-Peterburgsk. Bot. Sada 6: 164, 216. Jul-Aug 1879

2/25–32. Neillia (15–17; eastern Himalayas, China, mountains in West Malesia), Physocarpus (10–15; northeastern Asia, eastern and western North America). – Eastern Himalayas and China to northeastern Asia, mountains in West Malesia, North America. Placentation apical. Ovule one per carpel, apotropous (sometimes up to five per carpel, pleurotropous). Micropyle in Physocarpus bistomal. Fruitlets hard, shiny.

Sorbarieae Rydb., N. Amer. Fl. 22: 239, 256. 12 Jun 1908

4/13. Adenostoma (2; A. fasciculatum: western United States, northern Baja California in northwestern Mexico; A. sparsifolium: southern California, northern Baja California), Chamaebatiaria (1; C. millefolium; western United States), Sorbaria (c 9; temperate East Asia), Spiraeanthus (1; S. schrenkianus; Kazakhstan, Kyrgyzstan?). – Central to East Asia, western United States, northwestern Mexico. Leaves usually compound (in Adenostoma simple). Cuticular wax crystalloids in Adenostoma, Chamaebatiaria and Sorbaria transversely ridged, as rodlets. Carpel in Adenostoma single. Placentation apical. Ovules usually epitropous. Micropyle in Adenostoma, Chamaebatiaria and Sorbaria bistomal. Obturator (of different types) usually present. Fruit in Adenostoma an achene.

Spiraeeae DC., Prodr. 2: 541. Nov (med.) 1825 [‘Spiraeaceae’]

8/90–110. Aruncus (1; A. dioicus; temperate regions on the Northern Hemisphere), Holodiscus (1; H. discolor; western Canada, western United States, Mexico, Central America, Colombia), Kelseya (1; K. uniflora; Rocky Mountains in western United States), Eriogynia (1; E. pectinata; Rocky Mountains in northwestern North America), Petrophytum (3; P. caespitosum, P. cinerascens, P. hendersonii; Rocky Mountains in western North America), Sibiraea (4; S. angustata, S. laevigata, S. tianschanica, S. tomentosa; the Balkan Peninsula, Siberia, Central and East Asia), Spiraea (80–100; temperate regions on the Northern Hemisphere south to the Himalayas and Mexico), Xerospiraea (1; X. hartwegiana; Mexico). – Temperate regions on the Northern Hemisphere south to mountains in Colombia. Vestured pits present. Nodes 1:1 (unilacunar with one leaf trace). Stipules absent. Placentation in Spiraea, Holodiscus and Aruncus apical. Ovules six to eight per carpel, epitropous, unitegmic. Obturator present in Spiraea, Holodiscus and Aruncus. Fruit in Holodiscus an achene.

Kerriodae D. Potter, S. H. Oh et K. R. Robertson in D. Potter et al. in Plant Syst. Evol. 266: 38. 2007

7/10. Central to East Asia, western and southeastern North America, northwestern Mexico.

Exochordeae Schulze-Menz ex Reveal J. Bot. Res. Inst. Texas 4(1): 215. 29 Jul 2010 (Osmaronieae Rydb., N. Amer. Fl. 22: 240. 1918, nom. illeg.)

3/5. Oemleria (1; O. cerasiformis; southwestern Canada, western United States), Exochorda (1; E. racemosa; Central Asia to China), Prinsepia (3; P. sinensis, P. uniflora, P. utilis; the Himalayas to northern China and Taiwan). – Central Asia to the Himalayas and East Asia, western North America. Phellogen superficial. Medulla septated. Stipules caducous. Stylodia lateral. Obturator developed from ovary wall. Fruit a drupe or a septicidal capsule (in Exochorda also adaxially dehiscing). x = 8.

Kerrieae Focke in Engler et Prantl, Nat. Pflanzenfam. III, 3: 12, 27. Dec 1888

4/5. Coleogyne (1; C. ramosissima; southwestern United States, northwestern Mexico), Rhodotypos (1; R. scandens; China, the Korean Peninsula, Japan), Neviusia (3; N. alabamensis: southeastern United States; N. cliftonii: Shasta County in northern California), Kerria (1; K. japonica; temperate East Asia). – Temperate East Asia, southwestern and southeastern United States, northwestern Mexico. Carpels one to five. Megasporangium in Rhodotypos protruding. Fruit achenetum-like. n = 8 or 9 (Neviusia). Sorbitol and flavones present in Neviusia. Ellagic acid not found in Neviusia.

Pyrodae C. S. Campbell, R. C. Evans, D. R. Morgan et T. A. Dickinson in D. Potter et al., Plant Syst. Evol. 266: 39. 2007

22/700–>730. Ectomycorrhiza present. Phellogen superficial. Wood rays often narrow. Colleters present. Carpels (one or) two to five, antesepalous or median carpel abaxial, adnate to base of hypanthium, often connate. Gynoecial ring primordium present. Placentation basal (not when ovules numerous). Ovules (one or) two per carpel, anatropous, apotropous, bitegmic. Micropyle sometimes bistomal. Funicular obturator papillate. Exotestal cells thickened, with lignified walls, often mucilaginous. Mesotesta thick, sclerotic. x = (9, 15–)16–17. Flavone-C-glycosides present.

Gillenieae Maxim. in Trudy Imp. S.-Peterburgsk. Bot. Sada 6: 164, 222. Jul-Dec 1879

1/2. Gillenia (2; G. stipulata, G. trifoliata; southeastern United States). – Perennial herbs. Leaves compound. x = 9. – Gillenia is sister to the remaining Pyrodae.

Pyreae Baill., Hist. Plant. 1: 442, 475. Jan-Jul 1869

21/695–>725. x = 15, 17 (aneuploid reduction from x = 18). GBSSI present in four copies.

Kageneckia clade

2/4. Kageneckia (3; K. angustifolia, K. lanceolata, K. oblonga; Peru, Chile), Lindleya (1; L. mespiloides; Mexico). – Mexico, Peru. Chile. Ovules four to numerous per carpel, pleurotropous.

Vauquelinia clade

1/2. Vauquelinia (2; V. californica, V. corymbosa; southwestern United States, northwestern Mexico). – Tanniniferous cells pervasive. Capsule septicidal; carpels dehiscing adaxially (and partially abaxially).

Pyrinae Dumort., Fl. Belg.: 92. 1827 [‘Pyreae’]

18/690–>720. Dichotomanthes (1; D. tristaniicarpa; southwestern China), Eriobotrya (11–15; the Himalayas, East Asia, mountains in West Malesia), Rhaphiolepis (5; R. ferruginea, R. indica, R. lanceolata, R. major, R. salicifolia; East and Southeast Asia), Sorbus (130–140; temperate regions on the Northern Hemisphere), Amelanchier (c 20; temperate regions on the Northern Hemisphere south to Mexico and Guatemala), Pyrus (15–20; Europe, the Mediterranean, temperate Asia), Aronia (c 40; the Himalayas to Japan and Sumatra, eastern North America, Central America), Malus (c 40; temperate regions on the Northern Hemisphere), Cydonia (1; C. oblonga, the Caucasus, Kurdistan), Docynia (1; D. indica; eastern Himalayas, southern China, northern Thailand), Pseudocydonia (1; P. sinensis; China), Chaenomeles (3; C. cathayensis, C. japonica, C. speciosa; China, the Korean Peninsula, Japan), Cotoneaster (>260; temperate regions in the Old World), Pyracantha (7; P. coccinea: northeastern Spain to northern Iran; P. crenulata: the Himalayas; P. angustifolia, P. atalantioides, P. crenatoserrata, P. rogersiana: southwestern to central China; P. koidzumii: Taiwan), Crataegus (140–150; temperate regions on the Northern Hemisphere), Osteomeles (1–3; O. anthyllidifolia, O. schwerinae, O. subrotunda; China, the Ryukyu islands, Taiwan, the Cook Islands, Tonga, Pitcairn Island, Rapa Iti, the Hawaiian Islands), Hesperomeles (11; Central America, Colombia to Peru and Bolivia), Chamaemeles (1; C. coriacea; Madeira). – Temperate, alpine and arctic regions on the Northern Hemisphere and southwards to mountains in Sumatra and Bolivia. Leaves usually simple (in Osteomeles and species of Sorbus compound). Stipules deciduous. Usually half epigyny (in Dichotomanthes hypogyny). Carpels sometimes laterally free (in Dichotomanthes single). Outer integument five to 14 cell layers thick. Inner integument three to six cell layers thick. Hypantium carnose in fruit. Endocarp usually absent.

Cladogram (simplified) of Rosaceae based on DNA sequence data (Potter & al. 2007).

Distribution Temperate regions in North America and Eurasia, tropical Africa, India, Central and northern South America, with their highest diversity in temperate regions.

Fossils Leaves, pollen grains and fruits of Ulmaceae are abundant in Cenozoic layers from the Paleocene onwards in the Northern Hemisphere. Cedrelospermum, consisting of fossilized samaras, has been described from the Eocene of North America and from the Late Eocene to mid-Miocene of Europe. Late Cretaceous records of Ulmaceae are questioned.

Habit Bisexual, monoecious, andromonoecious or polygamomonoecious, evergreen or deciduous trees or shrubs. Prophylls usually basal.

Vegetative anatomy Ectomycorrhiza often present. Phellogen ab initio superficial. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements sometimes very long libriform fibres usually with simple (sometimes bordered) pits, septate (also vasicentric tracheids). Wood rays uniseriate to multiseriate, usually homocellular. Axial parenchyma usually paratracheal scanty, aliform, winged-aliform, confluent, vasicentric, or banded (sometimes apotracheal diffuse). Wood sometimes fluorescent. Wood elements often storied. Tyloses frequent. Secondary phloem stratified into hard fibrous and soft parenchymatous non-fibrous layers. Sieve tube plastids usually S type (in Ulmus Pc type with protein crystalloids); sieve tube elements with extruded nucleoli and non-dispersive protein bodies. Nodes 3:3, trilacunar with three leaf traces. Laticifers and latex absent. Heartwood sometimes with gum-like substances. Mucilaginous idioblasts and canals present. Cystoliths globose (calciumcarbonate?), usually without pegs (cellulose expansions of cell wall, covered by crystals; in Hemiptelea with pegs). Epidermal cell walls often with calciumcarbonate or silica. Prismatic calciumoxalate crystals frequent.

Trichomes Hairs usually rigid, unicellular (not micropapillar) or multicellular, simple, often with swollen base or calcified/silicified (hairs in Ampelocera smooth); glandular hairs?

Leaves Usually alternate (usually distichous; rarely opposite), simple, entire, usually with laterally (rarely vertically) conduplicate-plicate ptyxis. Stipules usually lateral and non-axillary, usually extrapetiolar (one of two stipules in Ulmus intrapetiolar), usually caducous; leaf sheath absent. Petiole vascular bundle transection? Leaf base often asymmetrical. Venation pinnate, usually craspedodromous; secondary and tertiary veins usually proceeding into leaf teeth (urticoid teeth; not in Ampelocera). Stomata anomocytic or paracytic. Cuticular wax crystalloids? Domatia as pockets or hair tufts, or absent. Epidermis and mesophyll often with mucilaginous idioblasts. Leaf margin usually coarsely serrate (rarely entire). Leaf usually with rigid unicellular hairs with swollen base, or glabrous.

Inflorescence Axillary, usually cymose (often fascicle; rarely racemose, or flowers solitary).

Flowers Actinomorphic, small to very small. Hypanthium present. Hypogyny. Sepals (three to) five (to nine), usually with imbricate (sometimes valvate) aestivation, usually spiral, persistent, free or partially connate. Petals absent. Nectary absent. Disc absent.

Androecium Stamens (four or) five (rarely eight, ten, or twelve), antesepalous. Filaments filiform, usually free, often adnate to sepal bases. Anthers dorsifixed, often versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to quadrinucleate cells. Female flowers sometimes with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains 4–5(–7)-stephanoporate, shed as monads, usually bicellular (in Ulmus tricellular) at dispersal. Exine tectate, with granular infratectum, rugulate, spinulate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, usually unilocular (due to pseudomonomery; in Ulmus bilocular). Stylodia usually two (in some species of Zelkova sometimes one). Stigmas decurrent, ventral, papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation subapical. Ovule one per ovary, usually anatropous (sometimes orthotropous), pendulous, epitropous, bitegmic, crassinucellar (to tenuinucellar?). Micropyle usually bistomal (in Ulmus endostomal), not Z-shaped. Outer integument four to six cell layers thick. Inner integument approx. four cell layers thick. Hypostase present. Megagametophyte usually monosporous, Polygonum type (in Ulmus tetrasporous, usually Drusa type, rarely Adoxa type). Antipodal cells sometimes proliferating (forming up to ten cells in Holoptelea). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad (Ulmus) or onagrad (Holoptelea).

Fruit A samara (in Ulmus with reticulately veined wings; in Planera with coriaceous appendages; in Zelkova an unwinged nut).

Seeds Aril absent. Seed flattened. Testa thin, indistinct. Exotestal cells elongate, usually not thickened (in Holoptelea with thickened walls). Endotesta? Tegmen? Perisperm not developed. Endosperm sparse or absent. Embryo straight (in Zelkova curved), well differentiated, without chlorophyll. Cotyledons two, flattened or longitudinally folded. Germination phanerocotylar.

Cytology n = 14, 28, (30) 42 (x = 14) – Polyploidy occurring. Chromosomes often with terminal or subterminal diffuse and complex centromers.

DNA Trans-spliced intron present in mitochondrial gene nad1. Deletion of 69 bp in plastid gene ndhF.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin) and their glycosides, cyanidin, proanthocyanidins (prodelphinidins), alkaloids (in Ulmus), sesquiterpene lactones, and lignans present. Saponins sometimes present? Ellagic acid and cyanogenic compounds not found. Raffinose and stachyose present in phloem exudates. Aluminium accumulated in some species.

Parietariaceae Bercht. et J. Presl, Přir. Rostlin: 261. Jan-Apr 1820 [’Parietariae’]; Urticales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 260. Jan-Apr 1820 [‘Urticeae’]; Urticopsida Bartl., Ord. Nat. Plant: 91, 102. Sep 1830 [’Urticinae’]; Cecropiaceae C. C. Berg in Taxon 27: 39, 23 Mar 1978; Urticanae Takht. ex Reveal in Novon 2: 237. 13 Oct 1992

Distribution Cosmopolitan except polar areas, with their largest diversity in tropical Asia.

Fossils Leaves and achenes of Urticaceae are known from the Cenozoic. Many Late Cretaceous fossils have been attributed to the Urticaceae (from Central Europe etc.), although none of them is unambiguous.

Habit Usually monoecious, polygamomonoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees, shrubs, lianas or suffrutices, perennial or annual herbs (in Coussapoa and Poikilospermum usually epiphytes). Young stems and branches often quadrangular in cross-section. Some arborescent species have buttresses. Cecropia often live in symbiosis with ants.

Vegetative anatomy Phellogen ab initio usually superficial (in, e.g., Urtica cortical). Primary medullary rays often narrow, alternating with wide ones. Vessel elements with simple perforation plates; lateral pits alternate, usually simple (sometimes bordered) pits. Imperforate tracheary xylem elements very long and strong libriform fibres, usually with simple (sometimes bordered) pits, usually septate. Wood rays usually multiseriate (sometimes uniseriate), usually heterocellular (sometimes homocellular). Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, or banded. Wood sometimes fluorescent. Wood elements usually storied. Tyloses often abundant. Secondary phloem stratified into hard fibrous and soft parenchymatous non-fibrous layers. Sieve tube plastids usually Ss type, with one to three starch grains, or S₀ type (cell nuclei in Laportea with non-dispersive P-protein bodies); sieve tube elements with extruded nucleoli and non-dispersive protein bodies. Nodes usually 3:3, trilacunar with three leaf traces (sometimes bilacunar or pentalacunar?). Laticifers and latex cells usually absent (rarely present in bark); latex usually absent (white latex rarely present). Cystoliths usually present (absent in, e.g., Cecropia), globose (calciumcarbonate?) or elongate (with pegs?). Prismatic calciumoxalate crystals abundant; druses sometimes present; raphides present in Dendrocnide.

Trichomes Hairs unicellular (usually micropapillate) or multicellular, simple, bristle-like and non-stinging, often curled, arachnoid or hooked, or stinging (in Urticeae with, e.g., 5-hydroxytryptamine); pearl glands rarely present.

Leaves Alternate (spiral or distichous) or opposite, simple, entire or (sometimes deeply) trilobate to septalobate, sometimes anisophyllous (in some species of Pilea peltate), with laterally or vertically conduplicate ptyxis. Stipules lateral, intrapetiolar or interpetiolar, often sheathingly connate or branch-enclosing, in Cecropia and Musanga very large; absent in several genera of Parietarieae); leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Leaf base often asymmetrical. Venation pinnate or palmate; secondary and tertiary veins proceeding into non-glandular teeth (urticoid teeth). Stomata usually anomocytic or anisocytic (sometimes helicocytic or paracytic). Cuticular wax crystalloids? Epidermal cells often with elongate to punctiform cystoliths (cystoliths sometimes absent); epidermal cell walls often with calciumcarbonate or silica. Mesophyll usually with mucilaginous idioblasts and ducts, often with calciumoxalate druses. Leaf margin serrate, sinuate or entire.

Inflorescence Axillary, panicle, spicate, capitate, umbellate or catkin- or spadix-like cymose (flowers rarely solitary), often with male inflorescence raceme- or catkin-like and female inflorescence capitate, often with involucroid bracts. Inflorescence axis in Elatostemateae flattened, discoid and fleshy. Inflorescence sometimes composed of spikes and surrounded by large involucroid caducous bract. Prophylls usually basal.

Flowers Usually actinomorphic (female flowers often zygomorphic), small to minute. Pedicel often articulated. Usually hypogyny (rarely half epigyny). Sepals in male flowers (one to) four or five (or six), with valvate aestivation, persistent, free or connate below (rarely entirely connate); in female flowers three to five, valvate, often unequal in size, often accrescent in fruit, free or entirely or almost entirely connate (rarely rudimentary or absent). Petals absent. Nectary absent. Disc absent.

Androecium Stamens (one to) four or five (or six), antesepalous, usually as many as sepals. Filaments usually inflexed in bud and at anthesis recurved with lightning rapidity (in Cecropia etc. usually erect), usually free from each other, free from tepals. Anthers basifixed to dorsifixed, usually non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits), usually explosively dehiscent. Tapetum secretory. Female often with staminodia, sometimes scale-like (in Pilea and other Elatostemateae often enhancing fruit dispersal by discharging ripe achenes through instantaneous straightening of staminodial filaments).

Pollen grains Microsporogenesis simultaneous. Pollen grains (1–)2–6(–15)-porate, shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, spinulate.

Gynoecium Pistil composed of two connate carpels (one of which entirely or almost entirely degenerating and sterile). Ovary usually superior, unilocular (pseudomonomery), often oblique or asymmetrical. Style single, simple, short, or absent. Stigma capitate, penicillate, ligulate or filiform (rarely peltate), papillate, Dry type. Male flowers usually with pistillodium.

Ovules Placentation basal or subbasal. Ovule one per carpel, anatropous, hemianatropous or orthotropous, ascending to upright, epitropous, bitegmic, crassinucellar. Funicle present or absent. Micropyle usually endostomal (in Parietaria bistomal). Outer integument approx. two cell layers thick. Inner integument approx. two cell layers thick, sometimes obturator. Hypostase present in at least Urtica. Parietal tissue four to six cell layers thick. Nucellar cap two to four cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes proliferating. Endosperm development ab initio nuclear. Endosperm haustorium chalazal (at least in Urtica). Embryogenesis asterad.

Fruit Usually a nutlet or an achene (rarely a drupe), often surrounded by (sometimes adnate to) persistent and accrescent (sometimes carnose) calyx; often a syncarp consisting of connate ovaries from several flowers.

Seeds Aril absent. Testa thin, crushed. Exotesta usually perforated. Some testal and tegmic layers persistent. Perisperm not developed. Endosperm copious to sparse, often oily or proteinaceous (sometimes starchy), or absent. Embryo small, straight, well differentiated, without chlorophyll. Cotyledons two, usually flattened (sometimes thick). Germination? Adventitious embryony and polyembryony occurring in Elatostema.

Cytology x = 6–14, 16, 19, 21 – Polyploidy frequently occurring. Centromers median and subterminal? Protein bodies present in nucleus.

DNA Trans-spliced intron present in mitochondrial gene nad1. Mitochondrial coxI intron present in Pilea.

Phytochemistry Flavonols (kaempferol, quercetin), flavone-C-glycosides, cyanidin, alkaloids, caffeic acid derivatives, polyacetate-derived arthroquinones, and furanocoumarins present. Ellagic acid, saponins and cyanogenic compounds not found.

Use Ornamental plants, textile plants (fibres of Boehmeria nivea, Girardinia, Laportea, Urtica, etc.), medicinal plants (Parietaria etc.), fruits (Cecropia, Pourouma), vegetables, timber.

Systematics Urticaceae are sister to Moraceae.

Cecropieae are sister-group to the clade [Urticeae+Elatostemateae] (Hadiah & al. 2008).

Cecropieae Dumort., Anal. Fam. Plant.: 17. 1829

29/435–485. Leucosyke (c 50; Malesia to Polynesia), Coussapoa (50–55; tropical South America), Cecropia (c 60; southern Mexico, Central America, the West Indies, tropical South America), Musanga (2; M. cecropioides, M. leo-errerae; tropical Africa), Myrianthus (4–7; M. arboreus, M. holstii, M. preussii, M. serratus; tropical Africa), Pourouma (25–30; Central America, tropical South America); ‘Parietaria’ (c 20; tropical to temperate regions on both hemispheres; paraphyletic; incl. Gesnouinia?, Soleirolia?), Gesnouinia (1; G. arborea; the Canary Islands; in Parietaria?), Soleirolia (1; S. soleirolii; Italy, islands in western Mediterranean; in Parietaria?); Forsskaolea (6; F. angustifolia, F. candida, F. hereroensis, F. procridifolia, F. tenacissima, F. viridis; southeastern Spain, the Canary Islands, the Cape Verde Islands, Africa, the Arabian Peninsula, India), Didymodoxa (3; D. caffra, D. capensis, D. catira; southern and eastern tropical Africa to northern Ethiopia), Droguetia (7; D. ambigua, D. debilis, D. gaudichaudiana, D. hildebrandtii, D. humbertii, D. iners, D. leptostachys; tropical to subtropical Africa, northeastern India to Yunnan, Taiwan and Java; incl. Australina?), Australina (2; A. flaccida, A. pusilla; Ethiopia, Kenya, southeastern Australia, New Zealand; in Droguetia?); Oreocnide (c 18; China, Japan, tropical Asia to New Guinea); Phenax (14; tropical America); Chamabainia (1; C. cuspidata; tropical Asia, Taiwan), Gonostegia (5; G. hirta, G. parvifolia, G. pentandra, G. quinquenervis, G. triandra; southern China, Southeast Asia to northern Australia), ‘Pouzolzia’ (c 35; tropical regions on both hemispheres; non-monophyletic), Neodistemon (1; N. indicum; tropical Asia), Hemistylus (4; H. boehmerioides, H. brasiliensis, H. macrostachya, H. odontophylla; tropical South America), Rousselia (2; R. guilhoti, R. humilis; Central America, the West Indies, Colombia), Pipturus (35–40; the Mascarene Islands, tropical Asia to New Guinea, Northern Territory and Queensland, Melanesia, Polynesia incl. the Hawaiian Islands), Neraudia (5; N. angulata, N. kauaiensis, N. melastomifolia, N. ovata, N. sericea; the Hawaiian Islands); ’Boehmeria’ (65–80; tropical regions on both hemispheres, subtropical regions on the Northern Hemisphere; polyphyletic), Debregeasia (3; D. longifolia, D. orientalis, D. saeneb; Ethiopia, tropical and subtropical Asia), Astrothalamus (1; A. reticulatus; West and Central Malesia), Archiboehmeria (1; A. atrata; Southeast Asia, southern China), Sarcochlamys (1; S. pulcherrima; tropical Asia), ’Cypholophus’ (15–30; Malesia to New Guinea and islands in western Pacific, Taiwan; polyphyletic). – Warm-temperate to tropical regions on both hemispheres.

[Urticeae+Elatostemateae]

Urticeae Lam. et DC., Syn. Plant. Fl. Gall.: 184. 30 Jun 1806

10/205–210. Nanocnide (2; N. japonica, N. lobata; China, northern Vietnam, the Korean Peninsula, Japan, Taiwan), Zhengyia (1; Z. shennongensis; Hubei province in central China), Urtica (c 80; nearly cosmopolitan), Dendrocnide (37; tropical Asia to New Guinea, eastern Queensland, eastern New South Wales and islands in the Pacific), Discocnide (1; D. mexicanus; Mexico, Guatemala), Girardinia (3; G. bullosa, G. diversifolia, G. palmata; East and Northeast Africa, tropical and subtropical regions in Africa and Asia); ‘Laportea’ (22; tropical and subtropical regions on both hemispheres, East Asia, eastern North America; diphyletic), ‘Urera’ (c 35; tropical and southern Africa, Madagascar, the Hawaiian Islands, tropical America; non-monophyletic; incl. Obetia?, Poikilospermum?), Obetia (8; tropical and southern Africa, Madagascar, the Mascarene Islands; in Urera?), Poikilospermum (c 20; eastern Himalayas, Southeast Asia, Malesia; in Urera?). – Subcosmopolitan. Stinging hairs are a synapomorphy of Urticeae. – Laportea is diphyletic, according to Deng & al. (2013). Zhengyia is sister-group to Urtica. The clade [Urtica pilulifera + U. neubaueri] is sister to all other species of Urtica. Urera is paraphyletic.

Elatostemateae Gaudich. in H. L. C. de Saulces de Freycinet, Voy. Uranie, Bot.: 493. 6 Mar 1830 [’Elatostemeae’]

7–8/950–>1.000. ’Elatostema’ (c 300; tropical regions in the Old World, one species, E. rugosum, in New Zealand; non-monophyletic; incl. Procris?), Procris (c 15; tropical regions in the Old World; in Elatostema?); Gyrotaenia (5; G. crassifolia, G. microcarpa, G. myriocarpa, G. spicata, G. trujilloana; the West Indies), Myriocarpa (18; southern Mexico, Central America, tropical South America), Meniscogyne (1; M. thorelii; Laos, Vietnam), Lecanthus (3; L. peduncularis, L. petelotii, L. pileoides; tropical regions in the Old World), Petelotiella (1; P. tonkinensis; Indochina), Pilea (600–>700; tropical and subtropical regions on both hemispheres). – Pantropical. – Water-storing hydrenchyma present in succulent species of Pilea.

Unplaced Urticaceae

Haroldiella (2; H. rapaensis, H. sykesii; French Polynesia, Rapa Island), Metatrophis (1; M. margaretae; Rapa Island).

Cladogram of Urticaceae based on DNA sequence data (Sytsma & al. 2002).

Cladogram (simplified) of Urticaceae based on DNA sequence data (Hadiah & al. 2008).

Cladogram (simplified) of Urticaceae based on trnL/F sequence data (Hadiah & al. 2008). Girardinia is sister to Dendrocnide, according to Deng & al. (2013).

Phylogeny (simplified) of Urticaceae based on DNA sequence data (Wu & al. 2013, Treiber & al. 2016)

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”The Nitrogen Fixing clade”

ROSALES Bercht. et J. Presl

BARBEYACEAE Rendle

CANNABACEAE Martinov

DIRACHMACEAE Hutch.

ELAEAGNACEAE Juss.

MORACEAE Gaudich.

RHAMNACEAE Juss.

ROSACEAE Juss.

ULMACEAE Mirb.

URTICACEAE Juss.