The families of flowering plants.
IncludingAlchemillaceae J.G. Agardh, Amygdalaceae D. Don, Annulaceae Dulac, Cercocarpaceae J.G. Agardh, Cliffortiaceae Mart., Coleogynaceae J.G. Agardh, Drupaceae S.F. Gray, Dryadeae (Dryadaceae) S.F. Gray, Fragariaceae Rich. ex Nestle., Lindleyaceae J.G. Agardh, Malaceae Small, Neilliaceae Miquel, Pomaceae S.F. Gray, Potentilleae (Potentillaceae) Trautv., Prunaceae Burnett, Rhodotypaceae J.G. Agardh, Sanguisorbeae, (Sanguisorbaceae) Loisel., Spiraeaceae Bertuch, Ulmariae (Ulmariaceae) S.F. Gray
Habit and leaf form. Trees, or shrubs, or herbs. The herbs usually perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting, or climbing (sometimes); the climbers scrambling. Trees and shrubs leptocaul (often with ‘short shoots’). Helophytic, or mesophytic, or xerophytic. Leaves deciduous (usually), or evergreen;alternate (except Rhodotypos); spiral; ‘herbaceous’, or leathery; nearly always petiolate; sheathing, or non-sheathing. Leaf sheaths when leaves sheathing, with free margins. Leaves not gland-dotted; simple, or compound; epulvinate; when compound ternate, or pinnate, or palmate. Lamina when simple dissected, or entire; pinnately veined (usually), or palmately veined (e.g. Alchemilla); cross-venulate. Leaves stipulate (usually), or exstipulate (in Spiraea etc.). Stipules intrapetiolar (often adnate to the petiole); free of one another; scaly, or leafy. Lamina margins crenate, or serrate, or dentate, or entire (rather infrequently). Vegetative buds scaly. Leaves without a persistent basal meristem. Domatia occurring in the family (from 6 woody genera); manifested as pits, or pockets, or hair tufts.
Leaf anatomy. Hydathodes present (occasionally), or absent. Mucilaginous epidermis present, or absent. Stomataanomocytic.
Lamina dorsiventral (usually), or centric (Crataegus azarolus). Cystolithsabsent. Minor leaf veins without phloem transfer cells (6 genera, including both herbaceous and woody).
Stem anatomy. Cork cambium present, or absent (?); initially deep-seated, or superficial. Nodes tri-lacunar (usually), or unilacunar, or penta-lacunar to multilacunar. Cortical bundles absent. Medullary bundles absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent. Xylem with tracheids (mostly), or without tracheids (Prunoideae); with vessels. Vessel end-walls simple, or scalariform and simple. Vessels without vestured pits. Wood parenchyma typically apotracheal, or apotracheal and paratracheal (there being some scanty-paratracheal recorded in a few genera). Pith with diaphragms (occasionally), or without diaphragms.
Reproductive type, pollination. Fertile flowershermaphrodite. Unisexual flowers absent. Plants hermaphrodite (usually). Pollination anemophilous (occasionally, e.g. Poterium), or entomophilous (usually); commonly pollinated by flies; mechanism conspicuously specialized (occasionally), or unspecialized.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes, in panicles, in racemes, in corymbs, in umbels, and in fascicles. The ultimate inflorescence unit when flowers aggregated cymose (usually), or racemose. Inflorescences terminal, or axillary; pseudanthial (occasionally), or not pseudanthial. Flowers small to large; often fragrant; usually regular; cyclic, or partially acyclic. Sometimes the gynoecium acyclic. Flowers tetracyclic to polycyclic. Floral receptacle developing a gynophore (rarely), or with neither androphore nor gynophore; markedly hollowed (Pomoideae, most Prunoideae, some Rosoideae (e.g. Rosa)), or not markedly hollowed (flattish or only slightly concave in Spiraeoideae, convex or swollen in some Rosoideae (e.g. Fragaria, Rubus)). Free hypanthium present (nearly always, the flower generally more or less perigynous), or absent. Hypogynous disk present (often), or absent; intrastaminal.
Perianthwith distinct calyx and corolla (usually), or sepaline; (5–)10(–20); 2 whorled (usually), or 1 whorled (when apetalous); usually isomerous. Calyx (3–)5(–10) (usually green and sepaloid); 1 whorled; polysepalous, or gamosepalous; regular; usually imbricate; when determinable, with the median member posterior. Epicalyx present (often, its members alternating with the calyx and seemingly representing its stipules), or absent. Corolla when present, (3–)5(–10); 1 whorled; polypetalous; usually imbricate; regular; white, or yellow, or red, or pink (but not blue).
Androecium (1–)10–100 (usually ‘many’). Androecial members branched (often, in the sense that they are often disposed in pairs), or unbranched; when determinable, ‘at least sometimes’ maturing centripetally (Cronquist 1981);free of the perianth (but usually attached to a hypanthium); free of one another, or coherent; often more or less 5 adelphous, or 10 adelphous, or 15 adelphous (etc. — in 5-adelphous whorls); 1–5 whorled (?). Androecium exclusively of fertile stamens (normally), or including staminodes (in cultivars). Staminodes when present, 5–50 (several to ‘many’); external to the fertile stamens; when present, usually petaloid. Stamens (1–)20–100 (usually ‘many’); reduced in number relative to the adjacent perianth to isomerous with the perianth (rarely), or diplostemonous to polystemonous (usually); often with an outer whorl of 10 in five antesepalous pairs; inflexed in bud. Anthers dorsifixed; versatile (mostly), or non-versatile (rarely, e.g. Filipendula); dehiscing via longitudinal slits, or dehiscing via pores (terminal, occasionally); introrse (mostly), or latrorse (e.g. Potentilla); unilocular (sometimes, e.g.Alchemilla), or bilocular; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with more than one middle layer (2 or 3). Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; 3 aperturate (usually), or 4–9 aperturate (?or more, rarely); colporate (or colporoidate, usually), or foraminate (Sanguisorba); hardly ever spinulose; 2-celled (in 14 genera).
Gynoecium 1–50 carpelled. Carpels isomerous with the perianth, or reduced in number relative to the perianth, or increased in number relative to the perianth. The pistil when other than apocarpous, 1 celled, or 2–5 celled. Gynoecium monomerous, or apocarpous, or syncarpous (Maloideae); of one carpel, or eu-apocarpous, or semicarpous, or synovarious (Maloideae); superior, or partly inferior, or inferior (occasionally even combining an inferior ovary with free carpels). Carpel when monomeric or apocarpous non-stylate, or stylate; apically stigmatic, or with a lateral style, or with a gynobasic style; when apocarpous 1–2 ovuled (usually), or 3–10 ovuled (Spiraeoideae).Placentation when apocarpous or one-carpelled marginal, or apical.Ovary when syncarpous, 2–5 locular (Maloideae). Styleswhen syncarpous, 2–5; free. Stigmas wet type, or dry type; papillate, or non-papillate; Group II type, Group III type, and Group IV type. Placentationwhen syncarpous, axile. Ovules 1(–2) per locule; pendulous, or ascending; non-arillate; anatropous (nearly always), or hemianatropous, or campylotropous; unitegmic, or bitegmic; crassinucellate. Outer integument contributing to the micropyle, or not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization, or fusing simultaneously with the male gamete. Antipodal cells formed; 3; proliferating (sometimes, e.g in Alchemilla), or not proliferating; ephemeral (usually), or persistent. Synergids pear-shaped. Endosperm formation nuclear. Embryogeny asterad.
Fruit fleshy, or non-fleshy; an aggregate (when carpels free), or not an aggregate. The fruiting carpels (when carpels free) commonly coalescing into a secondary syncarp (with small achenes or drupelets), or not coalescing. The fruiting carpel when apocarpous dehiscent, or indehiscent; a follicle, or an achene, or drupaceous, or baccate. Fruit when syncarpous dehiscent (rarely), or indehiscent; a capsule (Lindleyella), or a berry (sometimes in the guise of a pome), or a drupe; enclosed in the fleshy receptacle (or attached to it), or enclosed in the fleshy hypanthium, or without fleshy investment. The drupes with separable pyrenes, or with one stone. Seeds non-endospermic (nearly always), or endospermic (copiously so in Physocarpus).Perisperm absent. Seeds winged (rarely, e.g. Exochorda), or wingless. Cotyledons 2 (usually expanded and flat). Embryo achlorophyllous (31/64), or chlorophyllous (Rhaphiolepis umbellata); straight, or curved, or bent. Micropyle not zigzag.
Seedling.Germination phanerocotylar (e.g. Crataegus, Malus), or cryptocotylar (e.g. Prunus persica).
Physiology, biochemistry. Cyanogenic (very commonly), or not cyanogenic. Cynogenic constituents phenylalanine-derived, or leucine-derived. Alkaloids absent (nearly always), or present (very rarely). Iridoids not detected. Proanthocyanidins present, or absent; cyanidin (with the sole exception of a single species of Potentilla). Flavonols present (nearly always); kaempferol and quercetin (nearly always). Ellagic acid variously present (numerous Rubineae, Potentillineae, Dryadineae, Cercocarpeae, Ulmarieae, Sanguisorbieae, Roseae), or absent (numerous Spiraeoideae, Pomoideae, Kerrieae, Prunoideae). Arbutin present, or absent. Ursolic acid present. Saponins/sapogenins present, or absent. Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose (a quite wide sample found consistently depauperate in oligosaccharides). Inulinnot found. C3. C3 physiology recorded directly in Fragaria, Malus, Physocarpus,Potentilla, Prunus, Rosa, Rubus, Sorbus, Spiraea. Anatomy non-C4 type (Potentilla, Prunus, Rosa).
Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = 7–9, 17 (or more).
Taxonomy.Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Rosiflorae; Rosales. Cronquist’s Subclass Rosidae; Rosales. APG 3 core angiosperms; core eudicot; Superorder Rosanae; fabid; Order Rosales.
Species about 2000. Genera about 100; Acaena, Adenostoma, Agrimonia,Alchemilla, Amelanchier, Aphanes, Aremonia, Aria,Aruncus, Bencomia, Brachycaulos, Cerocarpus, Chaenomeles,Chamaebatia, Chamaebatiaria, Chamaemeles, Chamaemespilus,Chamaerhodos, Cliffortia, Coleogyne, Coluria, Cormus,Cotoneaster, Cowania, Crataegus, Cydonia, Dalibarda,Dichotomanthes, Docynia, Docyniopsis, Dryas, Duchesnea,Eriobotrya, Eriolobus, Exochorda, Fallugia, Filipendula,Fragaria, Geum, Gillenia, Hagenia, Hesperomeles,Heteromeles, Holodiscus, Horkelia, Horkeliella, Ivesia,Kageneckia, Kelseya, Kerria, Leucosidea, Lindleya,Luetkea, Lyonothamnus, Maddenia, Malacomeles, Malus,Margyricarpus, Mespilus, Neillia, Neviusia, Nuttalia, Oemleria, Orthurus, Osteomeles, Pentactina, Peraphyllum,Petrophytum, Photinia, Physocarpus, Polylepis, Potanina,Potentilla, Poterium, Prinsepia, Prunus, Pseudocydonia,Purshia, Pyracantha, Pyrus, Rhaphiolepis, Rhodotypos,Rosa, Rubus, Sanguisorba, Sarcopoterium, Sibbaldia,Sibiraea, Sorbaria, Sorbus, Spenceria, Spiraea,Spiraeanthus, Stephanandra, Taihangia, Tetraglochin,Torminalis, Vauquelinia, Waldsteinia, Xerospiraea.
Economic uses, etc. Edible fruits from Malus spp. (apples), Prunus spp. (apricot, cherry, nectarine, peach, plums, prune, sloe), Cydonia (quince), Pyrus (pear), Eriobotrya (loquat), Rubus (blackberry, boysenberry, loganberry, rasberry), Fragaria (strawberry), Mespilus (medlar); nuts (Prunus — almond); many ornamental trees and shrubs or hedge-plants, e.g. Spiraea, Photinia, Kerria, Cotoneaster,Pyracantha, Crataegus, Sorbus, Rhodotypos, Prunus,Rosa, Potentilla; and some widely naturalized pests (e.g. from Acaena,Crataegus, Cotoneaster, Pyracantha).