Including Antirrhin(ac)eae DC. & Duby, Aragoaceae D. Don, Hebenstreitiaceae Horan., Limosell(ac)eae J.G. Agardh, Melampyraceae Lindl., Paulowniaceae Nak., Pedicularidaceae(-`lares') Juss., Personaceae Dulac, Rhinanthaceae(`-oideae') Vent., Selaginaceae Choisy, Sibthorpiaceae D. Don
Excluding Ellisiophyllaceae
Habit and leaf form. Shrubs and herbs (mostly), or trees, or lianas. `Normal' plants, or plants of very peculiar form (when parasitic). Leaves well developed (usually), or much reduced (e.g. the parasitic Harveya, Hyobanche), or absent (e.g. the root parasite Lathraea). Totally parasitic (Lathraea), or partially parasitic (commonly, concentrated in the Rhinantheae), or autotrophic. When parasitic, on roots of the host. Annual, or perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Climbing (sometimes), or self supporting (mainly); the climbers stem twiners, or petiole twiners. Hydrophytic (e.g. Ambulia), or helophytic, or mesophytic, or xerophytic; when hydrophytic, rooted. Leaves of aquatics emergent. Heterophyllous (e.g. Hebe, Hydrotriche), or not heterophyllous. Leaves alternate, or opposite, or whorled; when alternate spiral, or four-ranked; `herbaceous', or leathery, or membranous (rarely); petiolate to sessile, or perfoliate (occasionally); connate (occasionally?), or not connate (usually); sheathing, or non-sheathing; simple; epulvinate. Lamina dissected, or entire; if dissected pinnatifid, or much-divided (e.g. submerged leaves in Hydrotriche, Limnophila); pinnately veined. Leaves exstipulate. Lamina margins entire, or crenate, or serrate, or dentate. Leaves without a persistent basal meristem.
Leaf anatomy. Hydathodes present (occasionally), or absent. Stomata present; mainly confined to one surface, or on both surfaces; anomocytic, or anisocytic.
Adaxial hypodermis present (rarely), or absent. Lamina dorsiventral, or isobilateral. Cystoliths present (occasionally), or absent. Minor leaf veins with phloem transfer cells (9 genera, e.g. Antirrhinum, Rhinanthus), or without phloem transfer cells (16 genera, e.g. Pedicularis, Scrophularia, Verbascum).
Stem anatomy. Cork cambium present, or absent; initially when present. deep-seated, or superficial. Nodes unilacunar. Primary vascular tissue usually in a cylinder, without separate bundles; centrifugal. Secondary thickening developing from a conventional cambial ring. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls simple. Vessels without vestured pits. Wood partially storied (VPI, Penstemon), or not storied; parenchyma if present, apotracheal, or paratracheal (usually very sparse or absent). Sieve-tube plastids S-type. Pith with diaphragms, or without diaphragms.
Reproductive type, pollination. Hermaphrodite. Entomophilous. Pollination mechanism conspicuously specialized (with loose-pollen mechanisms), or unspecialized.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in `inflorescences'; in cymes, in racemes, and in spikes. The terminal inflorescence unit cymose, or racemose. Inflorescences terminal, or axillary; mainly racemes, spikes and thyrses, terminal peloric flowers common. Flowers bracteate; bracteolate; small to medium-sized; very irregular (usually - apart from peloric terminal flowers), or somewhat irregular (e.g. Verbascum). The floral asymmetry involving the perianth and involving the androecium, or involving the androecium. Flowers cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk present.
Perianth with distinct calyx and corolla; 8-10(-13); 2 whorled; isomerous, or anisomerous. Calyx 4 (the posterior member missing, or the anterior pair united), or 5; 1 whorled; gamosepalous; unequal but not bilabiate, or bilabiate, or regular; persistent; imbricate, or valvate; when K5, with the odd member posterior. Corolla 4 (the posterior pair united), or 5(-8); 1 whorled; gamopetalous; imbricate, or valvate; bilabiate (usually), or unequal but not bilabiate, or regular (Verbascum, etc.); spurred (sometimes), or not spurred.
Androecium (4-)5 (posterior member sometimes missing), or 2(-3) (sometimes the lower pair reduced or missing). Androecial members adnate (to the corolla); usually markedly unequal; free of one another; 1 - whorled. Androecium exclusively of fertile stamens, or including staminodes. Staminodes when present 1 (the posterior member), or 2-3. Stamens (2-)4(-5); usually didynamous; reduced in number relative to the adjacent perianth, or isomerous with the perianth; oppositisepalous. Anthers connivent (in pairs), or separate from one another; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer (1 or 2); of the `dicot' type. Tapetum glandular. Pollen grains aperturate; 2-7 - aperturate; colporate (commonly, or colporoidate), or colpate; 2-celled.
Gynoecium 2(-3); syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 2(-3) locular. Gynoecium median; stylate. Styles 1; attenuate from the ovary, or from a depression at the top of the ovary; apical. Stigmas 1; 1-2 - lobed; wet type, or dry type; papillate; Group II type and Group III type. Placentation axile. Ovules 2-50 per locule (i.e. to `many'); pendulous (Selaginaceae); non-arillate; anatropous, or campylotropous, or hemianatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type, or Allium-type, or Drusa-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 2 (Mimulus, one being binucleate), or 3; not proliferating; ephemeral to persistent. Synergids pear-shaped, or hooked. Hypostase present, or absent. Endosperm formation cellular. Endosperm haustoria present (usually), or absent; when developed, chalazal and micropylar (usually), or chalazal, or micropylar. Embryogeny onagrad, or solanad.
Fruit non-fleshy (usually), or fleshy (rarely); dehiscent (usually), or indehiscent (rarely), or a schizocarp (Lagotis); when non-schizocarpic, a capsule (usually), or a berry. Capsules septicidal (usually), or loculicidal, or poricidal (occasionally). Seeds endospermic. Endosperm oily. Seeds small; not conspicuously hairy; winged, or wingless (often angled); without amyloid. Embryo well differentiated (usually), or weakly differentiated (Lathraea). Cotyledons 2 (reduced but distinguishable in Lathraea). Embryo chlorophyllous (3/3), or achlorophyllous (12/26); straight to curved.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic (rarely), or not cyanogenic. Cynogenic constituents phenylalanine-derived. Alkaloids present, or absent (mostly). Iridoids recorded (commonly, including Selaginaceae); carbocyclic and seco-compounds. Proanthocyanidins absent. Flavonols to all intents and purposes, absent. Ellagic acid absent (13 species, 9 genera). Saponins/sapogenins present, or absent. Aluminium accumulation not found. C3 and C4. C3 recorded in Agalinis, Antirrhinum, Castilleja, Gratiola, Linaria, Lindenbergia, Mimulus, Orthocarpus, Pentstemon. C4 recorded in Anticharis. Anatomy non-C4 type (Agalinis, Castilleja, Gratiola, Limnophila, Linaria, Orthocarpus, Penstemon, Scrophularia).
Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = 6 (or more).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Lamiiflorae; Scrophulariales. Cronquist's Subclass Asteridae; Scrophulariales. Takhtajan's Subclass Asteridae; Lamianae; Scrophulariales. Species 3000. Genera about 280; Alectra, Alonsoa, Antirrhinum, Aptosimum, Bartsia, Calceolaria, Castilleja, Celsia, Chelone, Collinsia, Cymbalaria, Digitalis, Euphrasia, Gerardia, Gratiola, Hebenstretia, Halleria, Linaria, Maurandia, Melampyrum, Mimulus, Paulownia, Pedicularis, Penstemon, Rhinanthus, Rhodochiton, Russelia, Scrophularia, Selago, Sutera, Torenia, Tozzia, Verbascum, Veronica, Wightia, Zaluzianska, etc.
Economic uses, etc. Many are poisonous, a few are (e.g. Digitalis) or have been officinal, Halleria has edible fruit (umbinza).
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