Including Asperifoliaceae Reichb., Heliotropiaceae Schrad., Onosm(at)aceae Horan., Scorpiaceae Dulac
Excluding Ehretiaceae, Wellstediaceae
Habit and leaf form. Trees, or shrubs, or herbs, or lianas (a few); without essential oils. Autotrophic. Annual to perennial (often hispid or scabrid); with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting (usually), or climbing. Helophytic, or mesophytic, or xerophytic. Leaves alternate, or alternate and opposite (then opposite below); flat; petiolate to sessile; non-sheathing; not gland-dotted; simple; epulvinate. Lamina entire; usually narrow, linear to lanceolate. Leaves exstipulate. Lamina margins entire (always?). Leaves without a persistent basal meristem. Domatia recorded, or not recorded.
Leaf anatomy. Abaxial epidermis papillose (e.g. Cordia spp), or not papillose (?). Mucilaginous epidermis absent. Stomata on both surfaces (usually); usually anomocytic. Hairs usually present (the herbs especially characterized by hispid leaves); eglandular, or eglandular and glandular. Multicellular hairs uniseriate; branched, or unbranched. Urticating hairs absent.
Adaxial hypodermis absent. Lamina dorsiventral, or isobilateral; without secretory cavities. Cystoliths commonly present (at the bases of the hairs). The mesophyll commonly containing calcium oxalate crystals. The mesophyll crystals raphides, or druses, or solitary-prismatic. Midrib conspicuous. Main veins vertically transcurrent, or embedded. Minor leaf veins with phloem transfer cells (usually - in 17 genera), or without phloem transfer cells (Caccinea, Heliotropium).
Stem anatomy. Secretory cavities absent. Cork cambium usually present; initially deep-seated, or superficial. Nodes unilacunar. Primary vascular tissue in a cylinder, without separate bundles; centrifugal. Cortical bundles absent. Medullary bundles absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. The secondary phloem not stratified. `Included' phloem absent. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls horizontal; simple (usually), or reticulately perforated. Vessels without vestured pits. Primary medullary rays wide (in woody species), or narrow. Wood ring porous to semi-ring porous; storied, or partially storied, or not storied; parenchyma apotracheal.
Reproductive type, pollination. Hermaphrodite, or gynodioecious (e.g. in Echium). Predominantly entomophilous; via hymenoptera.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in `inflorescences'; in cymes. The terminal inflorescence unit cymose (coiled at first). Inflorescences terminal, or axillary, or leaf-opposed; usually coiled cincinnial, sometimes double; not pseudanthial. Flowers bracteate; bracteolate; usually regular, or somewhat irregular (Echium and relatives); 5 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk present, or absent.
Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous. Calyx 5; 1 whorled; polysepalous, or gamosepalous (basally). Degree of gamosepaly, maximum length joined/total calyx length less than 0.25. Calyx basally appendaged (e.g. Myosurus), or neither appendaged nor spurred; imbricate, or open in bud, or valvate (rarely). Corolla 5; 1 whorled; appendiculate (often, with a corona of scales from the throat protecting the nectar), or not appendiculate; gamopetalous. Degree of gamopetaly 0.5-0.8. Corolla imbricate, or contorted; campanulate to hypocrateriform; unequal but not bilabiate, or regular; white, or yellow, or pink, or blue.
Androecium 5. Androecial members unbranched; adnate (to the corolla); all equal, or markedly unequal; free of one another, or coherent; 1 - whorled. Androecium exclusively of fertile stamens. Stamens 5; not didynamous, not tetradynamous; isomerous with the perianth; oppositisepalous (epipetalous at the mouth, or on the tube). Filaments appendiculate, or not appendiculate. Anthers cohering, or separate from one another, or connivent (sometimes somewhat coherent at base and apex); dorsifixed to basifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate; appendaged, or unappendaged. The anther appendages when present, basal. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer; of the `dicot' type. Tapetum glandular. Pollen grains aperturate; 3-20 - aperturate (to `many'); colpate, or porate, or colporate, or colpate and colporate (alternating); 2-celled, or 3-celled.
Gynoecium 2 (usually), or 4-5 (in Trigonotis); syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 2 locular (rarely), or 4 locular (usually, via false septa), or 10 locular (via false septa, in Trigonotis). Gynoecium median. Styles 1; from a depression at the top of the ovary; `gynobasic', or apical (in Heliotropoideae). Stigmas 1-2; when simple, 1-2 - lobed; nearly always dry type; papillate; Group II type (usually), or Group III type (rarely). Placentation axile to basal. Ovules 2 per locule (usually separating into one-ovuled portions); horizontal to ascending; epitropous; non-arillate; anatropous, or hemianatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type, or Allium-type (sometimes). Polar nuclei fusing prior to fertilization, or fusing only after one has been fertilized (?). Antipodal cells formed; 3; not proliferating; ephemeral, or persistent. Synergids pear-shaped, or hooked. Endosperm formation cellular to nuclear. Endosperm haustoria present, or absent; micropylar (Heliotropium). Embryogeny onagrad, or chenopodiad.
Fruit fleshy, or non-fleshy; when dry dehiscent, or indehiscent, or a schizocarp; when schizocarpic, comprising nutlets, or comprising drupelets; of separable pyrenes (two or four nutlets), or a drupe (1-4 seeded). Seeds endospermic, or non-endospermic. Cotyledons 2. Embryo achlorophyllous (14/17); straight, or curved.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, biochemistry. Cyanogenic (rarely), or not cyanogenic. Alkaloids present, or absent. Iridoids absent. Proanthocyanidins absent. Flavonols present (usually), or absent (Echium); kaempferol and quercetin (usually), or quercetin. Ellagic acid absent (11 species, 9 genera). Saponins/sapogenins absent. Inulin recorded (Cynoglossum). C3 and C4. C3 recorded in Arnebia, Coldenia, Heliotropium, Lappula, Lithospermum, Moltkiopsis, Onosmodium, Trichodesma. C4 recorded in Heliotropium. Anatomy C4 type (Heliotropium), or non-C4 type (Arnebia, Cynoglossum, Echiochilon, Heliotropium, Heterocaryum, Lappula, Lithospermum, Myosotis, Onosma, Onosmodium, Trichodesma, Vaupelia).
Geography, cytology. Holarctic, Paleotropical, Neotropical, Cape, Australian, and Antarctic. Temperate to tropical. Cosmopolitan, but fewer in cool temperate and tropical regions, and with a strong Mediterranean concentration. X = 4-12.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Solaniflorae; Boraginales. Cronquist's Subclass Asteridae; Lamiales. Takhtajan's Subclass Asteridae; Lamianae; Polemoniales. Species 2000. Genera about 120; Actinocarya, Adelocaryum, Afrotysonia, Alkanna, Amblynotus, Amphibologyne, Amsinckia, Anchusa, Ancistrocarya, Anoplocaryum, Antiotrema, Antiphytum, Arnebia, Asperugo, Auxemma, Borago, Bothriospermum, Brachybotrys, Brunnera, Buglossoides, Caccinia, Carmona, Cerinthe, Chionocharis, Choriantha, Craniospermum, Cryptantha, Cynoglossopsis, Cynoglossum, Cynoglottis, Cysostemon, Dasynotus, Decalepidanthus, Echiochilon, Echiostachys, Echium, Elizaldia, Embadium, Eritrichium, Gastrocotyle, Gyrocaryum, Hackelia, Halacsya, Heliocarya, Heliotropium, Heterocaryum, Huynhia, Ivanjohnstonia, Ixorhea, Lacaitaea, Lappula, Lasiarrhenum, Lasiocaryum, Lepechiniella, Lepidocordia, Lindelophia, Lithodora, Lithospermum, Lobostemon, Macromeria, Maharanga, Mairetis, Mattiastrum, Mertensia, Metaeritrichium, Microcaryum, Microula, Mimophytum, Moltkia, Moltkiopsis, Moritzia, Myosotidium, Myosotis, Neatostema, Nesocaryum, Nogalia, Nomosa, Nonea, Ogastemma, Omphalodes, Omphalolappula, Omphalotrigonotis, Onosma, Onosmodium, Oxyosmyles, Paracaryum, Pardoglossum, Patagonula, Pectocarya, Pentaglottis, Perittostema, Plagiobothrys, Pseudomertensia, Psilolaemus, Pteleocarpa, Pulmonaria, Rindera, Rochefortia, Rochelia, Rotula, Saccellium, Scapicephalus, Selkirkia, Sericostoma, Sinojohnstonia, Solenanthus, Stenosolenium, Stephanocaryum, Suchtelenia, Symphytum, Thaumatocaryum, Thyrocarpus, Tianschaniella, Tiquilia, Tournefortia, Trachelanthus, Trachystemon, Trichodesma, Trigonocaryum, Trigonotis, Ulugbekia, Valentiniella.
Economic uses, etc. Ornamentals, pot herbs, dyes for wood, stone, medicines, wines and cosmetics, and some important honey plants.
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