The families of flowering plants
~ Former Liliaceae, Xanthorrhoeaceae-Asphodeloideae of APG III.
Including Aloaceae (Aloëaceae, Aloeaceae) J.G. Agardh
Habit and leaf form. Herbs (mostly), or shrubs, or arborescent (some of the woody forms with trunks up to several metres). Plants succulent, or non-succulent. Perennial; with a basal aggregation of leaves (commonly), or with terminal aggregations of leaves (when woody); rhizomatous (always?). Helophytic, or mesophytic, or xerophytic. Leaves small to very large; alternate; spiral (usually), or distichous; herbaceous, or leathery, or fleshy, or leathery and fleshy; sessile; sheathing. Leaf sheaths with free margins. Leaves normally orientated; simple. Lamina entire; linear, or lanceolate, or ovate, or subulate (etc.); parallel-veined (but the veins often invisible externally); without cross-venules. Lamina margins entire, or serrate, or dentate (and often with an apical spine). Leaf development graminaceous.
Leaf anatomy. The leaf lamina dorsiventral, or centric. Stomata present; on both surfaces; anomocytic, or tetracytic, or paracytic (rarely). The mesophyll containing mucilage cells (with raphides); containing crystals. The crystals raphides (abundant), or raphides and solitary-prismatic. Minor leaf veins without phloem transfer cells (Asphodelus).
Axial (stem, wood) anatomy. Secondary thickening absent, or anomalous (e.g. Aloë). The anomalous secondary thickening when present, from a single cambial ring. The axial xylem with vessels (rarely), or without vessels.
The vessel end-walls scalariform.
Root anatomy. Roots with velamen (in some genera), or without velamen. Root xylem with vessels; vessel end-walls mostly simple.
Reproductive type, pollination. Plants hermaphrodite (usually). Floral nectaries present. Nectar secretion from the gynoecium (via septal nectaries).
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in inflorescences; in racemes, or in spikes. The ultimate inflorescence units racemose. Inflorescences scapiflorous; terminal; simple or compound racemes or spikes; espatheate. Flowers bracteate; small to large; regular to very irregular; when irregular, zygomorphic. The floral irregularity involving the perianth, or involving the perianth and involving the androecium. Flowers 3 merous; cyclic; pentacyclic. Perigone tube present (often, as a long, commonly curved tube), or absent.
Perianth with distinct calyx and corolla, or of tepals; 6; free, or joined; 2 whorled (3+3); isomerous; sepaloid and petaloid, or petaloid; similar in the two whorls, or different in the two whorls; green, or white, or red, or pink, or yellow, or purple, or purple and brown (not blue or violet). Tepal apex trichomes (TAT) absent (11 genera).
Androecium 6. Androecial members free of the perianth; all equal, or markedly unequal; free of one another; 2 whorled (3+3). Androecium exclusively of fertile stamens. Stamens 6; diplostemonous; alterniperianthial. Anthers dorsifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. The endothecial thickenings spiral. Microsporogenesis simultaneous. Pollen shed as single grains. Pollen grains aperturate; 1 aperturate; sulcate; 2-celled.
Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil 3 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 3 locular. Gynoecium stylate. Styles apical. Stigmas 1; dry type (usually), or wet type. Placentation axile. Ovules 2–40 per locule (2 to rather numerous); usually arillate; hemianatropous, or anatropous (nearly orthotropous in Aloë and Asphodelus); bitegmic; crassinucellate. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (Asphodelus, and with filiform apparatus). Endosperm formation helobial.
Fruit non-fleshy (nearly always), or fleshy (Lomatophyllum); dehiscent; a capsule. Capsules loculicidal. Seeds endospermic. Endosperm oily. Seeds winged, or wingless. Embryo well differentiated. Cotyledons 1. Embryo achlorophyllous (1/1); straight. Testa usually encrusted with phytomelan; black (usually), or grey, or brown (variable in Bulbine).
Seedling. Hypocotyl internode present, or absent. Mesocotyl absent. Seedling collar not conspicuous. Cotyledon hyperphyll compact; non-assimilatory. Coleoptile present, or absent. Primary root ephemeral.
Physiology, phytochemistry. CAM. C3 physiology recorded directly in Asphodelus, Bulbine. CAM recorded directly in Aloë, Astroloba, Bulbine, Gasteria, Haworthia, Poellnitzia. Inulin recorded (Gibbs 1974). Cyanogenic, or not cyanogenic. Anthraquinones detected (Aloë, Asphodeline, Asphodelus, Bulbine, Eremurus, Kniphofia, Simethis); polyacetate derived. Saponins/sapogenins absent. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent.
Geography, cytology. Holarctic, Paleotropical, Neotropical, Cape, Australian, and Antarctic. Widespread Old World, clearly centred in southern Africa. X = (6)7.
Taxonomy. Subclass Monocotyledonae. Dahlgren et al. Superorder Liliiflorae; Asparagales. APG 3 core angiosperms; Superorder Lilianae; non-commelinid Monocot; Order Asparagales (as a synonym of Xanthorrhoeaceae-Asphodeloideae).
Species about 800. Genera 20–24; Aloë (including Aloinella Lemée), Guillauminia, Lemeea, Asphodeline, Asphodelus, Astroloba, Bulbine, Bulbinella, Bulbinopsis, Chamaealoë, Chortolirion, Eremurus, Gasteria, Haworthia, Jodrellia, Kniphofia, Lomatophyllum, Paradisea (or Hyacinthaceae, or Anthericaceae?), Poellnitzia, Simethis, Trachyandra.
General remarks. Other than differing tendencies in numerous, often conspicuous overlapping characters, Asphodelaceae as represented by this compiled description differ from the far more coherent Xanthorrhoeaceae sensu stricto (q.v.) only in lack of secretory cavites in leaf laminae, arillate ovules, a compact cotyledon hyperphyll, positive records of inulin and absence of flavonols.
Illustrations. • AloŽ brevifolia: Bot. Reg. 996, 1826. • Habit (AloŽ). • Kniphofia burchellii: as Tritoma, Bot. Reg. 1745, 1836. • Simethis planifolia: as S. bicolor, Eng. Bot. 1541 (1869). • Eremurus, Gasteria, Haworthia, Knophofia (Chittenden).
We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.
Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 19th October 2016. delta-intkey.com’.