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Iridaceae in sub-Saharan Africa
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Published In: Species Plantarum 1: 36. 1753. (1 May 1753) (Sp. Pl.) Name publication detailView in BotanicusView in Biodiversity Heritage Library

Project Name Data (Last Modified On 4/28/2016)
Acceptance : Accepted
Taxon Profile     (Last Modified On 12/17/2016)
Description: Deciduous (rarely evergreen) geophytes. Corm globose to depressed-globose, axial in origin, tunics of fine or coarse fibres, leathery or woody, rarely papery. Catapylls 3, pale and membranous. Stem well developed, usually terete or rarely angled or slightly winged, simple or branched, often flexuouse, rarely scabrid to velvety. Leaves solitary to several, mostly basal and forming a 2-ranked fan or some cauline, rarely borne on a separate shoot, sometimes dry at flowering, blade sometimes reduced or lacking and leaves thus entirely sheathing, unifacial, usually with a definite midrib, plane and linear to lanceolate, rarely round to oval in section, occasionally the margins or midrib raised or winged and leaves thus X- or H-shaped in section, sometimes terete and longitudinally 4-grooved, cauline leaves usually few and reduced, rarely scabrid to velvety or pubescent; margin with vascular bundle and epidermal cells unspecialized. Inflorescence a spike, usually secund or in two ranks; bracts green and soft or firm, sometimes dry above, usually large, inner smaller than outer and 2-toothed or notched apically. Flowers usually zygomorphic, rarely radially symmetric, long-lived, bilabiate and funnel–salver-shaped or rarely rotate, variously coloured but often pink to mauve, lower tepals usually with contrasting nectar guides, unscented or variously scented, often closing at night, with nectar from septal nectaries; perianth tube funnel-shaped or cylindric, short or long; tepals subequal or unequal with dorsal largest and arched to hooded over the stamens, the lower three narrower and forming a lip, sometimes clawed and shortly united. Stamens usually unilateral and arcuate, rarely symmetrically disposed; filaments inserted well below top of perianth tube, included or exserted. Ovary globose; style filiform, usually unilateral and arcuate, 3-branched distally, branches oblanceolate-conduplicate,  and expanded above, often  bilobed. Capsules leathery,  usually slightly inflated and raltively large, ovoid to ellipsoid, segments recurving strongly after dehiscence. Seeds discoid with a broadly circumferetial papery wing, rarely wing obsolete, then subglobose or angled, domed on seeed body and concave on wing. Pollen monosulcate-operculate, operculum 2-banded, exine perforate-scabrate. Basic chromosome number x = 15, other numbers 14, 12, 11.
Etymology: from the Latin, gladiolus, diminutive of gladius, sword, for the sword-shaped leaves.
Revisionary account:
General Notes: Species about 270, mainly sub-Saharan Africa and Madagascar, also Europe and the Middle East, about 169 in southern Africa, centred in the winter-rainfall zone of southwestern Western Cape, with secondary centres of diversity in the eastern African highlands and in Shaba Province of Congo. As in most genera of Iridaceae with a significant representation of species in both summer- and winter-rainfall southern Africa there is minimal overlap among the species from the two rainfall regimes or in those from tropical Africa. Gladiolus species are most common in temperate habitats, mainly grasslands and open savanna in areas of summer rainfall and fynbos and other shrublands in the winter-rainfall region.

Gladiolus is highly variable in both floral and vegetative morphology but most species have characteristic ovoid, leathery, somewhat inflated capsules and distinctive seeds with a broad circumferential wing. The style is also diagnostic, with oblanceolate-conduplicate branches that are expanded above into broad, bilobed tips. The basic chromosome number, x = 15, indicates that the genus is palaeopolyploid. Most species are diploid, 2n = 30 but the almost pan African G. dalenii is heteroploid, with tetraploid, hexaploid, and occasionally diploid populations.

The affinities of Gladiolus have long been uncertain but molecular phylogenetic analyses suggest that the genus is most closely allied to the monospecific Melasphaerula of the southern African winter-rainfall zone. The two genera have little morphological similarity. A molecular phylogenetic genetic analysis of Gladiolus, with an emphasis on the southern Arican and Mediterranean species, proposes an origin for the genus in southern Africa during the Miocene, followed by range extensions into tropical Africa, Madagascar and the Mediterranean. The high diversity in the southern African winter-rainfall region is interpreted as a result of the long history of the genus in the region, combined with comparatively low rates of extinction and higher rates of ecological speciation.

The large number of generic synonyms reflects the floral diversity in the genus, much of which is now understood to represent adaptation to various specialized pollination systems. Pollination by large anthophorine bees foraging for nectar is the most common system in the genus but many southern African species have long-tubed white to pink flowers adapted for pollination by long-proboscid flies. Bird pollination is frequent in southern and tropical African species, as is moth pollination by large settling moths or hovering sphinx moths. Other species are pollinated by large butterflies, notably in southern Africa, where red-flowered species are pollinated largely or exclusively by the satyrid butterfly, Aeropetes. Radial symmetry has evolved in a handful of species in both southern and tropical Africa, and these species appear mostly to be pollinated by female bees foraging for pollen.

The floral diversity is matched by numerous vegetative adaptations, notably reduction in leaf number and the development of winged midribs and/or margins, or reduction of the entire leaf blade. In some species flowering occurs before the leaves are produced or after they have dried, thereby separating the growth and reproductive phases of the life cycle.

The current classification of Gladiolus in southern Africa recognizes seven sections, distinguished by the insertion of the foliage leaves on the stem, the shape and thickening of the blades, the shape and number of the flowers, and the shape and size of the capsules. The molecular phylogenetic analysis of the genus by Valente is only partially congruent with this classification and some marked anomalies make it difficult to interpret. However, many of the series or species groups recognized by Goldblatt & Manning are retrieved in their phylogeny, providing independent support for species relationships at these lower taxonomic levels. Provisionally, therefore, the slightly modified version of Goldblatt & Manning's classification by proposed by Manning & Goldblatt (2009) is followed here.


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1 Plants of tropical Africa (Zimbabwe and Angola to Ethiopia and all of West Africa) Gladiolus group Afro-tropical species
+ Plants of southern Africa arranged by sections (2)
2 (1) Leaves broadly to narrowly lanceolate, usually at least 4 mm wide, often with well-developed secondary veins, these sometimes finely and closely set; flowers mostly slightly bilabiate with the lower tepals not clawed but sometimes strongly clawed (3)
+ Leaves linear to terete, usually less than 5 mm wide, usually without well-developed secondary veins, if broader then without well developed central vein; flowers usually strongly bilabiate with lower tepals narrow and clawed (6)
3 (2) Leaves in a 2-ranked fan, glabrous or puberulous; corm tunics often papery (4)
+ Leaves usually superposed, blades often reduced, sheaths and sometimes blades pubescent or villous, rarely scabird; corm tunics of flattened fibres Gladiolus sect. Linearifolii
4 (3) Flowers usually small; capsules less than 15 mm long; seeds smaller, usually less than 6 mm long, with wings often reduced or unevenly developed Gladiolus sect. Linearifolii
+ Flowers usually larger: capsules usually more than 20 mm long; seeds larger, usually more than 8 mm long, with well-developed wings (5)
5 (4) Flowers rarely white to pink, sometimes red but then lower tepals with white transverse bands or base Gladiolus sect. Ophiolyza
+ Flowers usually cream to pale or deep pink or salmon-orange, lower tepals usually with red spade- diamond- or spear-shaped markings, often with white centre, or median red streak, flowers sometimes red but then with longitudinal markings Gladiolus sect. Blandi
6 (2) Leaves usually several, at least three basal (7)
+ Leaves (2)3(4), one or two basal and others superposed and inserted serially up stem; stems unbranched; spike usually often strongly flexuouse or sometimes stiffly erect; nectar guides on lower tepals usually of fine, longitudinal dark streaks and spots over a pale area or transverse bands of pale colour; capsules usually ellipsoid to ovoid and acute; flowers odten fragrant, except when long-tubed and pink or orange to red, sometimes nocturnal Gladiolus sect. Homoglossum
7 (6) Leaf midrib and margins usually heavily thickened, the blades sometimes centric and cross-shaped in section or terete and with four longitudinal furrows or grooves; corm tunics fibrous, often accumulating in a neck around base of stem; flowers rarely lighty scented; capsules usually oblong and slightly 3-lobed above; seeds often with wings reduced, vestigial or sometimes lacking Gladiolus sect. Heterocolon
+ Leaf midrib lightly thickened and raised but margins usually unthickened, rarely ridged or corrugated; corm tunics membranous, papery, woody or fibrous but rarely accumulating in a neck around base of stem; flowers often highly fragrant except red-flowered species; capsules usually ellipsoid and acute, rarely obovoid and 3-lobed above; seeds broadly and evenly winged Gladiolus sect. Hebea

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