Systematics of the hypervariable Moraea tripetala complex (Iridaceae: Iridoideae) of the southern African winter rainfall zone

Field and laboratory research has shown that the Moraea tripetala complex of western South Africa, traditionally treated as a single species, sometimes with two additional varieties, has a pattern of morphological and cytological variation too complex to be accommodated in a single species. Variation in floral structure, especially the shape of the inner tepals, degree of union of the filaments, anther length and pollen colour form coherent patterns closely correlated with morphology of the corm tunics, mode of vegetative reproduction, and in some instances capsule and seed shape and size. The morphological patterns also correlate with geography, flowering time and sometimes habitat. It is especially significant that different variants of the complex may co-occur, each with overlapping or separate flowering times, a situation that conflicts with a single species taxonomy. We propose recognizing nine species and three additional subspecies for plants currently assigned to M. tripetala . M. grandis , from the western Karoo, has virtually free filaments and leaves often ± plane distally; closely allied M. amabilis , also with ± free filaments and often hairy leaves, is centred in the western Karoo and Olifants River Valley. Its range overlaps that of M. cuspidata , which has narrowly channelled, smooth leaves, linear inner tepals spreading distally and filaments united for up to 1.5 mm. M. decipiens from the Piketberg, M. hainebachiana , a local endemic of coastal limestone fynbos in the Saldanha District, M. ogamana from seasonally wet lowlands, and early flowering M. mutila constitute the remaining species of the complex in the southwestern Western Cape. M. helmei , a local endemic of middle elevations in the Kamiesberg, namaqualand, has small flowers with short, tricuspidate inner tepals. All but M. amabilis and M. mutila are new species. We divide M. tripetala sensu stricto into three subspecies: widespread subsp. tripetala , subsp. violacea from the interior Cape flora region, and late-flowering subsp. jacquiniana from the Cape Peninsula and surrounding mountains.

The Afro-Eurasian and largely sub-Saharan genus Moraea Mill. (Iridaceae: Iridoideae) comprises ± 215 species of cormous geophytes (goldblatt & Manning 2009& Manning , 2010. Although florally diverse, the genus is recognized by a bifacial and channelled (rarely terete) leaf blade (isobilateral leaves are ancestral for Iridoideae) and a corm of a single internode derived from a lateral bud in the axil of the lowermost cataphyll. The majority of Moraea species have Iris-like flowers with clawed tepals, prominent, spreading outer tepal limbs marked with nectar guides, and flattened, petaloid style branches to which the anthers are appressed. unlike Iris L., the tepals of Moraea are normally free and the filaments are at least partially united or rarely secondarily free in a few species (always free in Iris). Species of Moraea with other flower types (goldblatt 1986b; 1998), usually with subequal tepals, reduced style branches and, in some species, a perianth tube, are derived and limit the utility of floral characters in circumscribing the genus.
Relatively widespread in the southern Africa winter-rainfall zone, Moraea tripetala is one of the more common species of subg. Vieusseuxia (D.Delaroche) goldblatt (± 30 spp.). The group is distinguished by the flowers usually lasting three days (vs. fugaceous and lasting less than a single day, the plesiomorphic condition), a single, channelled foliage leaf, and inner tepals variously modified in most species into trilobed or linear structures, occasionally reduced to short hair-like cusps or rarely absent (goldblatt 1976a; 1986a). As currently understood (goldblatt 1976b) M. tripetala is recognized by a blue, violet or purple perianth, reduced inner tepals that are usually represented by short, hair-like cusps 2-3 mm long, filaments either free or shortly united for 1-2 mm (1/4 to 1/3 their entire length) and an associated short style with the style branches appressed to the stamens for almost their entire length. Variation in the colour and texture of the corm tunics, length and shape of the leaves, in the size and shape of the inner tepals, and the occasional development of pubescence on the stems and leaves, was documented in some detail by goldblatt (1976b), who at that stage preferred not to accord any of the variant forms taxonomic recognition. goldblatt (1976b) also noted that M. tripetala was variable cytologically but the three populations sampled were diploid with a basic chromosome number of x = 6, this shared with all species of subgen. Vieusseuxia.
During extensive field work in western southern Africa over the past decade we have encountered a wide range of plants currently identified as Moraea tripetala. The patterns of variation in the flower, especially the form of the inner tepals, and associated differences in corm, capsule and seed morphology, and in the means of asexual reproduction have led us to the conclusion that recognition of a single species for the complex does not reflect biological reality. Particularly difficult to reconcile with a single species treatment for M. tripetala is the co-occurrence of two morphological variants, either flowering virtually side-by-side, or in bloom several weeks or months apart in the same locality, and some-times differing in their habitat. With this in mind we have undertaken an extensive review of the variation in M. tripetala. We conclude that a better treatment of the complex, which we consider monophyletic, is to recognize nine species, with M. tripetala subdivided into three subspecies.

MATERIALS AnD METHODS
We first made field observations across the entire range of the Moraea tripetala complex, noting in addition to floral morphology, details of corm tunics, mode of vegetative reproduction, and whenever possible capsule and seed morphology. These observations were then integrated with an examination of specimens in herbaria with important holdings of southern Africa flora, BOL, K, MO, nBg, PRE, and S (abbreviations following Holmgren et al. 1990), plus a study of type material of the named variants in the complex. Abbreviation of author names follows Brummitt & Powell (1992).
Chromosome counts were determined from mitotic squashes using root tips. Material for the original counts reported here was prepared according to the protocol described by goldblatt & Takei (1993). The vouchers are housed at the Missouri Botanical garden Herbarium (MO). Counts are based on samples of three to four individuals and, following standard practice in plant cytology, are assumed to represent an entire population.

RESuLTS
The lectotype of Moraea tripetala (designated by goldblatt 1976b), is a Thunberg collection from the southwestern Cape, with filaments united for ± 1 mm, inner tepals reduced to linear, acute cusps ± 3 mm long, and a linear, narrowly channelled, glabrous leaf. Corms are lacking in the type material but we are confident that a range of plants recorded from Aurora on the west coast of the Western Cape eastward to Knysna and mainly flowering from August to the end of September accord with the type. These often have slightly sweet-scented flowers and the inner tepals vary from short hair-like vestiges to linear structures up to 5 mm long. Apart from plants matching the type relatively well, there is a range of somewhat to significantly different populations both in the southwestern Cape and in the interior, in the western Karoo and Little Karoo. We describe the variant populations below, dealing first with plants from the near southwestern Cape followed by those from the interior.
Variant 1: the first significant variant in the southwestern Cape is an early flowering taxon, mostly blooming from mid-August to early September, often with relatively broad leaves. These are sometimes plane distally, 3-5(-7) mm wide and laxly twisted, ± half as long to ± as long as the stem, or rarely exceeding it. Plants occur at low elevations on clay or loamy soils. The inner tepals are linear to narrowly lanceolate, up to 12.5 mm long, and differentiated into an erect portion (equivalent to the claw of the outer tepals) and an outspread portion (equivalent to the limb) that is either slightly expanded at the base or, in some populations (e.g. Goldblatt 2310 MO), with two obtuse, lateral lobes, rendering the limb ± trilobed. The filaments are united for 1.0-1.5 mm, rarely only ± 0.5 mm. In two collections the leaves and stem of some plants are thinly hairy (e.g. Nordenstam & Lundgren 1998 MO, nBg, S;Acocks 1912 S). This variant was first recognized at species rank by C.F. Ecklon (1827) as Vieusseuxia mutila and later by J.g. Baker (1904) as Moraea punctata but has remained poorly understood. Moraea monophylla (Baker 1906) and M. tripetala var. mutila (Baker 1896) represent the same variant.
Plants from the immediate area of Sir Lowry's Pass, east of Cape Town, flowering in August, have inner tepals ± 10 mm long, consisting of a limb with a short central tapering cusp and short rounded lateral lobes. They are somewhat out of range for variant 1 and have a narrow, channelled, smooth leaf (e.g. Loubser 872 nBg; Goldblatt 2506 MO). These plants may belong here or alternatively may be hybrids with M. unguiculata or another species: we discuss them below in more detail.
Variant 2: a second southwestern Cape variant is the plant g.J. Lewis (1941) named Moraea tripetala var. jacquiniana (= M. jacquiniana Schltr. ined.). Often shorter in stature, 140-300 mm high and with smaller flowers than in typical M. tripetala, it usually has exceptionally slender leaves, mostly 2-3 mm wide, V-shaped in cross section. When dry the blades are closely folded together and appear terete. Flowering in this plant is from mid-november to January on the Cape Peninsula (Lewis 1950), but sometimes in late October elsewhere. Illustrated in Flora of the Cape Peninsula (Maytham Kidd 1950), the flowers are often dark purple, but sometimes light mauve to ± blue as in M. tripetala sensu stricto, and as far as known, consistently have white nectar guides. The linear inner tepals are 5-6 mm long (e.g. Pillans 10279 BOL, MO), thus slightly more than 1/2 as long as the outer tepal claws and reaching to between the base and middle of the anthers. The outer tepals have limbs 11-16 mm long and claws 8-11 mm long (vs. limbs 12-18 and claws 10-12 mm long in M. tripetala sensu stricto). At several localities, notably in the southern Cape Peninsula, at Jonkershoek and in the grabouw area, both typical M. tripetala and var. jacquiniana occur, the former blooming two to four months earlier, late July to September. M. pulchra Eckl. (1827) from 'Hottentotshollandkloof' collected in flower on 25 november represents the earliest name for this plant at species rank.
Variant 3: restricted to lowland wetlands of the southwestern Cape and flowering in September, this variant has an unusual, narrow ovary, cylindric rather than ovoid, 10-13 mm long, and darkly lined vertically on locules, and nearly cylindric capsules (15-)20-24 mm long, somewhat angled on the locules. Typical M. tripetala has an ellipsoid ovary 6-10 mm long and capsules 8-14 mm long, both round in cross section. The flowers of this variant are also unusual in having the outer tepal limbs pale blue with darker veins radiating from a yellow nectar guide. Plants are always relatively short, rarely exceeding 200 mm, have leaves usually shorter than the stems, and small corms with blackish, wiry tunics. There is also invariably a small cormlet in the axil of the foliage leaf, and plants often grow in small groups, the result of vegetative reproduction from these cormlets. Typical M. tripetala does not normally produce cormlets either in the leaf axils or at the corm base, and is always a solitary plant, favouring well-drained habitats. Variant 3 extends from Voëlvlei in the north to Harmony Flats at Strand in the south.
Variant 4: a last variant in the southwestern Cape is a vegetative apomict, restricted to limestone pavement or calcareous sands in the Saldanha Bay area, and blooming in August and early September, rarely later. Relatively short, up to 300 mm high, plants have lilac or pale to mid-blue flowers with pale yellow nectar guides, outer tepals 27-29 mm long and unusual inner tepals ± 4 mm long, spindle-shaped, sometimes oblanceolate and obscurely 3-lobed, tapering distally with the apex curving inward, sometimes the inner distal surface pilose. unusual for the complex, it has pale grey-blue anthers and off-white or yellow pollen. Capsules never develop to maturity and are shed several days after flowers fade and seeds are not formed. Microscopic examination of pollen confirms that the grains are all malformed. Propagation of new plants is accomplished by the production of several cormlets born in the axil of the foliage leaf and sometimes the lowermost sheathing leaf. non-flowering individuals produce similar cormlets at the base of the main corm. This variant co-occurs with typical M. tripetala, which blooms in the same sites in late September and early October, and has paler blue flowers and dark blue or purple anthers and red pollen typical of the species.
Variant 5: some plants from the Piketberg assigned to Moraea tripetala by goldblatt (1976b) have large inner tepals 7-10 mm long with distally expanded limbs that are ± trilobed and tapering to an attenuate, twisted tip. These plants are also unusually tall, reaching up to 450 mm, but the flowers are relatively small for the complex, with outer tepals 20-23 mm long and claws 8-9 mm long. The filaments, only ± 4 mm long, are united for ± 2 mm. In general appearance these plants resemble M. unguiculata Ker gawl., but the filaments, united for less than 1/2 (slightly more than 1/3) their length, are characteristic of the M. tripetala complex. Both typical M. tripetala and white-flowered M. unguiculata occur locally in the Piketberg, where the former has the short, hair-like inner tepals typical of the species.
Variant 6: outside the immediate southwestern Cape, in the western Karoo, plants included in Moraea tripetala usually have free filaments (sometimes united basally for < 0.4 mm), inner tepals reduced to small hair-like cusps mostly 1-2 mm long, a linear leaf, shallowly channelled below but often plane distally, and light brown corm tunics of medium-textured to soft fibres. Plants always form clonal colonies, the result of vegetative propagation by the production of two or more cormlets at the base of the flowering stem that replace the parent corm, and/or by axillary cormlets (Figure 14).
Populations of this variant fall into two main groups. The first of these, from the western Karoo and the Olifants River Valley and surrounding mountains, mostly from lighter, sandstone or shale-derived soils, has smaller flowers with outer tepals 23-25 mm long (claws 10-14 mm), anthers 4.5-7.0(-8.0) mm long and ± as long as to 1.5 × as long as the filaments, and either white to yellow pollen (eastern populations) or red pollen (western populations) ( Figure 14). Pollen colour is lost in older collections but our observations are based on examination of living plants across almost the entire range of this morph. The 5(6)-sided seeds are 1.7-2.0 × ± 1 mm and stand out in having pale, spongy, ridged angles and an extended pale spongy mass at the micropylar end. Seeds of typical M. tripetala and related variants are 1.0-1.3 × ± 1 mm, with sharply raised, pale angles and a finely wrinkled surface through which the underlying dark brown seed body is evident. This variant matches the type of M. amabilis Diels.
Plants closely resembling this variant growing on sandstone pavement in the Bokkeveld and gifberg-Matsikamma Mtns are often particularly short, usually < 180 mm high. Capsules and seeds in these plants are consistent with this variant.
A second western Karoo variant comprises large-flowered plants, mostly of dolerite-derived soils, with outer tepals 30-40 mm long (claws 12-18 mm long), anthers 8-11 mm long and ± twice as long as the filaments, and consistently red pollen ( Figure 15). The capsules are 16-19 mm long, narrowly ellipsoid-oblong with a markedly thickened apical rim. Seeds in the few fruiting collections that we have been able to examine have exceptionally large, brown seeds, 2-3 × 1.8-2.3 mm.
Variant 7: also in the western Karoo and adjacent interior southern Cape, another variant of the complex has linear inner tepals (8-)10-15 mm long, ascending below and spreading distally, filaments united for 0.5-1.8 mm, and narrowly channelled leaves with the leaf halves often appressed. The relatively large corms have tunics of wiry, usually thickened fibres, and cataphylls that decay into a particularly prominent collar of vertical fibres around the base. The filaments are (3-)4-6 mm long and the anthers, 4-8 mm long, with red pollen. Late flowering, these populations seldom bloom before the last week of September and only in late October at higher elevations in the Swartberg Mtns. Plants are sympatric in the Roggeveld and Klein Roggeveld with the western Karoo morph of variant 6, which has short, hair-like inner tepals and pale corm tunics, and their flowering times often overlap (e.g. Goldblatt & Porter 13461 and 13462, MO, nBg, from the Farms Fortuin and nuwerus). The few plants we have seen with mature capsules have distinctive seeds with a spongy testa slightly thicker on the angles.
Variant 8: plants from the Kamiesberg, namaqualand, discovered only in 2009 by Cape Town botanist n.A. Helme, and evidently belonging to the Moraea tripetala complex, have filaments united in the lower 1.5 mm but differ notably in their short, trifid inner tepals, represent the last significant variant of the complex (Figure 4). Apart from the unique inner tepals, the flowers are small, with outer tepals 21.0-23.5 mm long bearing bright yellow, velvety nectar guides. The outer tepal claws have a pair of marginal teeth just below their apices.
Even with the exclusion of these variants, the remaining populations of Moraea tripetala are still variable, and we discuss the major patterns below under that species. Extending from Aurora and the Piketberg on the west coast to the Cape Peninsula and thence across the southern Cape to Knysna, plants have moderately-sized, pale blue, purple or violet flowers with white or yellow nectar guides (many populations are variable for flower colour), inner tepals hair-like and rarely > 4 mm long, and the ovary is ellipsoid and 8-10 mm long. Vegetative reproduction via axillary or basal cormlets is rare. Among these populations, plants from the southern Cape sometimes lack inner tepals entirely and those from the Langeberg foothills may have free filaments. neither trend is consistent as far as we can determine. From the few fruiting specimens available, seeds of what we consider typical M. tripetala are consistently small, with pale, raised, ± winged angles.

DISCuSSIOn
Species, subspecies or merely ecotypes?: the immediate question our observations raise is whether or not any of the several morphological variants should be recognized taxonomically, thereby reducing the otherwise florally distinctive Moraea tripetala to just one of several taxa that are only moderately distinctive florally and not easily distinguished without examination of corms, capsules, and seeds. We conclude that the patterns of variation that we now recognize renders the single species solution unacceptable, particularly because of the correlated character differences in the corm, leaf, flower, and seeds, and the examples of sympatry of two variants at some sites, either flowering at the same time or flowering weeks to months apart. Moreover, the range of habitats and flowering times would be remarkable for a single species and unacceptable for the lack of conformity with any but the broadest morphological species concepts. Similarly, treatment of the major variants as subspecies violates any biological species concept. We conclude that all the distinctive entities that co-occur and have overlapping flowering times should be treated as separate species. Likewise, co-occurring variants that have flowering times separated by weeks or months require taxonomic recognition if they can be readily identified by more than one unique morphological marker. This framework has been the underlying philosophy in our decision to dismember Moraea tripetala sensu goldblatt (1976b). We also note that Baker (1896) in Flora capensis recognized one variety in the species and later added two more (Baker 1904(Baker , 1906. Lewis (1950)  Floral and reproductive biology: as might be expected in species with similar flowers, all members of the complex share the same pollination system, large bodied, mostly anthophorine bees (usually Amegilla and Anthophora spp.), or sometimes Apis mellifera (goldblatt et al. 2005). Our impression is that the species of the complex are remarkably successful and it is rare to see plants later in the season without multiple capsules with full complements of seeds; autogamous selfing can be dismissed because in these iris-type flowers the anthers do not reach the stigmatic lobes (goldblatt 1998) and we infer that insect mediated pollen transfer is necessary to achieve pollination.
Propagation by seed is complemented in several species either by production of cormlets in the leaf or cataphyll axils and at the base of the corm (Moraea hainebachiana, M. ogamana) or sometimes by production of two new corms in place of the parent corm (Moraea amabilis and often M. grandis). Enhanced vegetative reproduction in these species results in plants forming clonal colonies. These two vegetative propagation strategies are uncommon elsewhere in the M. tripetala complex. The lowland wetland populations we treat as M. ogamana consistently produce a single cormlet in the foliage leaf axil but the vegetative apomict, M. hainebachiana produces multiple cormlets in the foliage leaf axil as well as at the corm base, a feature particularly well-developed in non-flowering individuals. This last species produces malformed pollen, and the capsules are invariably shed soon after flowers wilt, without the ovules developing into seeds.
Chromosome cytology: as noted by goldblatt (1976a), chromosomal karyotypes are variable for populations then included in Moraea tripetala. Sampling is limited (Table 1) but we note the following. M. amabilis, M. cuspidate, and M. grandis have the longest three chromosome pairs either metacentric or sub-metacentric ( Figures 1A, B) and a small satellite on the fourth longest ± acrocentric pair (goldblatt 1971 and Table 1, as sat type 1). This karyotype is the common one in subgen. Vieusseuxia. The karyotype of M. mutila has small satellites on the two longest, submetacentric chromosome pairs (sat type 3) ( Figure 1C). In contrast, plants of M. tripetala subsp. tripetala and subsp. violacea (Table 1), have very large satellites on a long, telocentric chromosome pair (sat type 2). In three of four populations of subsp. tripetala sampled a second large satellite ( Figure  1D) is evident on a relatively short acrocentric chromosome pair (sat type 2a) (poor preparation in the fourth sample makes it uncertain whether the second pair of satellites is present or not). These examples, inadequate though they are, show that chromosomal rearrangements via inversion or translocation occur with some frequency in the complex and potentially provide a genetic basis for reproductive isolation because of meiotic disruption in any inter-populational hybrids. The three karyotypes also provide support of our revised taxonomy in showing different karyotypes, consistent within species. plants usually forming colonies due to vegetative reproduction by axillary cormlets and/or two new cormlets replacing parent corm; leaves widely channeled, often ± plane distally; plants sometimes velvety on stems and abaxial surface of leaves: 8a Flowers with outer tepal limbs (9-)11-18 × 10-14 mm, always as long as or longer than wide; anthers 4.5-7.0(-8.0) mm long and up to one and a half times as long as filaments; corm tunics usually of hard, coarse, light brown (  mm diam., exceeding stem and often trailing above, sometimes with axillary cormlet; sheathing cauline leaves 45-55 mm long, green with dry attenuate apices. Rhipidia mostly 3-5-flowered; spathes green with dry, brown, attenuate tips, inner 35-40 mm long, outer ± half as long as inner. Flowers pale to deep purple, outer tepal limb bases with white, wedge-or V-shaped, minutely papillate nectar guides dark purple in centre; outer tepals 20-23 mm long, claw 8-9 mm long, with wide, dark violet median stripe, limb 12-15 × 6-8 mm; inner tepals 7-10 long, violet, claw suberect, plane, expanded to 2 mm wide at apex, often into rounded lobes, limb 3-4 mm long, tapering to attenuate tip, laxly twisted distally. Stamens with filaments 3.5-4.0 mm long, united in lower ± 2 mm; anthers 4.5-6.0 mm long, reaching or shortly exceeding stigma lobe, dark purple-black; pollen orange-red. Ovary narrowly oblong-truncate, 7-9 mm long, usually exserted when flower mature; style ± 2 mm long, dividing ± 0.5 mm above top of filament column, branches, 7-9 × 1-2 mm; stigma shallowly bilobed; style crests narrowly wedge-shaped, ± 5 mm long, erect. Capsules narrowly ovoid, 8-11 mm long. Seeds 5(6)sided, 1.2-1.5 mm × ± 1 mm, facets slightly wrinkled, brown seed body visible though testa, angles between facets forming narrow, raised, pale, slightly spongy ridges. Chromosome number unknown. Flowering time: late Oct.-mid-nov. Figure 2.
Distribution: known only from the western half of the Piketberg in Western Cape (Figure 3), Moraea decipiens is evidently restricted to middle elevations of the range, rather than the emergent peaks, and based on our own collection, occurs on stony sandstone slopes in shallow pockets of soil among Restionaceae and low shrubs. none of the other three collections that we have seen include information about the habitat.
Diagnosis: named for the deceptive (decipiens = Latin deceiving) appearance of the flowers to those of M. tripetala, M. decipiens, nevertheless, differs in having the inner tepal claws distinctly expanded distally, sometimes into rounded lobes, with the limb linearattenuate and curved inward above. The inner tepals recall those of M. unguiculata and M. algoensis but unlike those species, the filaments of M. decipiens are only shortly united for ± 2 mm and the style branches diverge above the fused part of the filaments whereas in M. unguiculata and M. algoensis the filaments are united almost to their apices in a relatively thick column. The style branches of M. decipiens are relatively short, ± 7 mm long, unlike those in M. tripetala which are usually 9-12 mm long. The markings on the outer tepals are also notable, consisting of a wedge-or V-shaped white zone surrounding a dark purple basal mark that is continuous as a wide, longitudinal purple streak on the claw. Plants from Pakhuis Pass, flowering in late October (Helme 5726, nBg) bear a superficial resemblance to M. decipiens in size of the flowers, colour of the nectar guides and the broad inner tepals but the filaments are united for 2.5 mm and 6.5 mm long, thus longer than the anthers, which are just 4 mm long. More material of this plant is needed before an informed decision can be made about its status but we suspect it represents yet one more undescribed species of subgen. Vieusseuxia.
Typical Moraea tripetala itself also occurs at higher elevations on the Piketberg, e.g. Linder 414 BOL, from Levant Mtn where M. decipiens also occurs, but flowers some two to four weeks later. Early flowering M. mutila of the complex occurs in renosterveld on the lower eastern slopes of the range and has inner tepals broadly similar to those of M. decipiens, although narrower and longer. The population at the type locality was relatively large and plants showed no variation in critical features, thus unlikely to be hybrid between M. tripetala and M. mutila. Barker's collection of M. decipiens likewise comprises several plants (Barker 7563), all uniform for flower features, and the pollen appears normal under the microscope.  short branches, glabrous. Foliage leaf solitary, linear, channelled, remaining ± erect, ± 2 mm wide when opened flat; sheathing cauline leaves 2 or 3, 35-45 mm long, green, attenuate, becoming dry at tips. Rhipidia 2(3)-flowered; spathes green with dry tips, attenuate, inner 40-45 mm long, outer ± two thirds as long, entirely sheathing. Flowers pale blue to violet, outer tepal limbs darkly veined, with narrowly triangular, shortly velvety, yellow nectar guides at base of outer tepal limbs outlined pale blue to dark violet; outer tepals 21.0-23.5 mm long, claw 8-10 mm long, with a shallow raised tooth either side just below apex, limb 12.5-13.5 × 7-8 mm; inner tepals short, ± 4 mm long, 3-forked in upper 1 mm, inner lobe slightly larger than laterals, ± puberulous. Stamens with filaments ± 4 mm long, united in lower 1.5 mm, column puberulous; anthers ± 6.5 mm long, dark violet; pollen yellow. Ovary narrowly ellipsoid, ± 7 mm long, initially partly included, ultimately fully exserted; style ± 2 mm long, dividing at top of filament column, branches linear, 9 × 1 mm; style crests ± 7 mm long, erect. Capsules and seeds unknown. Flowering time: late Oct.-mid-nov. Distribution and habitat: known only from the Kamiesberg in central namaqualand (Figure 3), Moraea helmei has been collected just twice, from a "somewhat marshy site the top of the Langkloof south of the Farm Karas" and "from a seasonal seep south of Karas at the upper end of the Langkloof". Plants occur at an altitude of about 1 125 m. Conservation status is impossible to assess from the limited data available about the species, but the species is undoubtedly rare.
Diagnosis and relationships: the filaments, united for 1.5 mm and slightly more than one third their length, and the short style, ± 2 mm long, are consistent with the M. tripetala complex but the inner tepals of M. helmei are short and expanded distally into three short, subequal lobes (vs. linear and usually cusp-like in M. tripetala). The puberulous filament column appears to be unique in the complex. The trilobed inner tepals of M. helmei recall those of M. unguiculata, which has a longer, narrow, coiled central lobe and rounded lateral lobes, but the filaments of this species are united in a prominent column for more than half their length. A peculiarity of the outer tepal claws of M. helmei is the pair of shallow tooth-like projections just below the apices, a feature not known elsewhere in Moraea: the tepal claws in all other  species are straight or slightly bowed outward and have plane margins. Plants 15-30 cm high. Corm mostly 10-15 mm diam., evidently without cormlets at base, tunics of medium to coarse fibres, vertical members often thickened below into prominent claws. Stem simple, rarely 1-branched (1 of 28 plants seen), glabrous or occasionally pilose, sheathed below by brown, fibrous cataphylls, these often accumulating. Foliage leaf solitary, linear, shallowly channeled below, often becoming flat and slightly twisted in distal third, ± half to as long as stem, 3-5(-7) mm wide, glabrous or sometimes pilose abaxially or on margins; sheathing cauline leaves 20-40 mm long. Rhipidia mostly 2-or 3-flowered; spathes green, inner 35-40(-45) mm long, outer ± half as long as inner, rarely pilose. Flowers pale blue or white, nectar guides triangular, white to pale yellow with dark blue dots and often edged with darker color, velvety (or smooth?); outer tepal claws 9-12 mm long, limbs dipping at 45°, ovate, widest in distal third, 14-15 × ± 10 mm, margins recurving later, inner tepals ± linear to slightly expanded in middle, sometimes as rounded lobes, 7.5-12.5 mm long. Stamens with filaments 3.0-3.5 mm long, united for 0.5-1.5 mm; anthers 4.5-6.5(-7.5) mm long, pollen red or yellow. Ovary narrowly ellipsoid, 7-8 mm long; style 1-2 mm long, style branches 7-9 mm long, crests linear, 8-10 mm long. Capsules ellipsoid, 15-17 × ± 5 mm. Seeds ± 1.8 × 1.3-1.4 mm, pale straw-coloured, testa spongy, angles narrowed into wings. Chromosome number 2n = 12 (Table1). Flowering time: late Aug.-late Sept., depending on subspecies. Figure 5.
Distribution: largely a species of renosterveld, Moraea mutila is centred in the western lowlands of Western Cape, extending from the Cape Peninsula to Piketberg, and extending locally inland to the Tulbagh Valley (Figure 3). A collection said to be from the Olifants River Valley near Clanwilliam made by Arnold Penther is more likely mislabelled and was perhaps gathered en route from Porterville to Clanwilliam. At the very least the Olifants River Valley record requires confirmation. Soils on which M. mutila has been collected are described as shale and clay, consistent with renosterveld but probably also on granites, and plants we have seen grew on loamy clay. M. mutila is evidently rare on the Cape Peninsula and as far as we have been able to determine was first collected there by C.H. Bergius in 1817, then by Ecklon and by Zeyher in the 1820s, and later by Friedrich Wilms, Rudolf Marloth, and Harry Bolus but there are no recent records. nevertheless, it probably persists on clay soils, its preferred habitat, on the lower slopes of Lions Head and Signal Hill above Cape Town.
Possibly belonging here are pale blue-flowered plants from the western lower slopes of Sir Lowry's Pass (Loubser 872 nBg, Goldblatt 2506 MO) with the inner tepals trilobed as in M. mutila but the filaments are united for ± 2.5 mm and the style is 4 mm long. Alternatively these plants may be hybrids between M. tripetala and M. unguiculata.
Diagnosis: early blooming Moraea mutila (the name means mutilated, presumably due to the reduced inner tepals) is recognized by the pale blue, rarely white flowers with linear to narrowly lanceolate inner tepals 7.5-12.5 mm long, erect below and spreading distally. Often difficult to see in preserved specimens, the inner tepals are usually slightly wider in the midline and sometimes expanded into rounded lobes near the base of the spreading distal half. The flowers otherwise seem little different from those of M. tripetala but the pilose nectar guides are usually white or palest yellow and dotted with dark blue to purple, whereas the nectar guides of M. tripetala are more often yellow, edged with dark blue or purple. The leaf is typically relatively short, in most collections shorter than the stem, and widely channelled, sometimes plane in the distal half. Several specimens of two collections (Acocks 1912 andNordenstam &Lundgren 1998) have leaves velvety on the abaxial surface and velvety spathes. The few collections from the Cape Peninsula also have partly hairy leaves, either on the abaxial surface or on the margins. The pubescence is evidently variable in the populations that show the character, thus of limited value for identification, although we have not seen any collections of true M. tripetala with pubescence of any kind on the vegetative parts.
Taxonomic history: the earliest record of Moraea mutila is the plant described in detail by C.P. Thunberg (Dec. 1782) under the name Iris tripetala. Thunberg's plant had smooth, channelled leaves, a blue flower with yellowish nectar guides, and notably, inner tepals with very narrow claws, convex abaxially, and patent limbs at the base of which were two opposed lobes (dentibus duobus). Thunberg's M. tripetala has, however, no nomenclatural status or is at best superfluous as he cited I. tripetala L.f. (April 1782) in his account. The latter, described incompletely but said to have subulate (awlshaped or needle-like) inner tepals, is the basionym for M. tripetala. Thunberg actually collected multiple specimens of the M. tripetala complex from at least three sites but clearly had M. mutila in mind in his extended description of I. tripetala. Since his description conflicts with that of Linnaeus fil. we treat Thunberg's I. tripetala as a separate, but superfluous name.  September 1826September . nordenstam (1972 accepted the name as valid despite the brief diagnosis that mentions neither the leaf vestiture nor the form of the inner tepals, features critical to identification of the species. The epithet 'mutila' was taken from the manuscript name on Bergius's collection, now in the Stockholm Herbarium and annotated 'Iris nov. species mutila mihi' in Bergius's hand. We wonder whether Ecklon actually saw the Bergius specimen, which was probably already in Berlin when he arrived in Cape   Plants solitary, 180-300 mm high. Corm 15-25 mm diam., without cormlets at base, tunics of brown to almost black, tough, medium to coarse fibres. Stem smooth, simple or 1(2)-branched exceptionally with up to 6 branches, with cataphylls of medium textured, light brown, mostly vertical fibres forming a collar around base, sometimes accumulating in a dense mass. Foliage leaf solitary, linear(-linear-filiform), channelled, 1.5-3.0 mm wide, glabrous, V-shaped in section, leaf halves closely appressed in dry conditions; sheathing cauline leaves (30-)40-55 mm long, with dry attenuate tips. Rhipidia several-flowered; spathes green with attenuate, dry tips, inner (40-)50-80 mm long, outer ± half as long as inner, glabrous. Flowers pale blue, mauve or violet blue, outer tepal limbs lanceolate, with large white, velvety nectar guides dotted and usually edged with dark violet; outer tepal claws (8-)10-12 mm, usually speckled with dark blue to violet dots or purple in midline, limbs (12-)16-22 × 10-16 mm, inner tepals linear, (8-)10-15 mm long, ascending below, distal third to half usually spreading horizontally (or ± erect when short), often pilose at base. Stamens with filaments (3-)4-6 mm long, united basally for 0.5-1.8 mm; anthers 4-8 mm long, pollen red (rarely white). Ovary exserted, 11-15 mm long; style vestigial, branches 8-10 mm long; crests 5-8 mm long, linear, arching inward. Capsules ovoidellipsoid, 14-16 mm long. Seeds 1.8-2.0 × 1.3-1.4 mm, pale straw-coloured, testa spongy, thicker on angles. Chromosome number 2n = 12 (Table 1) Flowering time: mid-Sept. and Oct. Figure 6.

Moraea mutila appears again as
Distribution: a species of semi-arid habitats, Moraea cuspidata grows in both mountain renosterveld and dry, marginal fynbos, usually in sandy or sandy loam soils derived from sandstones of the Cape System or from the Beaufort Series of the Karoo System. It flowers late in the season-the earliest record is in mid-September but mid-to late October is usual. It is relatively poorly collected, not (we suspect) because it is rare, but because it grows in areas not much botanized at that time of year. In the late spring of 2009 we found the species at several sites we visited between Touw's River and Komsberg Pass in the Klein Roggeveld. The recorded range ( Figure  3) extends from the Bonteberg and Voetpadsberg near Touw's River through the Klein Roggeveld and southern Roggeveld to the Swartberg and higher mountains of the Karoo. An outlying record from Perdekloof, southeast of Oudtshoorn near Camfer (Goldblatt & Porter 12575), probably belongs here, although the flowers are exceptionally small, the inner tepals are 8-9 mm long, consistent with M. cuspidata. The apparent gap in the range between the Roggeveld and the Swartberg may be an artifact due to incomplete collecting.
Diagnosis: recognized immediately by the linear inner tepals up to 15 mm long (rarely less than 10 mm), spreading in the distal half ( Figure 6B), Moraea cuspidata (named for the long, cusp-like inner tepals) also usually has a particularly narrow foliage leaf, the leaf halves tightly appressed in dry conditions, and brown, sometimes almost black corm tunics, the vertical fibres of which are usually heavily thickened. In addition, the cataphylls tend to persist as a collar of stiff fibres around the base (not always present in herbarium material). The flowers are pale blue to violet with the large, velvety nectar guides spotted with dark blue on a white background. The filaments are usually united for 1.0-1.5 mm but occasionally less, and ± as long as or slightly shorter than the dark purple anthers that bear red pollen. We have seen only two collections with ripe capsules and these are relatively large, 14-16 mm long. The seeds of these two collections are also relatively large for the complex, 1.8-2.0 mm long and unique in having a pale, spongy testa much thickened into prominent ridges on the angles.
The species is sympatric at some sites and even coblooming with the smaller-flowered Moraea amabilis (also of the M. tripetala complex), but this species has a broadly channeled foliage leaf, sometimes plane distally and often pilose on the abaxial surface, and consistently short, hair-like inner tepals typically 1.5-2.0 mm long. At sites where we have seen the two growing together, anthers of M. amabilis have white pollen, contrasting with the orange-red pollen of M. cuspidata. The range of M. cuspidata also coincides in part with M. tripetala subsp. violacea, which extends from gydo Pass to the Hex River Valley and Touw's River, but we have not seen them growing near one another.  Plants (140-)250-450 mm high, glabrous on all vegetative parts. Corm mostly 8-15 mm diam., without cormlets at base, tunics of moderately coarse, wiry, usually dark grey fibres. Stem simple or 1-or 2(-6)-branched, sheathed below by brown cataphylls and sometimes with collar of brown fibres. Foliage leaf solitary (rarely 2 in subsp. violacea), leathery, narrowly channeled, C-or V-shaped in section, to 5 mm wide, usually exceeding stem and distally trailing; sheathing cauline leaves mostly 45-60 mm long. Rhipidia 3-several-flowered; spathes green with dry attenuate tips, inner 35-65(-70) mm long, outer ± half as long as inner. Flowers pale blue, purple, or violet, nectar guides triangular, yellow edged with violet, or white dotted with dark blue and edged with darker blue, lightly honey scented, unscented or strongly scented of carnation (subsp. violacea); outer tepals 20-30(-32) mm long, claws 9-12 mm long, limbs obovate, widest in distal third, 11-18 × ± 8-13 mm, plane, spreading at 45°, margins eventually recurved, inner tepals hair-like to ± linear, 1-4(-6) mm long, or lacking. Stamens with filaments 4-6 mm long, united for 0.5-1.5 mm, occasionally (southern populations) ± free; anthers 4-7 long, pollen usually orange-red, rarely white. Ovary 4-9 mm long; style 1-2 mm long, style branches 9-12 mm long, crests ± linear, 5-12 mm long. Capsules ellipsoid, 8-14 mm long. Seeds 1.0-1.3 mm × ± 1 mm, flat surfaces slightly wrinkled, brown seed body visible through testa, angles forming raised, golden-brown ridges. Chromosome number 2n = 12 (only subspp. tripetala and violacea counted) (Table 1). Flowering time: mainly occasionally in Oct. Figures 7,8. Distribution: widespread in Western Cape, Moraea tripetala extends from Aurora on the west coast to near Knysna in the east, and inland to Ceres and the Little Karoo ( Figure 9). Habitats vary from limestone flats in fynbos, to neutral and acid sands in sandveld, to loamy alluvium in coastal and montane fynbos. Interior populations from the Warm Bokkeveld (Ceres) and western Little Karoo, here segregated as subsp. violacea, occur in significantly drier habitats on clay soils in renosterveld. Populations segregated as subsp. jacquiniana are mostly montane and occur above 500 m but at lower elevations in the southern Cape Peninsula, and flower significantly later in the year. Although much of its lowland habitat has been lost to agriculture and its original range significantly reduced, M. tripetala still occurs extensively in undisturbed low to middle elevation sites. Interior populations have lost little of their original habitat except immediately around Ceres where orchards have replaced much of the original vegetation.
Diagnosis: in our revised, narrower circumscription, Moraea tripetala includes plants with reduced hair-like to linear inner tepals mostly 2-4, rarely up to 6 mm long, and filaments united for up to 1.5 mm and occasionally free. The outer tepal limbs are broadly ovate,  longer than wide and oriented ± 45° below horizontal. The corms have dark brown to grey tunics, usually composed of medium-textured fibres, and do not produce cormlets at the base. The linear leaves, normally exceeding the stem, are narrowly channeled, and in dry conditions the leaf halves lie parallel to one another. As far as we can determine, vegetative reproduction by production of cormlets does not occur in M. tripetala as now circumscribed. Flower colour ranges from pale to deep blue or violet or rarely light purple, sometimes even in the same population. nectar guides are often yellow edged with dark blue or purple, but occasionally white, then dotted with blue to purple. The seeds are small, and typical of the complex in being 5(6)-sided with the fac-ets slightly wrinkled and showing the brown color of the seed body ( Figure 7D). The angles are raised into narrow wings of pale straw colour. In contrast, M. amabilis, M. cuspidate, and M. grandis have larger seeds of more complex structure.
Variation: there are a number of notable variants among the plants we include in Moraea tripetala, one of the most important of which includes populations from the interior Western Cape on clays and shales from Ceres to the western Little Karoo. These plants have dark violet outer tepals with yellow nectar guides, are usually relatively short in stature, and have short stiff, hair-like inner tepals, mostly ± 2 mm long. The flowers we have examined alive have a strong clove or carnation-like scent. unusually for the complex, and for subg. Vieusseuxia as a whole, some populations of this morph have two foliage leaves (e.g. Dymond s.n. from near Ouberg Pass; Goldblatt 4184, 11438 from Op-de-Tradouw near Barrydale; Van Wyk 61 from the Voetpadsberg; Goldblatt & Snijman 6976A from Worcester). We recognize these populations of the interior Western Cape as subsp. violacea.
Elsewhere, Moraea tripetala has pale or deep blue, or sometimes purple outer tepals, either with yellow or white nectar guides, in the latter case usually speckled with dark blue or purple, and flowers are faintly sweetscented or odourless. It is also relatively tall (allowing for reduced stature in years of poor rainfall) and usually has hair-like to linear inner tepals 2-4(-6) mm long (occasionally the inner tepals are absent in southern Cape populations). Among these populations, those that flower from november to January on the Cape Peninsula and surrounding mountains stand out in their softly fibrous corm tunics, slightly smaller flowers, and usually very narrow leaves. The flowers also often have inner tepals 4-6 mm long. Typical M. tripetala in the same area flowers from August to early October. We think it useful to recognize these two sets of populations taxonomically but favour subspecies rank because the morphological differences between them and typical M. tripetala are small and not absolute. The taxon was treated by Lewis (1941Lewis ( , 1950 as var. jacquiniana.
Variation elsewhere across the wide range of Moraea tripetala seems to us less taxonomically significant and less consistent. We discuss this under subsp. tripetala.
History: although the type of Moraea tripetala was collected by C.P. Thunberg, the basionym Iris tripetala was described by Linnaeus fil. (April, 1782), several months before Thunberg (Dec. 1782) published his own, much more extensive description. goldblatt (1976b) selected as lectotype a specimen that had short, hair-like inner tepals and a long, narrow, channelled leaf, thus fixing the application of the name. Thunberg's plant is obviously a different species, and has the distinctive inner tepals of what is now M. mutila (see extended discussion under that species). The reduced inner tepals rendered M. tripetala so distinctive that it acquired few synonyms. De Candolle (1804), when transferring I. tripetala to Vieusseuxia, the genus then used for southern African species originally treated as Iris, renamed it V. tripetaloides, a morphologically more apt epithet. The name is superfluous as De Candolle cited I. tripetala L.f.  Plants 250-450 mm high, stem sometimes sheathed below by collar of brown fibres. Corm mostly 12-15 mm diam. Foliage leaf always solitary, linear. Inner spathes 45-65(-70) mm long. Flowers pale blue or purple with yellow or white nectar guides, faintly honeyscented or unscented; outer tepals 23-30(-32) mm long, claws 10-12 mm long, limbs 12-18 mm long,; inner tepals usually hair-like, 1-3(-5) mm long, or absent. Stamen filaments 4-6 mm long, united for 1.0-1.5 mm, occasionally (southern populations) ± free; anthers 4.5-7.0 long; pollen usually red or white. Ovary 8-9 mm long; style branches ± 12 mm long, crests mostly 8-12 mm. Capsules 12-14 mm long. Flowering time: late Aug.-mid-Sept., occasionally in Oct. (exceptionally July, e.g. Wurts 212). Figure 7.
Distribution: largely a plant of coastal forelands, subsp. tripetala extends from Aurora on the west coast of Western Cape to near Knysna in the east. It also occurs on lower mountain slopes, occasionally up to ± 700 m, and has been recorded on the Piketberg, the Hottentots Holland-Jonkershoek mountain complex, the Witzenberg at Ceres, and the lower southern slopes of the Langeberg (Figure 9). Populations along the southern Cape, notably in the De Hoop area, the Agulhas Peninsula (Goldblatt 2607), and some from near Caledon (Goldblatt 6196) occasionally lack inner tepals, or individual flowers may lack one or two of the inner tepals (e.g. Goldblatt & Manning 12248, east of Swellendam). Plants from the southern foothills and lower slopes of the Langeberg sometimes have free filaments (e.g. Thorne s.n. SAM, from garcia's Pass; Goldblatt 3741 from near Suurbraak). Subsp. tripetala is usually found in sandy habitats or on coastal limestone, thus in sandveld, strandveld and fynbos vegetation.
Distribution: subsp. violacea extends from gydo Pass and the Warm Bokkeveld though the southern foothills of the Hex River Mtns near Worcester inland to the Bonteberg, Touws River and the western Little Karoo near Barrydale (Figure 9). Populations with two foliage leaves appear to occur randomly across its range. Plants almost invariably occur on clay soils in renosterveld but we have also seen the species growing among low succulent shrubs. Plants 140-300 mm high, stem usually sheathed below by collar of fine, loosely netted fibres. Corms ± 10 mm diam., tunics of relatively fine fibres, not usually accumulating. Foliage leaf solitary, narrowly channelled or leaf halves appressed together thus appearing terete, exceeding stem and up to twice as long, 2-3(-5) mm wide (when dry often apparently ± 1.5 mm diam.). Inner spathes 40-50 mm long. Flowers usually purple or blue with triangular, white nectar guides spotted with dark blue and edged darker color, unscented, outer tepals 20-25 mm long, claws 9-11 mm long, limbs lanceolate, 11-16 × 8-11 mm, inner tepals hair-like, mostly 5-6 mm long, reaching or exceeding anther bases. Stamen filaments 4-6 mm long, united for 1.0-1.5 mm; anthers 5-6 long, pollen usually (always?) red. Ovary 4-6 mm long; style branches 10-11 × 2 mm long, crests linear, 5-8 mm long. Capsules mostly 10-14 mm long. Flowering time: mid-nov.-Jan., rarely in late Oct. Figure 8.

Representative specimens
Distribution: occurring entirely within the range of subsp. tripetala, subsp. jacquiniana extends from the Cape Peninsula to the Hottentots Holland and Houw Hoek Mtns and north to the mountains around Franschhoek, Paarl, and Wellington (Figure 9). There are also isolated records from the Piketberg, the mountains near Ceres, and in the vicinity of Citrusdal (Ecklon & Zey-her Irid 20), this last in need of confirmation. Although largely montane, subsp. jacquiniana occurs at quite low elevations in the southern Cape Peninsula. Plants grow in fynbos on stony sandstone slopes. Flowering is very late in the season, mostly november and December, but plants have been collected in January and sometimes in late October. Subsp. tripetala flowers in August and September, usually at lower elevations, sometimes in similar habitats.
Diagnosis: subsp. jacquiniana is poorly differentiated from subsp. tripetala but apart from the later flowering time it can be distinguished by the finely fibrous corm tunics, shorter rhipidial spathes 40-50 mm long, and slightly smaller flowers, usually deep violet, consistently with inner tepals 4-6 mm long (usually shorter or absent in subsp. tripetala). The outer tepals are typically 20-25 mm long, the ovary 4-6 mm long and capsules, 10-14 mm long, all smaller than in subsp. tripetala. As far as is recorded, the violet flowers consistently have white nectar guides whereas M. tripetala has yellow or less often white nectar guides, then speckled with blue dots. The colour illustrations of flowers of the two taxa can readily be compared in Wild Flowers of the Cape Peninsula (Maytham Kidd 1950).
The taxon was named in honor of n.J. Jacquin, whose monumental volumes of coloured illustrations of plants, many of them from the Western Cape and namaqualand, were an inspiration and major resource for accurate plant depictions, including his painting of three variants of Moraea tripetala (Icones plantarum rariorum vol. 2, t. 211), one of which perhaps was thought by Schlechter to represent what is now subsp. jacquiniana. The epithet was first used, without description, by Bolus & Wolley Dod (1903) Goldblatt & J.C.Manning,3217 (Vredenburg): Jacobsbaai, 1 km E of town on calcrete ridge, Erf 890, (-DD), 31 Aug. 2011, Claassens 95 (nBg, holo.;MO, iso.).
Distribution: restricted to the Saldanha District in Western Cape (Figure 11), Moraea hainebachiana is a narrow edaphic endemic of rocky, limestone flats and slopes and calcareous sands along the coast and adjacent hills. Plants grow in humus-rich pockets of loam between fractured limestone as well as in coarse calcareous sand. Populations extend from the Farm Trekoskraal north of Jacobsbaai through the limestone hills north of Saldanha to the southern end of the Donkergat Peninsula in West Coast national Park. Distinctive as the habitat is, typical M. tripetala grows in the same places, but blooms four to five weeks later. M. hainebachiana was evidently first collected in 1932 near Langebaan by the late g.J. Lewis, expert on the systematics of southern African Iridaceae. Her collection was referred without question to M. tripetala at the time. The probability that the species was distinct from M. tripetala was brought to our attention by Koos (Jakobus) Claassens of Jacobsbaai, who has made a thorough study of the local and strongly endemic flora of the Saldanha limestone areas. given its very narrow range, M. hainebachiana must be considered threatened by coastal development although it is probably secure at the southern end of its range in the West Coast national Park under current low wildlife stocking levels.
Diagnosis: named for the Hainebach family of Cape Town for their generous contribution to conservation of the Cape flora, Moraea hainebachiana at first seems to be fairly typical M. tripetala except for its low stature and slightly smaller flowers. On close examination, however, the species exhibits several unusual features. The first of these is the cluster of small, dark grey or bluish cormlets in the foliage leaf axil and at the base of the corm, the latter feature especially pronounced in nonflowering individuals. The foliage leaf is always inserted on the stem well above the top of the cataphyll and clearly separated from it whereas in other species in the complex, the insertion of the foliage leaf is concealed by the cataphyll. The flowers of M. hainebachiana are consistently pale violet to deep blue and have inner tepals 5-6 mm long, somewhat longer than is typical for M. tripetala and unusual in being expanded in the middle, thus spindle-shaped or sometimes oblanceolate and obscurely 3-lobed, and occasionally pilose on the inner (adaxial) surface. More significantly, the filaments are as long as or slightly longer than the anthers, whereas the anthers are slightly longer than the filaments in M. tripetala and in some species of the complex considerably so. The anthers are pale grey-blue and contain whitish or yellow pollen. Orange-red pollen is more frequent in the complex. The nectar guides of M. hainebachiana are also somewhat unusual, consisting of lines of dots on a whitish background with the edges not clearly defined as they are in M. tripetala in which the yellow or white nectar guides are typically edged in darker blue to violet. Vegetative reproduction does not occur in M. tripetala as circumscribed here and the production of multiple cormlets in leaf axils and at the base of the corm is unknown elsewhere in the M. tripetala complex.
Visiting the type locality at Jacobsbaai after flowering to examine capsules and seeds, we found all capsules poorly developed and ready to be shed, if not already fallen, before developing any seeds. We confirmed the condition for two more populations. Microscopic examination of pollen grains shows them to be malformed and evidently infertile. We conclude that M. hainebachiana is a vegetative apomict. Propagation is effected by cormlets that plants liberally produce.  Plants 150-200(-380) mm high. Corm 6-12 mm diam., tunics of hard, wiry, black fibres. Stem usually unbranched, 2 or 3 internodes long. Foliage leaf solitary, sometimes with second leaf from axillary cormlet, usually shorter than stem, rarely longer, 2-3 mm wide when opened flat, widely channelled or leaf halves almost folded together, smooth, apple-green, inserted at ground level or shortly below ground, usually bearing small axillary cormlet; sheathing cauline leaves (30-)45 mm long, green with dry attenuate tips. Rhipidia mostly 3-flowered; spathes green with dry brown attenuate tips, inner to 55 mm long, outer ± half as long as inner. Flowers pale blue, with triangular yellow nectar guides, with dark veins radiating from nectar guide, outer tepal claws 12-14 mm long, limbs ovate, ± 15 mm long, inner tepals hair-like, up to 4 mm long. Stamens with filaments ± 5 mm long, united for ± 1 mm; anthers ± 6 mm long, dark bluish black, pollen white. Ovary linear-cylindric, 10-13 mm, darkly lined vertically on locules and septa; style ± 2 mm long, branches ± 12 mm long, crests ± linear, 8-12 mm long. Capsules ± cylindric, (15-)20-24 × 2 mm. Seeds 0.8-1.5 mm diam., golden brown, shortly winged on angles. Chromosome number unknown. Flowering time: mid-Aug.-mid-Sept. Figure 12.
Distribution: restricted to the Western Cape lowlands between Voëlvlei and Strand and locally in the upper Breede River Valley between Wolseley and Botha, Moraea ogamana occurs in waterlogged stony ground ( Figure 11). Judging by the few collections, the species is rare. With rapid agricultural development in lowland Western Cape M. ogamana must be regarded as endangered, although its existence is secure in the Elandsberg nature Reserve at Bo-Hermon immediately south of Voëlvlei, a small part of its once much wider range.
Diagnosis: the narrow, linear-cylindric ovary 10-13 mm long, often partly included in the spathes, and pale blue flower with outer tepal limbs with yellow nectar guides and dark radiating venation immediately set Moraea ogamana apart in the M. tripetala complex. Associated with these features are the short stature, rarely exceeding 180 mm, fairly small corms, 7-12 mm diam., with wiry, black tunic fibres and a relatively short leaf, usually shorter than, but occasionally exceeding the stem. unlike most populations of M. tripetala in the Western Cape forelands, which have orange-red pollen, the pollen of M. ogamana is white. An additional, but somewhat trivial distinction is the style crests, up to 12 mm, which are unusually long for the relatively small flowers. More importantly, almost all specimens we have seen have a single, small cormlet in the foliage leaf axil. This modest level of vegetative reproduction results in plants sometimes growing in small clumps of three or four individuals. Correlated with the long, angular ovary, the capsules are ± cylindric and normally 20-24 mm long. Capsules of M. tripetala are ellipsoid-oblong and only 10-14 mm long. The species is named in honour of Ms naoka Ogama of Japan for her generous donation to the study of southern African Iridaceae.  Plants 150-250 mm high, growing in clonal colonies. Corm mostly 7-12 mm diam., with tunics of pale, medium to fairly coarse fibres (rarely soft relatively fine fibres), often replaced annually by two daughter corms. Stem simple or 1(2)-branched, glabrous or velvety, sometimes with cormlets in below-ground axils, enclosed at base by brown cataphylls, occasionally accumulating in a collar of fine fibres around base. Foliage leaf solitary, linear, channeled, usually broadly so, 2-5 mm wide, abaxial surface often velvety-pilose; cauline sheathing leaves ± 40-52 mm long, glabrous or velvety, green with dry apices. Rhipidia several-flowered; spathes green, dry and attenuate above, becoming dry entirely, glabrous or velvety, inner mostly 45-55 mm long, outer ± half to two thirds as long as inner. Flowers mostly purple or violet to dark blue (rarely pink, dull yellow or buff-brown), nectar guides velvety, white A B C speckled with purple or dark blue, often with larger dark mark at base, often slightly scented of honey or vanilla, outer tepals 20-31 mm long, claws pale mauve often with darker central line and spotted purple-black, 10-12 mm long, velvety, limbs ± broadly lanceolate, (9-)11-18 × 10-14 mm, apex obtuse-apiculate, inner tepals erect, hair-like, acute, 1.5-2.5 mm long. Stamens with filaments ± free, 3.5-6.0 mm long; anthers 4.5-7.0(-8.0) mm long, pollen white (often red in west of range, rarely pale blue). Ovary exserted from spathes, 6.0-9.5 mm long, flushed red; style vestigial, < 0.5 mm long, branches ± 10 mm long; crests linear, erect or arching inward, 6-9 mm long. Capsules narrowly ellipsoid, 11-16 mm long, sometimes with thickened apical rim. Seeds 1.7-2.0 × ± 1 mm, facets with spongy edges and prominent spongy micropylar crest. Chromosome number 2n = 12 (Table 1). Flowering time: mostly Sept.-mid-Oct. Figure 13.
Distribution: relatively widespread, Moraea amabilis extends from the Olifants River Valley to the Bokkeveld Escarpment and in an arc across the northern edge of the Roggeveld Escarpment and south through the Roggeveld and Klein Roggeveld to Worcester and the western end of the Little Karoo ( Figure 11). Plants grow in light clay soils or in sandy loam. Locally in the Bokkeveld Mtns and gifberg, plants grow in shallow sandy ground on sandstone pavement where they are usually dwarfed, we assume because of the shallow, nutrient poor soil. In the west of its range plants flower from late August to mid-September, but at higher elevations to the east, plants bloom from mid September into October, sometimes as late as the end of that month.
Diagnosis: Moraea amabilis is recognized in the Moraea tripetala complex by the virtually free filaments (united for < 4 mm) and inner tepals reduced to hair-like cusps up to 2.5 mm long, and by the distinctive mode of vegetative reproduction in which the parent corm is replaced annually by two new corms. Plants also often have a cormlet in the leaf axil and as a result form clonal colonies. This unusual characteristic is only evident if the plants are carefully extracted from the ground (see Figure 14A). The species is most likely to be confused with M. grandis, which also has virtually free filaments but that species is larger in all features, notably the ovary, 6-9 mm long in M. amabilis vs. 10-12(-15) mm in M. grandis, and anthers 4.5-7.0(-8.0) mm and up to half again as long as the filaments vs. 9-11 mm long and twice as long as the filaments in M. grandis. The corms of M. amabilis have pale (rarely dark) brown, coarsely fibrous tunics with prominent vertical elements in contrast to the finer tunic fibres in M. grandis. Apart from smaller flowers, M. amabilis often has white pollen in contrast to the consistently bright red-orange pollen of M. grandis ( Table 2).
Capsules of Moraea amabilis, typically 11-16 mm long, contain relatively small, sharply angular seeds, ± 2 mm at longest axis, with a prominent, pale, spongy micropylar crest ( Figure 14D). In contrast, capsules of M. grandis are significantly larger, up to 19 mm long, have a thickened apical ridge and the large, dark brown seeds are up to 3 mm long ( Figure 15D. A velvety to pilose pubescence is universal on the leaves, stem and spathes of western Karoo populations of M. amabilis but those from the Bokkeveld Mtns, gifberg and Cedarberg-Olifants River Mtns usually have smooth leaves although some notable exceptions include Goldblatt & Porter 13522 from Kransvlei and Levyns 10155 from Paleisheuwel. We place only moderate confidence in the pubescence character as some other members of the M. tripetala complex occasionally have hairy leaves (notably M. mutila), but we have seen no collections of M. grandis with hairy leaves. On the dolerite flats near Farm Keiskie, southeast of Calvinia, plants of M. grandis (Goldblatt et al. 13366) were entirely hairless, whereas M. amabilis growing nearby had pubescent leaves, stems and sheathing leaves (Goldblatt et al. 13365). The possibility that M. amabilis and M. grandis hybridize in the nieuwoudtville area where the two cooccur is discussed briefly under the latter species.
Plants from near Worcester match Moraea amabilis vegetatively, notably in their corm tunics and velvety leaves and stems but have unusual, buff-yellow, or dusty pink flowers (e.g. Goldblatt 4089). Plants from the Rabiesberg, not far distant, also have velvety stems and leaves and were described as having brown (we assume buff-brown) flowers. These colour variants are unique for the M. tripetala complex.
Moraea amabilis was described by Friedrich Diels in 1910 based on plants he had collected in September 1900 when he lived in Calvinia. goldblatt (1976b) erroneously designated Diels 626 as the holotype, overlooking the second collection cited in the protologue, Diels 1169 (incidentally no longer extant). The so-called holotype becomes the lectotype. Treated merely as a variant morph of M. tripetala by goldblatt (1976b), we now regard M. amabilis as one the most distinctive members of the M. tripetala complex in its habit of often producing two new corms in place of the single parent corm. Flowers of the type, evidently somewhat shrunken, have tepal claws 10-12 mm long and limbs ± 12-14 × 11 mm, an ovary ± 7 mm long, and the stamens have filaments ± 3.5 mm and anthers 5-6 mm long. Leaves and corms are lacking, but as these were described they were presumably present in the syntype, now lost, Diels 1169. The extant type material most closely matches the morph from the sandstone pavement in the Bokkeveld Mtns close to the edge of the escarpment. Plants 200-400 mm high, usually in small colonies. Corm mostly 10-15 mm diam, with tunics of light (rarely dark brown), relatively fine, firm or soft fibres, rarely accumulating in a thick mass; often producing two new corms to replace parent corm. Stem glabrous, rarely velvety to pilose, simple or 1(2)-branched, base sheathed by dry light brown cataphylls, these usually persisting as collar of fine fibres around base; usually producing one or more cormlets in leaf or cataphyll axils. Foliage leaf solitary, shallowly channelled, usually exceeding stem and up to twice as long, 3.5-5.5 mm wide, glabrous; cauline sheathing leaves green or becoming dry from tips, long-attenuate, 55-70 mm long. Rhipidia several flowered; spathes green with dry, attenuate tips, glabrous, inner 55-80 mm long, outer ± half as long as inner. Flowers pale blue to pale violet or light purple, nectar guides small, triangular, yellow (rarely cream) on a white background and edged dark blue, short-velvety (rarely glabrous), scented of vanilla, outer tepals 30-40 mm long, claws 12-18 mm long, velvety, limbs ± orbicular, widest above middle, usually ± as wide as long, 15-20(-23) × 15-20 mm, plane, obtuse-apiculate, spreading at 45-60°, inner tepals hairlike, erect, 1.5-3.5 mm long. Stamens with filaments ± free or united < 0.2 mm, 4.0-5.5 mm long; anthers 8-11 mm long, usually not reaching stigma lobes, pollen orange-red. Ovary (10-)12-15 mm long, slightly narrowed below apex; style vestigial, < 0.4 mm long, branches ± 15 mm, crests linear, 8-12 mm long. Capsules 16-19 mm long, narrowly ovoid-oblong with thickened rim. Seeds relatively large, 2-3 × 1.8-2.3 mm, facets rounded, angles with wing-like ridges. Chromosome number 2n = 12 (Table 1). Flowering time: mainly late Aug.-late Sept. Figure 14.
Distribution: restricted to the northern end of the western (winter rainfall) Karoo, Moraea grandis extends from the Langberg west of Loeriesfontein through the higher country between Loeriesfontein and Calvinia and along the Bokkeveld Plateau to the northern end of the Roggeveld Escarpment ( Figure 15). Plants favour heavier soils and are most often found in heavy red clay among dolerite rocks but also grow on lighter soils derived from tillite and shale.
Diagnosis: the most striking of the species of the Moraea tripetala complex, M. grandis has relatively large corms up to 15 mm diam, with tunics usually of fine, soft (rarely firm) fibres, and the largest flowers in the alliance. The outer tepals are 30-40 mm long (Table  2) and have ± orbicular limbs 18-23 × 15-20 mm, thus almost as wide as long, and have unusually small nectar guides, usually yellow (sometimes cream) on a white background, minutely spotted with dark blue, and usually edged with a darker colour than the rest of the tepal limb. The limbs are also widest in the upper third  and usually have an abrupt, comma-like tip. The flowers have a pleasant, vanilla-like odour. Like its relative, M. amabilis, it typically grows in colonies resulting from vegetative reproduction by axillary cormlets and the sometimes replacement of the parent corm by two daughter corms. It also resembles M. amabilis in having virtually free filaments (united for < 0.2 mm), but the anthers, 8-11 mm long, are at least twice as long as the filaments and sometimes 2.5 × as long, easily the largest in the M. tripetala complex. The pollen is consistently bright orange-red, not unusual for the entire group, and contrasting with the white pollen in most western Karoo populations of M. amabilis, which has smaller anthers 4.5-7.0 mm long and ± as long as or up to 1.5 × as long as the filaments.
Moraea grandis sometimes grows in close proximity to M. amabilis, and the two can be seen flowering together on the farm Keiskie south of Calvinia. Difficulty in distinguishing the two in the immediate vicinity of nieuwoudtville is probably the result of hybridization and introgression between the two species, which share the same large-bodied bees as pollinators.
Moraea grandis has been illustrated in several wild flower volumes notably in the Nieuwoudtville, Bokkeveld Plateau and Hantam wildflower guide (Manning & goldblatt 1997: 69) and The color encyclopedia of Cape bulbs (Manning et al. 2002: 19, 307). The photographs show the distinctive, broad outer tepal limbs oriented almost vertically in fully open flowers.