The Cape genus Micranthus ( Iridaceae : Crocoideae ) , nomenclature and taxonomy

The genus Micranthus (Pers.) Eckl., has traditionally been treated as comprising three species, all with virtually identical, bilaterally symmetric, deep or pale blue to white flowers arranged in crowded, 2-ranked spikes and with divided style branches, but differing in their foliage. Examination of plants in the field and herbarium shows that there are four additional species. M. filifolius Goldblatt & J.C.Manning, from the Caledon District of the southwestern Western Cape, has up to six, filiform leaves, the blades of at least the lowermost terete and cross-shaped in section, and usually pale blue-mauve flowers. M. simplex Goldblatt & J.C.Manning from high elevations on Zebrakop, Piketberg, has the smallest flowers in the genus, white but tinged lilac as they age, linear leaves up to 1.5 mm wide, and undivided style branches. M. cruciatus Goldblatt & J.C.Manning, from the northern Cedarberg and Bokkeveld Mtns, has up to four leaves, the lower with linear or terete blades with heavily thickened margins and central vein and relatively large flowers, unusual in having the style dividing at the mouth of the perianth tube into particularly long branches, these deeply divided as is typical of the genus. M. thereianthoides Goldblatt & J.C.Manning, from the Paardeberg south of Malmesbury, is unique in the genus in having flowers with an elongate perianth tube. We also document the occurrence of large populations of putative hybrids at some sites. We provide a complete revision of Micranthus with original observations on leaf anatomy, pollen morphology and reproductive biology and discuss its confused taxonomic and nomenclatural history and that of the three common species of the genus, known for over 150 years. In so doing, we neotypify Gladiolus alopecuroides L. (1756) [= Micranthus alopecuroides (L.) Eckl. (1827)], type of the genus, and choose lectotypes for M. plantagineus Eckl. var. junceus Baker (1892) and Gladiolus fistulosus Jacq. Now with seven species, Micranthus remains endemic to the Cape flora region, extending from its extreme northern limit in the Bokkeveld Mtns south-eastwards to Port Elizabeth. We also deal with the genera Paulomagnusia Kuntze and Beilia Kuntze with which Micranthus has sometimes been associated, although both are nomenclatural synonyms of Thereianthus G.J.Lewis, a genus close allied to Micranthus. 1 B.A. Krukoff Curator of African Botany, Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.goldblatt@mobot.org. 2 Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, 7735 Claremont, Cape Town. E-mail: j.manning@ sanbi.org.za. 3 Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209. 4 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, USA. Email: roy.gereau@mobot.org. 128 Bothalia 43,2 (2013) three species growing together locally with only very small habitat differences, if any, so habitat preferences are far from absolute. Several additional populations of Micranthus extend the range of leaf morphology in the genus. Plants at high elevations in the Piketberg (Goldblatt & Manning 10172, MO, NBG) have narrow, straight to falcate leaves ± 1 mm wide with one or more strongly thickened veins, small, white flowers fading pale lilac, and undivided style branches. These plants grow in an unusual habitat for Micranthus, crevices and shallow pockets of soil on wet sandstone rocks. A second series of populations (e.g. Goldblatt 10438 MO, NBG) from the northern Cedarberg and Bokkeveld Mtns has long, slender, linear leaves with a heavily thickened main vein and equally thick margins, thus often cross-shaped in transverse section (Figure 1D). These slender, often tall, plants also stand out in having the style dividing at the mouth of the perianth tube, with unusually long style branches divided for less than a FIGURE 1.—Leaf anatomy in Micranthus. A, M. alopecuroides, Elandsberg Nature Reserve, Goldblatt & Manning 13616; B, M. filifolius, Kogelberg, no voucher; C, M. filifolius, Drayton, Goldblatt & Manning 13623; D, M. cruciatus, Pakhuis, Goldblatt & Porter 13766; E, M. tubulosus, Rondebosch Common, Goldblatt & Manning 13620; F, M. plantagineus, Drayton, Goldblatt & Manning 13632. Scale bar: 500 μm. A B C

The genus Micranthus (Pers.)Eckl., has traditionally been treated as comprising three species, all with virtually identical, bilaterally symmetric, deep or pale blue to white flowers arranged in crowded, 2-ranked spikes and with divided style branches, but differing in their foliage.Examination of plants in the field and herbarium shows that there are four additional species.M. filifolius Goldblatt & J.C.Manning, from the Caledon District of the southwestern Western Cape, has up to six, filiform leaves, the blades of at least the lowermost terete and cross-shaped in section, and usually pale blue-mauve flowers.M. simplex Goldblatt & J.C.Manning from high elevations on Zebrakop, Piketberg, has the smallest flowers in the genus, white but tinged lilac as they age, linear leaves up to 1.5 mm wide, and undivided style branches.M. cruciatus Goldblatt & J.C.Manning, from the northern Cedarberg and Bokkeveld Mtns, has up to four leaves, the lower with linear or terete blades with heavily thickened margins and central vein and relatively large flowers, unusual in having the style dividing at the mouth of the perianth tube into particularly long branches, these deeply divided as is typical of the genus.M. thereianthoides Goldblatt & J.C.Manning, from the Paardeberg south of Malmesbury, is unique in the genus in having flowers with an elongate perianth tube.We also document the occurrence of large populations of putative hybrids at some sites.We provide a complete revision of Micranthus with original observations on leaf anatomy, pollen morphology and reproductive biology and discuss its confused taxonomic and nomenclatural history and that of the three common species of the genus, known for over 150 years.In so doing, we neotypify Gladiolus alopecuroides L. ( 1756) [= Micranthus alopecuroides (L.) Eckl.( 1827)], type of the genus, and choose lectotypes for M. plantagineus Eckl.var.junceus Baker (1892) and Gladiolus fistulosus Jacq.Now with seven species, Micranthus remains endemic to the Cape flora region, extending from its extreme northern limit in the Bokkeveld Mtns south-eastwards to Port Elizabeth.We also deal with the genera Paulomagnusia Kuntze and Beilia Kuntze with which Micranthus has sometimes been associated, although both are nomenclatural synonyms of Thereianthus G.J.Lewis, a genus close allied to Micranthus.Bothalia 43,2 (2013) three species growing together locally with only very small habitat differences, if any, so habitat preferences are far from absolute.
Several additional populations of Micranthus extend the range of leaf morphology in the genus.Plants at high elevations in the Piketberg (Goldblatt & Manning 10172, MO, NBG) have narrow, straight to falcate leaves ± 1 mm wide with one or more strongly thickened veins, small, white flowers fading pale lilac, and undivided style branches.These plants grow in an unu-sual habitat for Micranthus, crevices and shallow pockets of soil on wet sandstone rocks.
A second series of populations (e.g.Goldblatt 10438 MO, NBG) from the northern Cedarberg and Bokkeveld Mtns has long, slender, linear leaves with a heavily thickened main vein and equally thick margins, thus often cross-shaped in transverse section (Figure 1D).These slender, often tall, plants also stand out in having the style dividing at the mouth of the perianth tube, with unusually long style branches divided for less than a quarter of their length.They grow in seasonally marshy sites in peaty soil, often in moss on sandstone pavement, and appear to flower particularly well after fire.
A third series of populations from the Caledon District of Western Cape has up to 6 leaves with linear to filiform blades, often cross-shaped in section (Figure 1B, C), with the bases persisting in a well-developed fibrous neck.These plants grow on stony, loamy clay or sandy soils, usually in well drained sites that are dry at flowering time.
Lastly, a population from the Paardeberg south of Malmesbury, only discovered in 2012, has hollow leaves reminiscent of those of Micranthus tubulosus but is unique in the genus in having dark blue flowers with an elongate perianth tube, 20-22 mm long, thus more than twice as long as in any other species of Micranthus.
Consistent treatment of the genus suggests that these divergent populations should logically be recognized as separate species.The circumscriptions of the existing species cannot be expanded to accommodate these plants.We describe these new species as Micranthus cruciatus Goldblatt & J.C.Manning, M. filifolius Goldblatt & J.C.Manning, M. simplex Goldblatt & J.C.Manning and M. thereianthoides Goldblatt & J.C.Manning.Other variants, which we believe are interspecific hybrids, occur locally and we discuss these below.One of them, evidently Micranthus plantagineus × M. tubulosus, is particularly common at the foot of the Elandskloof Mtns.Plants have a flexuose stem, terete, hollow leaves and short spikes of up to 10 flowers, and appeared at first to be a separate species, so different were they from their putative parents.
We review the nomenclature of Micranthus, choose types for the two species currently lacking designated types, and present a systematic revision, thus dealing with collections that do not accord with the current circumscriptions of the three species included in the genus.We also deal with Paulomagnusia Kuntze (1891).When described, Paulomagnusia included two species, one a Micranthus and the other, P. spicatus (L.) Kuntze, now the type species of Thereianthus G.J. Lewis (1941).Our revision includes new observations on leaf anatomy, pollen morphology, and reproductive biology and pollination; these presented following the generic description and nomenclature.

TAXONOMIC HISTORY AND RELATIONSHIPS
Micranthus is most closely allied to Thereianthus, also endemic to the Cape flora region.Lewis (1950) first pointed out an unusual, specialized feature shared by the two genera, namely that the lowermost foliage leaf is inserted on the flowering stem as it is in Lapeirousia Pourret, also in tribe Watsonieae Klatt, rather than on the corm.This means that the corm tunics are formed solely from the cataphylls, without any contribution from the leaf bases as is found in Watsonia Mill.and some other members of the tribe.Molecular systematic studies of plastid DNA sequences confirm the immediate relationship of the two genera, which together are sister to Watsonia plus Pillansia L.Bolus (Reeves et al. 2002;Goldblatt et al. 2008), with Lapeirousia (sens lat.) retrieved as member of a second clade of the tribe, which includes Cyanixia Goldblatt & J.C.Manning and Savannosiphon Goldblatt & Marais.The close relationship of Micranthus and Thereianthus is reflected in their largely shared taxonomic and nomenclatural history.
Note: although Persoon's (1805) infrageneric taxa appear at first to be unranked, the preface (ix) to his Synopsis has the following statement: Melius autem judicavi, eas species (nonnullis forte tamen excipiendis) ab aliis leviter in charactere aberrantes, imprimis si genus minus amplum sit, sub divisione peculiari aut SUBGE-NERE, quo etiam nonnula Botanicorum recentium genera relata sunt, comprehendere, ne ultra necessitatem genera multiplicentur.This is a clear statement that his infrageneric taxa are subgenera.[We have judged it better to include those species (with some exceptions) that in their character(s) are only slightly different from others, especially if the genus is not very large, under the 'particular division' or subgenus (which are some of genera of recent botanists), so that genera are not multiplied beyond necessity.]Deciduous geophytes.Corm axillary in origin, subglobose, rooting from below; tunics coarsely fibrous.Leaves few, the lower 2 or 3 cataphylls, lowermost foliage leaf longest, inserted on stem above corm, blades either plane with a definite main vein and falcate or lanceolate with margins moderately to heavily thickened, or ± tubular and hollow, or terete and ± solid with heavily thickened central vein and margins separated by narrow longitudinal grooves.Stem erect and straight or ± flexuose, simple or few-to several-branched.Inflorescence a congested, 2-ranked spike, usually weakly rotated; bracts short, overlapping, with leathery or dry central portion and broad membranous margins, inner forked apically and shorter than to ± as long as outer.Flowers zygomorphic, lasting several days, blue to violet, mauve, white or flushed lilac, scentless or pleasantly scented, with nectar from septal nectaries; perianth tube short, curving outward, ± cylindric below, flaring in upper half; tepals ± equal, dorsal slightly larger and arching over stamens, lower tepals extended ± horizontally.Stamens unilateral and arcuate; filaments slender, free; anthers oblong, held under the dorsal tepal, splitting longitudinally.Ovary ovoid, sessile; style branches slender, usually deeply divided and recurved, or barely notched at apex.Capsules woody, small, narrowly ovoid-ellipsoid or urn-shaped with ovules in lower fourth.Seeds 2-4(5) per locule, 3(4)-sided below, elongate, widest at micropylar end with micropylar crest and micropyle above base, tapering and pointed at chalazal end, surface slightly wrinkled.Basic chromosome number x = 10.
The diagnostic features of Micranthus are the crowded, 2-ranked spike; small, bilaterally symmetric, tubular flowers; distinctive dry outer floral bracts with broad membranous margins; basal leaf inserted on the stem above the level of the corm (shared with Lapeirousia and Thereianthus); and small, narrow capsules, each locule containing up to four slender seeds almost as long as the locules and with a micropylar crest at the proximal end.Micranthus is unique among subfamily Crocoideae in having zonasulcate pollen grains (Figure 2).The sulci are distal as seen at the tetrad stage (S.Nilsson, pers. comm. Oct. 1996) and the zonasulcate condition in Micranthus is thus derived from the basic monosulcate grain by extension of the sulcus until it encircles the grain.Exine sculpturing is reticulate, grading to tectate-perforate close to the aperture margin.Among Crocoideae, only a few species of Thereianthus also have reticulate exine sculpturing (Manning & Goldblatt 2011).Most genera of Crocoideae, including Thereianthus, have sulcate pollen grains, with a pair (sometimes solitary) of narrow bands of exine (elongated opercula) lying parallel to one another along the long axis of the aperture.Other more complex apertures are known in Geissorhiza Ker Gawl.(Goldblatt & Manning 2009).Cyanixia and Zygotritonia have trisulculate grains.All these pollen types have tectate-perforate exine with small supratectal spinules.
Leaf marginal anatomy in species with plane leaves conforms to the norm for Watsonieae in combining unspecialized marginal epidermal cells and a marginal vein with a sclerenchyma cap below the epidermis.This condition prevails in Watsonia and the Lapeirousia clade (excluding L. corymbosa (L.) Ker Gawl.and its immediate allies), but notably not in Thereianthus or Pillansia, both of which lack a marginal vein or sclerenchyma strand below the unspecialized marginal epidermis (Goldblatt & Manning 1990;Rudall & Goldblatt 1991;Goldblatt et al. 2004;Manning & Goldblatt 2011).
Chromosome cytology: the basic chromosome number for Micranthus is x = 10.One population each of the four species counted, namely M. alopecuroides, the new M. filifolius (reported as M. junceus), M. plantagineus (as M. junceus) and M. tubulosus, are diploid, 2n = 20 (Goldblatt 1971;Goldblatt & Takei 1997).The base number and karyotype, consisting of one long and nine short chromosome pairs, are matched exactly in Thereianthus.The related genus Watsonia has x = 9 and a derived karyotype with two long chromosome pairs.Pillansia, the fourth and last genus of this lineage of Watsonieae, also has x = 10, with its single species tetraploid, 2n = 40 (Goldblatt 1977; not 44 as originally published by Goldblatt 1971).
Reproductive system, compatibility and pollination: virtually nothing has been reported about the reproductive system in Micranthus, but we infer that self-incompatibility and compatibility are important in the evolution and distribution of the genus.It is notable that three species, M. alopecuroides, M. plantagineus and M. tubulosus typically have all flowers producing a full complement of capsules and we infer self-compatibility and facultative autogamy for these species.In contrast, M. filifolius and M. thereianthoides exhibit lower capsule production and we infer self-incompatibility for these species.We are unable to infer compatibility relations for M. cruciatus and M. simplex as good fruiting material is not available.Significantly, the putatively selfcompatible species M. alopecuroides, M. plantagineus and M. tubulosus have the widest ranges in the genus, with M. plantagineus occurring over the entire range of the genus.M. filifolius has a modest range, entirely within the Caledon District of Western Cape but M. cruciatus, M. simplex and M. thereianthoides are local endemics, the latter two currently known from only one or few populations.Self-incompatibility is believed to be ancestral for Iridaceae (Goldblatt & Manning 2008).
In genera that we have studied, we have found relatively few species to be facultatively autogamous, and we assume such species are specialized.Reversals from self-compatibility to incompatibility are believed to be unlikely.Thus in Micranthus we infer that the self-compatible M. alopecuroides, M. plantagineus and M. tubulosus are derived for this character.The latter two species are also derived in their hollow leaves as outgroup comparison indicates that plane, isobilateral leaves are the plesiomorphic condition (present in most members of the family and universal in sister genus Thereianthus).Reproduction through aerial cormlets that replace flowering on the spike axis is also known only in these three species.The arrangement of species in our account reflects our belief that the self-incompatible species are closer to the ancestral stock of Micranthus.
The small flowers of all species except Micranthus thereianthoides are so similar in size, shape and colour (perianth tube 3-5 mm long) that they almost certainly share the same generalist pollination ecology, though we have only recorded insect visitors for M. alopecuroides, M. plantagineus and M. tubulosus.Goldblatt & Manning (2006) regarded these three species as having a generalist pollination strategy and insect visitors to these species include large-bodied bees (Apidae), bee-flies (Bombyliidae), hopliine beetles (Scarabaeidae: Hopliini) and butterflies.Among the latter are Pieris helice (Pieridae) (M.plantagineus) and Cynthia cardui and Colias electo (Pieridae) (M.tubulosus).New observations confirm to the generalist pattern with anthophorine bees and wasps including Delta cf.caffra (Eumenidae), a species of Sphecidae visiting M. plantagineus.The longer perianth tube of M. thereianthoides, 22-25 mm long, suggests a specialized pollination system using a long-proboscid pollinator, possibly a long-proboscid fly species or large butterfly.The nectar reward, retained in the lower part of the perianth tube, is only accessible to pollinators with a proboscis at least 18 mm long, thus excluding access to smaller butterflies, bees, bee-flies and wasps that visit flowers of other Micranthus species.

Key to the species
Note: plants with the lower part of the spike bearing smaller, paler floral bracts subtending one or more small cormlets may be Micranthus junceus or M. tubulosus or may be hybrids involving these two species or with M. alopecuroides and are not accommodated in the key.Plants mostly 200-450 mm high, base usually sheathed with collar of short fibres.Corm mostly 10-12 mm diam., tunics of dark brown, relatively coarse, reticulate fibres, drawn into short bristles above.Stem usually simple or 1-or 2(3)-branched, when unbranched often with one or more scales below base of spike, usually bearing 1 or more cormlets in axil of lowermost foliage leaf.Leaves 2-4(5), lowermost 1 or 2 plane, broadly to narrowly falcate (occasionally ± lanceolate) or linear, (2-)5-10(-15) mm wide, with moderately prominent main vein; margins slightly or occasionally heavily thickened (De Vos 2288), hyaline when dry; upper 1 or 2(3) leaves largely sheathing.Spike mostly 40-80-flowered, often much congested, with internodes 1.5-3.0mm long; bracts 5-7 mm long, outer with broad to narrow brown centre and translucent membranous margins, inner ± as long as outer, notched apically, translucent with 2 dark veins slightly broader toward base; lower or all nodes sometimes vegetative and then bracts paler in colour and subtending one (or more) cormlets in each axil.Flowers usually dark blue, sometimes pale blue, often lower third to fourth of tepals paler blue or white, distally edged with a thin darker blue line, unscented; perianth tube ± 5 mm long; tepals subequal, elliptic, 7-8 × ± 3 mm, with short narrow, claw-like base.Stamens with filaments ± 5 mm long, diverging in upper half; anthers oblong, 3-4 mm long, pale mauve; pollen white to pale blue.Style ± 7 mm long, mostly dividing between upper third of filaments and lower third of anthers; branches ± 1.2-1.6 mm long, divided for ± half their length.Capsules oblong to narrowly ovoid, ± 5 mm long but ± 4 mm long when dry. Seeds angular-elongate, 3.5-4.0mm long, 3 or 4 per locule.Flowering time: October in the north, November to December in the south.
Distribution: centred in the southwestern Western Cape, Micranthus alopecuroides has a relatively narrow range, extending from the Cape Peninsula north into the Olifants River Valley and east locally to Hermanus and Swellendam (Figure 3).Plants typically grow on welldrained clay or loamy, seasonally wet, slopes and flats but have also been recorded on sandy ground.The Olifants River Valley populations grow in thin clay or sandy gravel, often over rocky pavement that is totally dry even before flowering commences.
Diagnosis: Micranthus alopecuroides is distinctive in its plane (Figure 1A), sometimes very broad basal leaves; the blades lanceolate to linear or falcate, sometimes up to 12 mm wide or exceptionally to 15 mm in plants from the Roman's River area of the upper Breede River Valley.Exceptions are numerous and there are collections with ± linear leaves 3-5 mm wide and only up to 25 mm long (notably Purcell s.n., NBG).The flowers are typical of the genus, usually dark blue, with a perianth tube ± 5 mm long.The spikes of 40 or more flowers are often unusually congested with the internodes ± 1.5 mm long.
Populations from the Olifants River Valley and flowering in October, at least three weeks earlier than elsewhere, stand out in their relatively lax spikes with internodes 2.5-3.0 mm long (vs.± 1.5-2.0mm elsewhere) and relatively short, straight leaves, 5-8 mm wide with particularly prominent mucronate tips.The outer bracts of these plants also differ from those in populations to the south in their broader translucent margins, thus with a significantly narrower central band of green tissue (brown when dry).These plants are typically restricted to thin clay or light sandy soils over rocky pavement and represent a distinctive race of the species.
Hybrids: certain collections from sandy flats south of Malmesbury constitute a puzzle.They consist of plants with abnormally elongated spikes up to 120 mm long, more than 3/4 of their length bearing small, pale bracts each enclosing not a flower but a small cormlet.Only the top fourth of the spikes have properly formed, dark brown bracts subtending either pale blue flowers (e.g.Goldblatt & Manning 10431, MO, NBG, with tubular leaves) or deep blue flowers (Goldblatt & Manning 10432, MO, NBG, with plane leaves).The leaves, either tubular or plane with a central vein, correspond to Micranthus tubulosus or M. alopecuroides respectively.We conclude that these plants constitute hybrids or a hybrid swarm with M. alopecuroides as one parent and M. tubulosus or possibly M. plantagineus as the other.Microscopic examination of the pollen shows some apparently normal grains and others smaller than normal The status of these plants is uncertain but we suspect them to have a hybrid origin.
[Note 1.Daniel Solander, the unacknowledged author of Hortus kewensis published under William Aiton's name (1789), described the new species Ixia plantaginea 'foliis linearibus strictis, spica disticha imbricata', based on a collection of Francis Masson and, for reasons that are obscure, at the same time cited Linnaeus's Gladiolus alopecuroides in synonymy.The epithet plantaginea, alluding to the similarity of the inflorescence to that of Plantago L., is no more apt than Linnaeus's recalling the resemblance to the grass, Alopecurus L., and constitutes an illegitimate superfluous name.Nevertheless, the broad-leaved species remained known by the later epithet plantagineus, until well into the 20th century (e.g.Lewis 1950) despite the leaves being described as narrow and linear in the protologue.The epithet plantagineus was applied to M. alopecuroides by, among others, Ker Gawler (1803), who evidently did not realize it applied to two different species.Baker (1892Baker ( , 1896)), who also used the name M. plantagineus for M. alopecuroides, compounded this error and recognized M. plantagineus var.junceus not realizing that the type of the species was in fact identical with his new variety.][Note 2. Described in 1756 by Linnaeus as Gladiolus alopecuroides, with the brief diagnosis 'foliis linearibus, spica disticha imbricata,' the species was transferred to Micranthus by Ecklon (1827), when he raised Persoon's Gladiolus subg.Micranthus to generic rank.Of the three sheets identified as G. alopecuroides in the Linnaean herbarium, one [LINN 59.13] is a Sparrman collection post-dating the protologue, and the other two cannot be unambiguously related to the name.One [LINN 59.15] is M. tubulosus and the other [LINN 59.14] may be M. alopecuroides but is atypical in its large size, numerous branches and particularly broad leaves that hardly accord with the protologue [leaves linear].We prefer to choose a neotype: Barker 3384, which has relatively narrow leaves and conforms exactly to the protologue.This action unambiguously preserves the current application of the name to the plane-leaved species (Lewis 1950;Goldblatt & Manning 2000).] [Note 3. Gladiolus minutiflorus Schrank (1822) has been associated with Micranthus alopecuroides, which Schrank also recognized (as Gladiolus), but the description is vague (flowers small, secund, tepals subequal) and we are unable to determine the plant to genus with confidence, let alone to species.Schrank did, however, explicitly describe the leaves as short, striate and narrow, the lower ± 5 inches (125 mm) long.No authentic material has been located either at the Munich (M) or Brussels (BR) Herbarium, the institutions where the types of Schrank's species, where they exist, are believed to be located.]Leaves (2)3, plane, linear or falcate, ± 1 mm wide, usually with 1 or 2 prominent veins, margins thickened, hyaline when dry. Spike 16-to 40-flowered; bracts purple-brown with broad translucent, brown-flecked membranous margins, ± 5 mm long, inner bracts ± as long as outer, membranous with 2 dark keels, notched at apex.Flowers white fading to lilac, outer tepals tipped pale lilac, with subapical brown ridge on reverse; perianth tube ± 3 mm long; tepals oblong, ± 4 × 1.2 mm.Stamens with filaments ± 2.5 mm long; anthers oblong, ± 2.5 mm long.Style ± 7 mm long, dividing opposite middle of anthers; branches ± 1 mm long, barely notched at apex.Capsules oblong, slightly warty in distal half, ± 4 mm long.Seeds elongate-angular, ± 3 mm long.Flowering time: December to at least mid-January.
Distribution: known only from the slopes of Zebrakop, highest peak in the Piketberg, Micranthus simplex, like M. cruciatus, grows in shallow soils in moss or in rock crevices on wet sandstone rocks (Figure 3).The habitat remains moist as late as January when the species blooms.
Diagnosis: unusually small for the genus, stems of Micranthus simplex rarely exceed 180 mm and the white flowers with lilac-tipped outer tepals are distinctive, other species having flowers in shades of deep to pale blue or blue-mauve, or occasionally white.The inflorescence has the appearance of being relatively lax, the bracts of the lower flowers of the spike not overlapping those above them, but the upper bracts are as closely set as in other species.It is one of two species of Micranthus with consistently plane leaves; the other, M. alopecuroides, is a taller plant with congested spikes of 40 to 80 flowers and broader leaves mostly 5-12 mm wide.The flowers of M. simplex are the smallest in the genus, the perianth tube just 3 mm long and the short anthers ± 2.5 mm long.The short, undivided style branches, ± 1 mm long, are likewise unusual for Micranthus, other species of which normally have the style branches somewhat to considerably longer and divided for at least one third their length.Plants 300-800(-1200) mm high, base weakly sheathed by fine fibres.Corm 10-15 mm diam., tunics of fine to moderately coarse, dark brown, reticulate fibres.Stem simple or rarely branched, with solitary cormlet in axil of second leaf and sometimes also third leaf.Leaves (4)5 or 6, green or drying at flowering, lowermost 2 or 3 longest, blades 100-300(-800) mm long, 2.5-5.0 (-15) mm diam., tubular and hollow, sometimes inflated, acute-mucronate, upper leaves progressively shorter and narrower, uppermost bract-like and entirely sheathing.Spike 10-to 40(-70)-flowered, bracts brown with broad translucent membranous margins, 8-11(-15) mm long, as long as 1.5-2.0spike internodes, inner bracts slightly shorter than outer, forked apically, membranous with 2 dark keels broadened toward base.Flowers suberect, dark violet or purple, unscented; perianth tube ± cylindric, 22-25 mm long, tepals oblong, 5-6 × 1.5-2.5 mm, reverse of outer tepals with prominent subapical ridge.Stamens with filaments 6-8 mm long, exserted ± 3 mm; anthers oblong, 3.5-4.0mm long.Style 24-27 mm long, dividing between middle and slightly beyond anthers, branches ± 1.5(-2.0)mm long, divided for ± half their length.Capsules ovoid, smooth, 5-6 mm long, with ± 4 seeds per locule.Seeds elongate-angular, tapering to points at both ends, ± 3.5 mm long.Flowering time: January.Figure 4.
Distribution: a highly local endemic, Micranthus thereianthoides is restricted to the Paardeberg near Malmesbury (Figure 3), where it grows at mid to upper altitudes along the banks of seasonal streams, the corms usually wedged among granite rocks, sometimes in humic loam, where the plants are more robust.Plants are locally plentiful along several streams on the range.The long-tubed, violet flowers are evidently adapted to pollination by long-proboscid flies.The incomplete fruit set in wild plants suggests that M. thereianthoides is an obligate outcrosser.
Diagnosis: Micranthus thereianthoides closely resembles M. plantagineus and some forms of M. tubulosus in its cylindrical leaves but is unique in the genus in the relatively large floral bracts, 8-11 mm long, and most strikingly in its dark violet flowers with elongate, cylindrical perianth tube 22-25 mm long, thus ± twice as long as the bracts (Figure 4).The species appears never to develop cormlets in the floral bract axils.

Gladiolus fistulosus
Distribution: typically a species of lower slopes usually on clay and granite-derived soils but also on sandstone, Micranthus tubulosus is restricted to the western half of Western Cape.It extends north of the Cape Peninsula as far as the northern Cedarberg, where an early (1923) collection documents its occurrence at Heuningvlei, and no further east of the Peninsula than Suurbraak near Swellendam and the Agulhas Peninsula (Figure 5).Like other species of the genus, it blooms late in the season when the hollow, inflated leaves are often dry and brown.Plants from the Pakhuis Mtns growing in moist, sandy ground are exceptional in their small size (leaves up to 100 mm long) and require additional study.A much dwarfed fragment of Micranthus tubulosus, said to be from Garies (Caporn s.n., ex hort.Kirstenbosch (as Nat Bot Gard.915/15) in BOL) is unlikely to be from there as no other records of the genus from Namaqualand exist.
Diagnosis: the inflated, tubular, falcate leaves (Figure 1E) are diagnostic for the species, the spikes and flowers of which differ hardly at all from those of Micranthus alopecuroides.A particularly distinctive feature of the leaves is the prominent brown mucro at the obtuse to ± truncate apices.The leaves are often ± dry at flowering time- Marloth's (1915: plate 41) has a particularly apt illustration of the species.As in M. plantagineus, one or more cormlets may be produced in the lower axils of the spike, a phenomenon first noted by Ker Gawler (in Sims 1801) and later confirmed by Lewis (1950).The condition is more frequent, although not consistent, in M. plantagineus.Despite its apparently preferred habitat on relatively dry slopes, M. tubulosus can occasionally be found on moist sandy flats, sometimes co-occurring with M. alopecuroides and M. plantagineus (Wurts 519 consists of just such a mixture, M. tubulosus and M. plantagineus evidently found growing in close proximity).Hybrids between these two species at shared sites blur their usually clear foliar differences.We discuss putative hybrids between Micranthus tubulosus and M. alopecuroides or M. plantagineus in more detail below.
Plants collected near Saron (e.g.Schlechter 10618) are unusually small, mostly 100-150 mm but some just 70 mm tall, and have shorter, fewer-flowered spikes than usual.They appear linked to taller, more robust specimens by a range of intermediates.In contrast, plants from Gouda (Barker 9861), nearby, are exceptionally robust, up to 600 mm tall, with leaves almost as long, and the white flowers have tepals 10 mm long, the outer 2.5 mm wide.
A curious feature of Micranthus tubulosus is that populations may consist of a mixture of some plants with entirely fertile spikes and others with the lower part of the spike sterile (e.g.Ecklon & Zeyher Irid 192 and 190).All three specimens of Goldblatt 8711 and several of Purcell 43 have spikes sterile in the lower half.Particularly short leaves in Goldblatt 8711 are also puzzling but not unique.
History: Long known as Micranthus fistulosus (Jacq.)Eckl.(e.g.Baker 1896), based on Gladiolus fistulosus Jacq.( 1797), that combination was in fact not valid, though it was used as M. fistulosus Eckl.(a nomen nudum assumed to be a valid combination) by Baker (1892Baker ( , 1896)).By citing the basionym, Baker's use of the name M. fistulosus becomes a valid (albeit unintended) combination, also superfluous through his citing of valid earlier synonyms, including Gladiolus tubulosus Burm.f.(1768).Jacquin's illustration of G. fistulosus has two plants: we designate as lectotype the left hand one, which has dark blue flowers and leaves typical of M. tubulosus.The right hand plant, which has pale blue flowers and the upper leaf more typical of M. plantagineus, may be a hybrid with that species.The sterile lower nodes of the spike, bearing silvery bracts, are more typical of M. plantagineus and represent at least a different genotype from the right hand plant.Brown (1929) identified the type of Gladiolus tubulosus among specimens in Burman's herbarium, and realizing that it was an earlier name for M. fistulosus, provided the combination M. tubulosus.A fine illustra- Plants 180-300 cm high, base sheathed with sparse to well-developed collar of fibres.Corm 12-16 mm diam., tunics of relatively soft, fine or thicker fibres.Stem unbranched or rarely with single short branch, without cormlets in leaf axils.Leaves 4-6, green or beginning to dry from tips at flowering time, lowermost longest, reaching to middle of spike to shortly exceeding it, blade either ± terete and ± 1 mm diam.or ± plane and ± 2 mm wide, with heavily thickened central vein and margins, separated when dry by narrow longitudinal grooves, upper leaves shorter, with sheaths overlapping, uppermost 1 or 2 leaves sheathing for most of their length, with short free tips.Spike mostly 18-50-flowered, closely congested, lower bracts always subtending flowers; bracts mid to dark brown, ± 5 mm long, outer with broad translucent membranous margins, inner slightly shorter than outer, with 2 dark keels broadened toward base, notched apically.Flowers pale mauve or mid-blue, unscented; perianth tube ± 5 mm long; tepals oblong, ± 5 × 2.2-2.8mm.Stamens with filaments ± 5 mm long, exserted ± 2.5 mm; anthers oblong, ± 3 mm long.Style ± 7 mm long, dividing between base and middle of anthers; branches 1.0-1.6 mm long, divided for up to half their length, rarely only notched at apex.Capsules narrowly ovoid, smooth, 4-6 × ± 2 mm, with up to 4 seeds per locule.Seeds elongate-angular, mostly 3-sided, tapering to points at both ends, 3-5 mm long.Flowering time: mid-November to late February.Figure 6.
Distribution: centred in the Caledon District of Western Cape, Micranthus filifolius is largely coastal with populations recorded from Steenbras and Cape Hangklip eastward to Hermanus and inland to Shaw's Mtns, the lower slopes of Caledon Swartberg and east to Akkedisberg Pass and Elim (Figure 7).Collections are mostly from clay and clay-loam soils, occasionally from sandy sites, but even collections from the Klein River Mtns above Hermanus at elevations of up to 400 m are from a shale band.The species is particularly abundant after fire (e.g.Drewe 495, 1101) but will flower in unburned veld unless shaded out by taller vegetation.Plants bloom unusually late in the season, with most flowering collections made after mid-January, and two (Gillett 520; Levyns 11269) were in mid-to late February.We have confirmed late flowering at near-coastal sites ourselves but inland populations, as from Drayton Siding, east of Caledon, and Akkedisberg Pass, flower from mid-November to early January and are in fruit before any coastal populations come into flower.We suggest that this early flowering is due to warmer and drier conditions well inland of the coast.The coarser corm tunic fibres and collar of fibres around the base of the stems in these populations are perhaps adaptations to the drier habitat.Plants sometimes co-occur with or grow close to M. plantagineus, which is in fruit when M. filifolius begins to bloom, two or three weeks after the last flowers of M. plantagineus have faded, both at the coast and at inland sites.We have also found M. filifolius growing together with M. tubulosus.
Diagnosis: with its narrow leaves, the lowermost of which is linear or terete (Figure 1B & C), Micranthus filifolius is most like M. plantagineus in general appearance.It differs, however, from that species in several respects, particularly in the solid leaf blades of the Plants 300-450 mm high.Corm globose, 8-10 mm diam., tunics of brown, soft membranous layers not accumulating.Stem simple or 1-branched, usually with 1 or 2 small cormlets in lowermost leaf axil.Leaves (3)4 or 5, lower 3 linear to subterete, ± 1.5 mm wide, margins and midrib heavily thickened with narrow longitudinal grooves between (often cross-shaped in section with 4 narrow longitudinal grooves), reaching to base or middle of spike, uppermost leaf sheathing stem almost to base of spike, with short free portion.Spike up to 70-flowered; bracts brown or straw-coloured with broad translucent membranous margins, ± 4 mm long, inner ± as long as outer, membranous with 2 dark keels, apically notched.Flowers pale blue-mauve (drying ± white) or deep blue, perianth tube ± 3 mm long, tepals oblong, ± 4 × 1.5 mm.Stamens with filaments ± 5 mm long; anthers oblong-linear, ± 3 mm long.Style ± 4 mm long, dividing ± at mouth of tube opposite middle of filaments, branches ± 2.5 mm long, divided for ± one third their length.Capsules oblong, smooth, 5.0-5.5 mm long.Seeds elongate-angular, mostly 3-sided, tapering to points at both ends, ± 3 mm long.Flowering time: mid-November to late December, possibly lasting into January.
Distribution: restricted to the northern Cape flora region, Micranthus cruciatus is known from the northern Cedarberg immediately south of Pakhuis Pass and in the Bokkeveld Mtns southwest of Nieuwoudtville (Figure 7).Plants grow on rocky slopes, in seeps on thin sandy soil over sandstone pavement, flowering in December as the habitat dries out in the hot weather.No doubt the species is rare but the very few collections are probably due to its midsummer flowering when little plant collecting is undertaken.We suspect that M. cruciatus occurs in suitable sites between its few stations, thus in the southern Bokkeveld Mtns and the Gifberg/ Matsikamma Mtn complex and perhaps elsewhere in the Cedarberg.First collected by the late Elsie Esterhuysen in 1941 according to available records, M. cruciatus has elicited no attention until now and was assigned to the broadly similar M. plantagineus (as M. junceus) in herbaria.
Diagnosis: linear-to terete-leaved Micranthus cruciatus is immediately distinguished by its solid, narrow leaf blades, ± 1.5 mm wide, with heavily thickened veins and margins separated by narrow longitudinal grooves (Figure 1D).Leaves are either linear or terete becoming cross-shaped in section distally with only the margins and central vein thickened.Plants broadly resemble M. plantagineus although they are more slender than is usual in that species, which has hollow leaves 2-3 mm diam.and is conspicuous in the production of cormlets in the lowermost and sometimes upper leaf axils.In contrast, M. cruciatus has no more than one or two small cormlets, these borne in the axil of the lowermost leaf.The pale mauve-blue or sometimes dark blue flowers are typical of the genus in shape but notable in the short perianth tube, ± 3 mm long, in the style dividing at the mouth of the perianth tube and in the unusually long style branches up to 2.5 mm long, divided for up to one third their length.Most species of Micranthus have the style dividing opposite the base to middle of the anthers and style branches typically less than 1.6 mm long.The narrow, heavily thickened leaf in M. cruciatus is convergent with that in M. filifolius, but in other critical details the two are very different, the latter with coarsely fibrous corm tunics and a collar of fibres around the base of the stem.
It is noteworthy that typical, hollow-leaved M. plantagineus with dark blue flowers also occurs in the Pakhuis and Bokkeveld Mtns (e.g.Leipoldt 3596 BOL, NBG, PRE) as well as in the Cedarberg, but it has not been recorded growing near M. cruciatus and they evidently have somewhat different habitat preferences.Plants 200-400(-650) mm high, base without collar of fibres.Corm globose, 12-15 mm diam., tunics of dark brown, medium-textured, reticulate fibres drawn into fine points above.Stem erect, simple or with up to 9 short branches, with cormlets in axil of lowermost leaf and sometimes of upper cataphyll and rarely other leaf axils.Leaves (2)3( 4), green at flowering, lower-most leaf longest, blades terete or oval in section, hollow, 2.0-3.5 mm diam., smooth when fresh with translucent veins, when dry, veins appearing thickened with homologue of marginal vein pair more prominent, usually reaching to middle of spike to shortly exceeding spike, uppermost 1 or 2 leaves sheathing for most of length, with free part often slightly longer than sheath.Spike (16-)40-100-flowered, lower bracts sometimes subtending cormlets; bracts mid-to dark brown, ± 6 mm long but slightly smaller if subtending cormlets, outer with broad translucent margins, apices sharply acute and ultimately curved outward, inner bracts ± as long as outer, with 2 dark keels broadened toward base.Flowers usually dark blue, occasionally pale blue or white, evidently sometimes slightly sweetly scented, perianth tube 6-7 mm long, tepals oblong, with thickened subapical ridge on reverse, (4-)6-7 × ± 1.2 mm.Stamen filaments ± 5 mm long; anthers oblong, 3-4 mm long.Style ± 7 mm long, dividing between lower one third and middle of anthers (rarely ± 1 mm below anther bases), branches 1.0-1.8mm long, divided for one third to half their length.Capsules smooth, ± urn-shaped or narrowly ovoid, (3)4-5 × 2-3 mm, with (2)3 or 4 seeds per locule, 5 mm long.Seeds elongate-angular, 3(4)-sided, tapering to points at both ends.Flowering time: October to December (rarely in May).
Distribution: Micranthus plantagineus has a wide range across the Cape flora region, extending from the Bokkeveld Plateau near Nieuwoudtville south to the Cape Peninsula and east to Port Elizabeth (Figure 8).An isolated population from the Anysberg Nature Reserve in the Little Karoo (Vlok 2545) appears typical of the species except for the shorter perianth tube, ± 4 mm long.Plants typically grow in seasonally wet habitats, often in marshy sites, along streams, or at least in places that are waterlogged in the winter months.
Diagnosis: the elongate inflorescence with up to 100 flowers and a perianth that is often deep blue, but sometimes pale blue or white, are unexceptional for the genus and identification of Micranthus plantagineus depends on leaf morphology.The two to four leaves are straight, stiffly erect, hollow and terete to oval in section, ± 2-3 mm diam., and reach or shortly exceed the spike (Figure 1F).When alive the leaves are smooth with the veins evident as paler, translucent lines.On drying, the veins appear hyaline and the veins at the adaxial and abaxial poles are somewhat more prominent.In addition, the stem is often branched, and as many as four (exceptionally nine) short branches may be produced shortly below the base of the main spike, these seldom exceeding half the length of the main spike.Lewis (1950) noted that the lower flowers of the spike are often replaced by cormlets [as many as five may be present in an axil].That feature is not universal and many otherwise typical plants may have normal flowers and capsules from base to apex of the spike.A second characteristic feature of M. plantagineus is the presence of one or more cormlets in the lowermost leaf axil (not invariably present in other species) and occasionally in the axils of the upper cataphyll and one or more of the upper leaves.Plants lack a collar of fibres around the base (in contrast to superficially similar M. filifolius and M. tubulosus).
Collections of Micranthus plantagineus from Grootwinterhoek Forestry Station exemplify this situation: Goldblatt 10451 with dark blue flowers represents the robust morph of the species, but slender-leaved plants (Goldblatt 10452), growing adjacent to stands of the robust morph, have white flowers.Other slender-leaved M. plantagineus (Goldblatt10453) growing nearby have blue flowers.These last two collections consist of shorter, less robust plants, 200-250 mm tall, and have more slender, but still hollow leaves ± 1.5 mm diam.No other Micranthus species were found in the area making the possibility that hybridization has played a role in this pattern of variation unlikely.
We must also mention a collection made by T.M. Salter in May 1935 from Viljoen's Pass.The morphology conforms closely to Micranthus plantagineus in the three terete and hollow foliage leaves, stems without a collar of fibres at the base, cormlets in axils of all leaves and in floral dimensions.Thus the only difference we see is the flowering time, noted as anomalous on the specimen label by Salter.The marshy habitat likewise conforms to the species.A search for the population in May 2013 failed to find any Micranthus species in bloom at this time of year, but the site may simply be lost to farming activity or dam construction.
Putative hybrids between Micranthus plantagineus and M. alopecuroides are discussed under the latter species.
Related species: until now, several collections of plants with filiform-linear and plane or terete, but not hollow, leaves with a thickened central vein and margins (cross-shaped in section) have been included in Micranthus plantagineus (as M. junceus) in herbaria.We believe these are separate species.The several southern Western Cape populations with this leaf type, here referred to M. filifolius, always have four or five, rarely six leaves, the blades ± 1 mm wide vs, mostly three (rarely two or four) leaves in M. plantagineus.These plants have flowers with a consistently shorter perianth tube ± 4 mm long and shorter tepals, also ± 4 mm long vs, both perianth tube and tepals mostly ± 6 mm long in M. plantagineus and often a pale blue mauve to almost white (less often deep blue) perianth.M. filifolius is rarely branched and we have seen no specimens with the lower flowers aborted and replaced by cormlets, both common but not universal traits of M. plantagineus.
Other collections with this derived leaf type are known from the northern Cedarberg and Bokkeveld Mtns and are here treated as the new Micranthus cruciatus Goldblatt & J.C.Manning.These plants have only four, rarely five leaves, the two basal with linear or terete, four-grooved blades, ± 1.5 mm wide, similar to but broader than those of M. filifolius.An important associated character is the style, which divides at the mouth of the perianth tube into unusually long branches 2.0-2.5 mm long, divided for ± one third their length.Other species of Micranthus have the style dividing between the base and middle of the anthers and the style branches never exceed 1.5 mm.
History: long known as Micranthus junceus (Lewis 1950;Goldblatt & Manning 2000), M. plantagineus was evidently first recognized as a distinct species, called Phalangium spicatum by Burman (1768), at least as to the specimen in his collection (now at the Delessert Herbarium, Geneva).Burman provided no validating description, instead merely citing Plukenet's (1694) illustration in part 3 of the Phytogeographia.There is no accompanying text or even polynomial identifying the illustration, plate 310, f. 1., nor does the figure constitute a validating illustration with analysis.Phalangium spicatum Burm.f. is thus a nomen nudum and invalid.Plukenet's illustration is of a broad-leaved plant and does not, in our opinion, represent any species of Micranthus but is probably Ixia scillaris L.Even if any text associated with this illustration is found and if the name is lectotypified on the specimen rather than the Plukenet illustration, the combination M. spicatus (L.) Heyn.( 1847 Bothalia 43,2 (2013) unfortunately superfluous as Gladiolus alopecuroides was cited in synonymy and, likewise, transfers of Ixia plantaginea to Watsonia (Ker Gawler 1803), and Gladiolus Pers.(1805), are superfluous as both authors cited Gladiolus alopecuroides as synonyms.Ecklon (1827), however, intended to transfer the species to Micranthus, where it becomes valid and is treated as a new name from that date rather than a new combination based on I. plantaginea Aiton. Ecklon (1827) recognized M. alopecuroides as a separate species, the first author to differentiate it from M. plantagineus, but whether deliberately or by accident is uncertain.Baker (1892) described Micranthus plantagineus var.junceus, citing no specimens, but later listed several exsiccatae (Baker 1896), all of which were available to him in 1892.We choose a lectotype from among these, Zeyher 1611, a specimen in good condition and representative of the species.The taxon was raised to species rank by Brown (1929), who, at the time also identified Phalangium spicatum Burm.f. as the same species, at least as to the specimen in Burman's herbarium.M. junceus, a name used until now for this plant, becomes a synonym of M. plantagineus.

Hybrids
Interspecific hybrids are not uncommon in Micranthus and are likely to occur when two or more species co-occur.Most striking of the hybrids is that between M. plantagineus and M. tubulosus.The two species flower together at the foot of the Elandskloof Mtns in Elandsberg Nature Reserve and present a remarkable sight.The hybrids are locally very common growing with typical M. tubulosus and are always slightly shorter than the parent, 100-150 cm high, and like it have a well-developed collar of fairly coarse fibres around base.The other parent is less common, but present in small clumps, recognized by its erect habit, straight leaves and pale blue flowers.The hybrid is evidently fertile (plants in fruit have well developed capsules with apparently normal seeds) and stand out in having a slightly flexuose stem and narrower leaves than either parent.Unlike M. tubulosus, which they otherwise most closely resemble, hybrid individuals bear small cormlets at aerial nodes and sometimes at the base of the spike.We have seen similar hybrid plants near Elim where M. tubulosus and M. plantagineus also grew side-by-side.Less common are hybrids between Micranthus tubulosus and M. alopecuroides, but at Elandsberg Nature Reserve we noted both species growing close to one another with apparent hybrids among them.The putative hybrids have short, plane leaves, in outline like those of M. tubulosus but not round in section, although the leaves have an airspace between the two surfaces and lack the visible main veins of M. alopecuroides.
) (= Thereianthus spicatus) prevents the use of Burman's epithet at species rank in Micranthus.Curiously, Phalangium spicatum Houtt.(1780), typified by a good illustration, marks this as the first valid naming of M. plantagineus.Although seeming to refer to Burman's P. spicatum, Houttuyn makes it clear this is his species (Phalangium scapis spicatis mihi, i.e.Houttuyn) and that the Plukenet figure cited by Burman is an entirely different plant.As noted above, Heynhold's combination M. spicatus (L.) Heyn.bars transfer of Houttuyn's epithet to Micranthus.In Aiton's (1789) Hortus Kewensis, Daniel Solander, the unacknowledged author of the species in this work, described Ixia plantaginea based on a collection of Francis Masson.The sheet at BM includes two plants with narrow, stiffly erect, centric leaves (the Masson collection) and three specimens with shorter, plane leaves (Nelson 1777) that are M. alopecuroides.The Masson specimens conform to the diagnosis, 'foliis linearibus strictis, spica disticha imbricata [leaves linear, straight and upright] and constitute the holotype.The name is