New combinations in Antimima ( Ruschioideae , Aizoaceae ) from southern Africa

As a consequence of the emended description of Antimima N.E.Br. by Dehn (1989) and based on extensive comparison of all types of the genera Antimima and Ruschia Schwantes, and on studies of about 250 fresh collections of Antimima. 95 species are tranferred to that genus from Ruschia. The arguments for this step, and bnef diagnoses of the 100 known species of Antimima are given. One species is excluded.


INTRODUCTION
described the genus Antimima with a single species, A. dualis (N.E.Br.)N.E.Br.-this species was first named Mesembryanthemum duale N.E.Br.(Brown 1920) and was transferred to the genus Argxroderma N.E.Br.(Brown 1922), based on similari ties of leaf morphology and the light greyish white colouring of the leaves.When, after 18 years of cultiva tion of the plant, Brown saw a flower and realized that the species was not a member of the genus Argxroderma since it lacked the hypanthium characteristic of that genus (Figure 1 A, B).
The species had meanwhile also been described as Ruschia dualis L. Bolus (1929), but based on a different type.This name has been used for about 60 years, imply ing that the genus Antimima was a synonym of Ruschia Schwantes.
In a comprehensive survey covering all species of the Aizoaceae, material of types of all species in Ruschia and fresh material of ± 250 collections of Antimima was examined.As a result.95 species are here transferred to Antimima.With A. virgata transferred back to Ruschia, based on comparisons of fruits and the type.Antimima at present comprises KX) species.
The subdivision of the genus into subgenera will have to be dealt with on a much broader basis.M. Dehn (pers. comm. 1992) noticed that the circumscriptions of the subgenera need reconsideration, in the light of data which must be derived from extensive studies in flowers and leaves, especially heterophylly and Alicean mor phology of the epidermis.Features of flowers and leaf anatomy are therefore given sporadically in this treat ment.due to lack of data for complete comparisons.

CRITERIA FOR INCLUSION OF SPECIES IN ANTIMIMA Fruit
As demonstrated by Dehn (1989: 199).typical fruits of Antimima possess very large closing bodies filling and blocking the distal exit of the locule completely, i.e. no seeds can be expelled through this opening (Figure 1 C. F-H ).In contrast, fruits of Ruschia have small, hook shaped closing bodies which never form a complete blockage or barrier for the seeds.
In addition, expanding keels in fruits of Antimima are broad and laciniate.diverging distinctly in a ± radial direction and reaching almost to the tip of the valve (Figure 1C, H).In Ruschia. the expanding keels are typ ically shorter and are spread into an almost tangential direction, visible in the open capsule on the valve.Due to these structural differences, fruits of Antimima nor mally open completely (Figure IF), the valves often even recurving below the horizontal plane, whereas cap sules of most species of Ruschia open their valves into an erect position only.
Covering m embranes possess additional closing devices at their distal ends in both genera in principle.In fruits of Ruschia these devices appear mostly in the shape of closing rodlets.rarely as closing ledges or clos ing bulges.Closing rodlets can also occur in Antim im a, but in this genus closing ledges (Figure 1C.F) or the lack of any additional closing devices are more common (Figure 1G. H).The last-mentioned condition is absent in Ruschia.
In seven species of Antimima.the covering mem branes possess radial wings (Figure 1G. H) or other pro trusions on top.clearly visible in younger fruits.Similar features have been found in genera like Cheiridopsis N.E.Br., but never in typical fruits of Ruschia.E,A. pumila,Hartmann 30325 (HBG!), side view of fruiting heterophyllous branch: long connate sheath is papery and smooth, short erect free parts papillate.F, A. dolomitica, Dinter 3782 (BOL, iso.!), radial l.s.through capsule in middle of locule: straight covering membrane (CM) lying distally on big closing body (CB), which is also touched by closing ledge (CL).G, A. fenestrata, Hartmann < 5c Dehn 15626 (HBG!), radial l.s.through capsule in middle of locule: covering membranes sloping down from a high centre ending in a distal recurving pressed against big white closing body (CB); erect wing in nearly radial position stabilizing covering membrane.H, A. eendomensis, Dinter 5207 (B,holo!), part of open capsule with basally distant moderately diverging expanding keels (EK), small valve wings (VW) broadest in middle, very big closing bodies (CB) and covering membranes (CM) pressed against them; covering membranes with radial wings on top lowering towards centre I, A. distans, Hartmann et al. 20682: heterophyllous long shoot with several leaf pairs with short connate sheaths and free long parts (FLP) and connate subterminal leaf pair (CLP) forming long sheath with short free parts enclosing terminal leaf pair.ES, expanding sheet; P, placenta; V, valve; VR, valve rim; W, wing on top of cover ing membrane.
Fruits of the majority of species of Ruschia lack valve wings altogether, with a few exceptions such as Ruschia subgenus Spinosae.Capsules of Antimima are also most ly devoid of valve wings (Figure 1C), but in a number of species narrow, awn-shaped or even broader valve wings are found, always broadest in the middle and tapering towards both ends (Figure 1H).
Finally, fruits of Ruschia develop a long, deep fruit base whereas those of Antimima have rather shallow, mostly bowl-or funnel-shaped capsule bases (Figure ID. E).
Locule numbers are mostly five in both genera, six occurring in several species in Ruschia and in Antimima.

Leaves
Isophylly (i.e.all leaves of a plant are ± equal), is pre sent in all species of Ruschia.In contrast, the species of Antimima develop isophylly (Figure IB, D) or heterophylly on a plant, the latter either as a result of two leaf pairs (Figure IE) of different leaf shapes developing in one season, or as a result of differing leaf shapes devel oping in sequence along a side branch (Figure II).The first condition resembles that in Cheiridopsis subg.Cheiridopsis (Hartmann & Dehn 1987) or Mitrophxllum (Poppendieck 1976).Leaves developing in sequence along a side branch resemble those of Vanzijlia L.Bolus (Hartmann 1983).In both cases, the long-sheathed leaf pair forms a papery, persisting sleeve-like protective cover enveloping the subsequent, assimilating leaf pair during the dry season.The dry. persistent sheath consists mostly of the connate lower parts of the leaves of a pair (Figure IE.I), but in a few cases the leaves are separated to the base but stand closely together forming a protective sleeve-like cover.In these cases, the broad lower parts resemble sheaths that have been slit open, but they were never connate, and the upper parts resemble the free parts of sheathed leaves closely.In general, the epidermis of the sheaths is smoother than that of the free tips, and often the different leaf types differ in their epidermal structures.

Flowers
Flowers of both genera are rather small in the majori ty of species, rarely exceeding 20 mm in diameter (Figure 1A.B).The petals are white to pink or purple and often arranged in five, rarely six.distinctive groups.Filamentous staminodes are present, surrounding the central cone of stamens.In many cases, the tips elongate and recurve, and are often dark purple to almost black in colour.Characters of the flowers do not differ in princi ple between Ruschia and Antimima.neither do inflores cences, which can be solitary or in cymes of various sizes, typical of species but not of either genus.

Growth forms
After the removal of a number of compact species from Ruschia by Glen (1986: Ebracteola Schwantes.Acrodon N.E.Br.) and the re-establishment of Marlothistella Schwantes (Hammer 1995), Ruschia compris es almost exclusively, shrubby to creeping growth forms with long, visible internodes and branching at the distal ends of the stems.
In Antimima, growth forms are variable: compact to caespitose, rarely creeping, or with a compact centre with long shoots climbing into other shrubs.All shrubs with a marked distal branching, proposed for inclusion in Antimima (by Dehn on herbarium sheets), had to be excluded from that genus because of their fruit morphol ogy.which resembles that of Ruschia.

CORRELATIONS BETWEEN CHARACTER STATES
The possession of the typical fruit permits a definite inclusion of a given plant in Antimima.but if fruits on the type sheet are absent or incomplete, the knowledge of the correlation of characters can also help to establish the identity of the material.
In all heterophyllous species with capsules, the expanding keels are basally separated, broad and laciniate, and diverging towards the tips of the valves, not in a tangential position as found in Ruschia.Valve wings are absent or present, mostly forming flanges tapering towards the tips.The closing bodies are mostly large, but in some species small white ones are present.In one species no closing body is developed but the capsules show so many similarities with those of other species that the taxon is included here in Antimima.Furthermore, it is deduced that heterophylly is correlated with the Antimima type of capsule morphology, and as a conse quence, all heterophyllous species formerly belonging to the genus Ruschia are transferred to Antimima.
It should be noted that a revision of Ruschia down to species level is not yet available, the circumscription of that genus might therefore require some adjustment at a later stage.Nevertheless, based on the criteria given above.95 species are transferred here from Ruschia to Antimima.
Plants compact, caespitose or with compact centre and long shoots, isophyllous or heterophyllous.Leaves: cymbiform with convex sides to nearly Finger-shaped, often keeled in upper part; or mostly oval and very often connate for at least part of the length, leaf pairs differing in this feature, mucronate; epidermis cells Hat to papil late.wax cover smooth or papillae, with a smooth later al channel and wax flakes or platelets on rest of surface.Flowers: l(-3 ).rarely in well-developed cymes; bracts and bracteoles present; petals pink to purple, rarely white, often arranged in 5 (rarely 6) groups.Filamentous staminodes: mostly in a central cone surrounding sta mens closely.Capsules: 5. rarely 6 locules, stalked, always with persisting bracteoles.top with high or low valve rims, base mostly bowl-shaped, sometimes funnelshaped; covering membranes undulate, more rarely straight, mostly elevated in centre, in several species with radial or tangential wings or other protrusions on them: closing body large, rarely small, absent in one species; expanding keels broad, diverging and radial in distal part, reaching to tip of valve; valves with or with out mostly narrow valve wings.Plants forming flat mats, isophyllous.Leaves: form ing white bodies in the resting state.Capsule: top almost flat, base short, funnel-shaped, closing body round, white, rather small for genus; covering membranes con vex with a low distal rim. a closing ledge below it.bases of expanding keels just not touching each other at their bases, moderately diverging and tapering into short awns.Typical of species: leaves of a pair form small, brightly white bodies in the dry state.Plants compact, heterophyllous.Leaves: persisting yellow, papery sheath of one leaf pair around stem, free parts triquetrous, other leaf pair connate for 2-3 mm only, ending in recurved long awn; keels of leaves with distinct, large, long papillae.Capsules: typical of genus.
Plants caespitose, heterophyllous.Leaves: sheath of one leaf pair as a connate body with only short free tips during the resting period, whitish yellow, smooth, with a protruding roll along line where both leaves are fused in dry state; second leaf pair hardly connate, with elevations above subhypodermal idioblasts and a papillate epider mis.Capsules: typical of genus, valve wings narrow.17.A ntim im a crassifolia (L.Bolus) H.E.K.Hart mann, comb.nov.Mesembryanthemum and allied genera, part 3: 338 (1958).Type: Hall BOL25758 (BOL, holo.!).

Ruschia deflecta L Bolus ex Jacobsen: 491 ( 1970) orthographic variant
Plants cushion-forming, heterophyllous.Leaves: yel low sheaths of one leaf pair on short shoots connate for ± half their length, those on long shoots only connate for ± '/ft of their length, free parts of both types with distant, big papillae.Capsules: typical of genus, but closing bodies hood-shaped and broader than thick.A deviating single fruit on the type sheet with broad valve wings and a longer base belongs to Amphibolia, but not to the remain ing type material, which agrees with the description.
Plants compact to caespitose, isophyllous.Leaves: trigonous, mucronate.Capsules: typical of genus.Typi cal of species: fruits raised above plant and the thick root.
Plants compact with erect, long shoots, heterophyl lous.Leaves: sheath of one leaf pair long, almost smooth, with a bulge running down side where both leaves meet, and long, ± triquetrous free parts with some large, long papillae along keel only; other leaf pair only slightly con nate, free parts densely papillate.Capsules: typical of genus.Typical of species: stout, dark brown or maroon, long internodes (Figure II).
Plants com pact, highly branched, isophyllous.Leaves: trigonous but keeled, sides convex to straight, whitish grey.Capsules: typical ot genus; covering mem branes almost straight.The species can easily be con fused with A. turneriana.Characteristic of A. dualis are the strictly compact growth, retained also in cultivation, and the strictly solitary flowers (Figure 1A -D).Semi-compact dwarf shrublets, isophyllous.Leaves: triquetrous with long papillae.Capsules: typical of genus with narrow valve wings at the point where they touch edge of valve and with oblique wings on top of covering membranes, 6-loculed (Figure 1H).Dinter (1928: 86, 87)  Plants densely shrubby, isophyllous.Leaves: terete, apically rounded.Capsules: typical of genus; covering membranes straight; valve wings very narrow.Resembles species of Rhinephyllum in growth form, both occur in the same area, so that they can easily be confused.
Plants compact, white in the resting slate, heterophyl lous.Leaves: sheath of one leaf pair forming white body in dry season, smooth, keel markedly smooth, margins ciliate, completely embracing subsequent leaf pair with longer free parts with short papillae on them.Capsules: with closing body pointing into locule like a finger; expanding keels very similar to the short ones of A dualis.Plants with compact centre and long shoots, each internode with papery whitish apical rim, appearing corky helow and drying into rings, stems therefore resembling cones put one into the other, heterophyllous.
Leaves: sheath of one leaf pair turning papery for most of its length (± 6 mm) in the resting state, free apical part of ± 2 mm with very low papillae; second leaf pair en veloped during the resting period, ± 6-8 mm long, tri quetrous, epapillate; in both leaf types keel and margins rounded.Capsules: typical of genus.Compact shrubs forming tiny tree-like shapes, hetero phyllous.Leaves: sheath of one leaf pair long, smooth, white, embracing subsequent leaves at bases, these latter free for ± 7\ their length and withering in a peculiar way: hardened bundles along keel (and into the apical tooth) and margins form a triangular window frame and persist much longer than intermediate tissue so that finally tri angular openings are formed.Capsules: typical of genus; highly undulate covering membranes with very broad, horizontally arranged wings on top almost completely hiding covering membranes when seen from above (Figure 1G).Typical of species: windowed leaves drying white, with frames remaining on plants.Plants with compact centre with long shoots, erect at first, decumbent later, rooting at nodes some distance from the primary centre and developing a secondary com pact centre, heterophyllous.Leaves: sheaths of both types of leaves papery, white, disintegrating early; bases of sec ond leaf pair almost separated by a triangle of stem tissue, with low papillae, old leaves closely set on short shoots.Capsules: typical of genus.Typical of species: formation of secondary compact centres with adventitious roots.Plants with decumbent branches, heterophyllous.

Antimima fergusoniae
Leaves: sheath of one leaf pair connate for ± 72 its length, free parts papillate, mucro to 1 mm long; second leaf pair connate for 76 of its length, free parts papillate, longer papillae along keel and margins, these leaves forming repeatedly on fresh long shoots.Capsules: unknown.Flat shrubs smelling of fish when damaged, covered in spines derived from big dichasial inflorescences, iso phyllous.Leaves: trigonous.Capsules: with high valve rims extending over edge of capsule, otherwise with typ ical features.Plants always densely covered with spines, making it easy to recognise them.37. Antimima h erm (Schwantes) H.E.K.Hart mann, comb.nov.Plants compact, heterophyllous.Leaves: sheath of one leaf pair papery for most of its length; other leaf pair free (flowering plant on photograph showing free leaves only), as is the case during the growing and flowering period.Capsules: typical of genus; covering membranes with low protrusions in radial direction near centre of fruit.Placed by Schwantes (1928a: 302)
Plants with flat mats, heterophyllous.Leaves: sheath of one leaf pair smooth, white in its lower half, often suf fused purplish, tips disintegrating into thread-like protu berances, sheath enclosing subsequent leaf pair of papil late leaves connate only for ± lh of its length; surface of epidermis of outer leaf pair of elongate cells with cuticular folds covered by an almost continuous wax layer.Capsules: typical of genus, but top with low rims, 6loculed.Typical of species: laciniate upper parts of high ly connate leaf pairs persisting on plant for many years.In this feature, and in leaf surfaces, it resembles A. pygmaea most closely, differing in the truncate leat tips and in fruit morphology from the latter.Plants compact with long decumbent shoots, numer ous short shoots on them, heterophyllous.Leaves: sheath of one leaf pair whitish, keel of shorter free part continu ing as a row of long papillae on sheath and internode below; alternating leaf pair with a sheath only 1.5 mm long, free parts subulate.Capsules: typical of genus; expanding keels merging into expanding sheets.Similar in leaf shapes and sequence to A. fergusoniae.Plants compact, isophyllous.Leaves: triangular to tri quetrous, light grey, hard.Capsules: most similar to those of A. dualis, but closing body smaller and almost hook-shaped.Grows far from most other species of genus, settling in distinct, stony habitats on limestone and seemingly unable to explore adjacent areas with a sand cover where grass and trees predominate.Mesembryanthemum leucantherum L.Bolus in Annals of the Bolus Herbarium 4: 7 (1925).Ruschia leucanthera (L.Bolus) L. Bolus: 239 (1927b).Type: Tugwell BOL17WR (BOL, lecto.!, here designated); Russell NBGH/24 (BOL, paralecto.!).
Plants compact to caespitose, isophyllous.hypocotyl and roots thickened.Leaves: trigonous, mucronate.Cap sules: typical of genus, interior falling off persistent old outer bundles of capsule, leaving a prickly star on plant where a fruit has been; in young fruits narrow valve wings in angle where expanding keel meets edge of valve.In habitat, plants adorned by whitish capsules for most of year.Similar to A. dekenahi, but differing in wings on cover ing membranes and longer papillae along keel and mar gins of leaves.Plants with compact centre with long shoots and high ly branched, tufted short shoots, bearing flowers (and fruits) in groups, heterophyllous.Leaves: smooth, papery sheath of one leaf pair of ± 5 mm length enclosing base of subsequent leaf pair with sheath of ± 4 mm and free parts up to 15 mm long; epidermis cells of free parts almost flat, only slightly protruding.Capsules: typical of genus; covering membranes raised high near centre and with indications of radial protrusions, but these appear as if folded and not forming a wing proper.Typical of species: dense groups of short shoots developing on long shoots.

55.
Antimima menniei (L.Compact shrublets, heterophyllous with 3 different types of leaves.Leaves: most commonly sheaths w hitish yellow with reddish brown free tips; epidermis at tips with rounded, low papillae; leaves up to 8 mm long; at bases of side branches first leaf pair with a white-yellow sheath, but without any different free tips, ± 5 mm long; second leaf pair hardly connate, free parts with promi nent, long papillae giving leaves a rough appearance, ± 5 mm long.Capsules: very hard, differing from typical form by possession of small finger-shaped closing bod ies.Typical of species: formation of at least three differ ent leaf forms and development of four new branches under terminal fruit, in contrast to common pattern of two or single branch.Low shrublets, heterophyllous.Leaves: set densely, keels cartilaginous.Capsule: not known.(Tischer) H.E.K. Hartmann, comb. nov. Cheiridopsis minima Tischer in Succulenta 9: 145 (1927).Type: Anon.s.n.(HEID) Plants com pact, isophyllous.Leaves: trigonous.Capsules: typical of genus.Description of fruit permits the conclusion that the species belongs to Antimima, where in several species, centrally raised covering membranes occur combined with deep-set, white, flat closing bodies.Dark shrublets, outer long shoots decumbent, hetero phyllous.Leaves: long sheath of one leaf pair turning white, basally maroon, free parts 1-2 mm long, papillate; second leaf pair hardly connate, free parts 3 -4 mm long, papillate and ciliate along margins, latter leaves on long shoots, sheaths always at bases of short side shoots, on which both leaf types alternate regularly.Capsules: typi cal of genus.Typical of species: dark appearance as a result of dark stems which are well visible because leaves are very short.

Antimima minima
Distinguished from A. distans from the same area by much more slender stems and a broad cushion shape with age.The first description appeared in June 1928.being validly published at that time; the Latin description appeared in August 1928 (Bolus 1928c), but was super fluous at that time.Plants compact with long, erect, stiff shoots, isophyl lous.Leaves: boat-shaped with convex sides.Capsules: typical of genus.The name should be used for stout plants with few to some flowers per inflorescence from a restricted area north of the Gariep River in Namibia.
Plants compact with decumbent long shoots, hetero phyllous.Leaves: sheath of one leaf pair smooth, later white, thin, papery, keel markedly horny, ciliate to serru late; second leaf pair connate for 1-2 mm only, free parts ± 10 mm long, with distinct awn borne apically, with low papillae, keel ciliate to serrulate, wax in dense layers of vertical platelets on free parts.Capsules: typical of genus; expanding keels merging gradually into expand ing sheets.
The drawing of Duncanson (K!) shows a plant in the growing season with several long shoots (with prominent internodes) with shortly connate, spreading leaves, resembling those of long shoots of A. mathewsii very closely.Compact shrublets, heterophyllous.Leaves: sheath of one leaf pair white, ± 4 mm long, tree tips ± 3 mm long, covered by fine papillae, longer ones on margins; second leaf pair with a 1-2 mm long sheath, free parts ± 6 mm long, papillate, the latter on long shoots, short shoots beginning with a sheathed leaf pair, later both types alter nating regularly.Capsules: typical of genus.Smallerleaved version of A. peersii.(Schwantes)  Plants compact with long, erect, or decumbent shoots with dark purple to blackish internodes, several to many short shoots developing on distal nodes, arrangement resem bling m istletoe on a branch, heterophyllous.Leaves: one leaf pair with a wide, yellow sheath with only short free parts, ± 6 mm long overall; other pair united for ± '/4 of its length, trigonous, obtuse, up to 12 mm long; epidermis of free parts papillate.Capsules: typical of genus.Typical of species: blackish internodes and crowded short shoots bearing flowers.66. Antimima oviformis (L.Bolus) H.E.K.Hart mann, comb.nov.Mesembnanthemum and allied genera, part 2: 261 (1931).Type: Herre SUG93II (BOL.holo.').
Plants compact with bundles of erect long shoots, het erophyllous.Leaves: white, smooth sheath of one leaf pair connate for at least '/2 its length; second pair connate for ± 3 mm, free parts elongate-trigonous with rather large, dense papillae, with a dense cover of wax platelets transgrading into threads, underlying continuous wax breaking eventually into big plates.Capsules: without closing bodies, but with typical broad expanding keels and narrow valve wings broadest at middle, both features typical of genus.68.Antimima paucifolia (L.Bolus) H.E.K.Hart mann, comb.nov.
Plants compact with long shoots, heterophyllous.Leaves: sheath of one leaf pair white, often with a cop per-coloured tinge at its upper end.± 8 mm long, free parts ± 10 mm long; subsequent leaf pair only a little connate with ± 20 mm long free parts appressed to each other during resting state, long shoots with these latter leaves only, short shoots starting with sheathed leaf pair, leaf forms alternating later: epidermis of free parts papil late.Capsules: typical of genus; narrow valve wings and medium-sized closing bodies.Plants with erect, stiff branches with proleptic short sh<x)ts from a rather dense, caespitose centre.Capsules: typical of genus with 6 locules.
Plants compact with long shoots, heterophyllous.Leaves: sheaths o f one leaf pair ± 4 mm long and free parts ± 2 mm long, these dominant on long shoots; other leaf pair connate for ± 2 mm, free parts ± 4 mm long, all free parts with elevations and papillae, on short shoots, types o f leaf pairs alternating regularly, longer leaves persisting over several years.Capsules: typical of genus; closing bodies medium-sized.
The name Ruschia persistens was used for two differ ent species by Bolus (1932Bolus ( , 1963)), the latter being an illegitimate name in Ruschia.However, the epithet is available in the genus Antimima.Shrubs with many scars from old leaves, isophyllous.Leaves: subfalcate to subclavate.Capsules: typical of genus; valve wings in their basal halves broader than expanding keels.
The broad, tapering valve wings, resembling those of Eberlanzia schneideriana (A.Berger) H.E.K.Hartmann, are unusual in the genus.The species is placed here based on the broad, typical closing bodies and the undu late covering membranes with inconspicuous closing ledges only.
Plants com pact with long shoots.Capsules: incom pletely described; covering membranes touching closing body; expanding keel denticulate, but these features are typical of genus.
Plants compact with very long, thin, yellow long shoots in other bushes, heterophyllous.Leaves: one type of leaf pair forming a papery sheath-like cover of two parts ± 10 mm long, connate only for ± 1 mm, apical part papillate, subulate; second leaf pair ± 8-10 mm long, tri quetrous, subulate, papillate, mucro recurved.Capsules: typical of genus.Typical of species: pseudo-sheaths formed by one leaf pair consisting of almost free leaves, but turning papery for dry season like connate sheaths in other species.
Plants compact, heterophyllous.Leaves: sheath of one leaf pair connate; second leaf pair almost free, 5 -6 mm long, free parts keeled, margins and keel finely ciliate, all glaucous-green.Capsules: not known.Plants creeping with proleptic side branches as tiny lateral bodies, later forming a mat, heterophyllous.
Leaves: sheath of one leaf pair long, yellowish, rather smooth, enclosing larger part of subsequent leaf pair with a shorter sheath and longer free parts with papillae, in particular along margins and keel, often two or more leaf pairs of second type following each other, ± 3 mm long, body ± 2.5 mm diam.Capsules: typical of genus.Small-leaved version of A. brevicarpa sharing similar character states with that species, in particular the same type of fruit and ecology.Ruschia levynsiae (L Bolus) Schwantes: 58 (1949).Mesembryanthe mum levynsiae L Bolus: 256 (1927b).Type: Levyns NBG1373/26 (BOL, holo.!).
Plants com pact, heterophyllous.Leaves: smooth sheath on one leaf pair with short, spreading, papillate, rough tips; second leaf pair enclosed which unfolds dur ing rainy season showing two almost oval, spreading leaves, connate slighty at bases and exhibiting upper sur faces, a rare condition in compact Ruschioideae; epider mis papillate.Capsules: typical of genus.
The difference in appearence between the resting state, when the yellowish green leaf pair forming a round body with the free tips protruding, and the growing phase is striking-in season, the bright green leaf pair spreads so completely that the persistent sheaths arc hardly visi ble.This difference is the reason why Bolus described the species twice within two years, having seen Ruschia pumila in the resting state only, but Ruschia levynsiae growing, developing the described heterophylly (Figure IE).Bolus (1913: 150) had described M. pumilum, circa 5 cm altum', using the adjective as a descriptive word only, not as a formal epitheton to name a species, as she stated herself in 1929, when describing Ruschia pumila.M eanwhile, Fedde & Schuster (1918) had taken M esembryanthemum pum ilum L.Bolus as a validly described name, which it became only through their cita-tion of the basionym and the type, hence the authorship nearly smooth; margins and keel turning white with age, 'L.Bolus ex Fedde & Schuster'.keel continuing in a fold or narrow wing down sheath and stem below; second leaf pair w ith longer free parts with medium long papillae.Capsules: typical of genus; expand-79.Antimima pusilla (Schwantes)  Plants compact, heterophyllous.Leaves', very similar in so far as they both develop long, smooth sheaths, one leaf pair with low distant papillae on short free parts (this one persisting during dry season enclosing next leaf pair); other with distinctly longer papillae thus looking rougher, papillae elongated along margins.Capsules: with expanding keels merging into expanding sheets and both together covering lower half of valve (when open); closing body small with a hollow underneath into which placenta extends.The heterophylly may be overlooked easily, but the fact that during the resting time one leaf pair encloses the subsequent one indicates the different roles the two leaf pairs play.80. Antimima pygmaea (Haw.)H.E.K. Hartmann. comb. nov. Mesembryanthemum pvgmaeum Haw. in Supplementum plantarum succulentarum: 98 (1819).Ruschia pygmaea (Haw.)Schwantes: 92 (1928b).Type: drawing no.996/204 by Duncanson (K.lecto.!. here designated).
Plants caespitose, forming mats, heterophyllous.Leaves: sheath of one leaf pair developing into a conical body enveloping subsequent leaf pair, upper part disinte grating into a cracked cover through which the following leaf pair grows when rain starts to fall, tips of both leaf types pointed and triangular as seen from above; epider mis very smooth, covered by a thin rugose layer of wax.Capsules: typical of genus; very narrow valve wings, 6loculed.Typical of species: conical white bodies formed by leaf pairs in resting period becoming cracked with time; laciniate old leaves occur also in A. ivori.but those are truncate and appear like a trimmed edge.81.Antimima quarzitica (Dinter) H.E.K.Hart mann, comb.nov.Dinter in Feddes Repertonum 19: 151 (1923).Ruschia quarzitica (Dinter) Dinter & Schwantes: 69 (1929).Corpuscularia quarzitica (Dinter) Schwantes: 186 (1926).Type Dinter 3866 (B.holo.!).
Plants compact with long shoots, heterophyllous.Leaves: white, papery sheath of one leaf pair 3-4 mm long, free parts 4-5 mm long, with low papillae; other leaf pair united for ± 2 mm only, free parts up to 10 mm long, with long papillae on keel and margins, low papil lae of different sizes on sides, wax cover dense, consist ing of flakes often connected.Capsules: unknown.The heterophylly had not been recognized in the orig inal description, nor the differences in papillae.In both features, the species resembles A. mathewsii, which has thicker and more robust stems but is similar in the irreg ular branching pattern.Plants caespitose, heterophyllous.Leaves: sheath of one Low shrubs, heterophyllous.Leaves: sheaths of one leaf pair long, whitish with rather short free tips; epidermis leaf pair 4-5 mm long, free parts 1-2 mm long, papillae low; other leaf pair with sheath ± 4 mm long, free parts up to 9 mm long, on long shoots at each node in one sea son.Capsules: unknown.87.A ntim im a sobrina (N .E .B r) H .E. K.Hart mann, comb. nov. Mesembryanthemum sobrinum N.E.Br. in Kew Bulletin 1929: 60 (1929).Ruschia sobrina (N.E.Br.)Schwantes: 230 (1938).Type: Muir 4014 (K, holo.).
Plants compact to forming big mats, isophyllous.Leaves: finger-shaped to terete.Capsules: typical of genus; only dorsal bundles of fruit present, interior hav ing fallen out as a tumble fruit, tiny valve wings some times present.Resembles Argyroderma fissum (Haw.)L.Bolus in its decumbent growth and finger-shaped leaves; both species grow sympatrically and can easily be confused in their vegetative state.Since fruits remain on the plants in both species, a distinction in habitat is easier.Plants compact, heterophyllous.Leaves: long sheath in one type of leaf pair, free parts with some short, dis persed papillae; other leaf pair with short sheaths and long, papillate free parts.Capsules: with very narrow valve wings and small closing bodies.The type looks very similar to that of A. verruculosa, differing mainly in the diameter of the flower, a feature known to change during anthesis.Ruschia subtruncata L.Bolus var.minor L. Bolus: 286 (1954).Type: Acocks 17185 (BOL, holo.!).

89.
Plants compact with long shoots twining in other bushes, heterophyllous; side branches with scars and somewhat spiny remains of closely set leaf pairs.Leaves: one leaf pair with papery protective cover of two parts ± 10 mm long, connate for only ± 2 mm, apical part papil late; second leaf pair enveloped during resting period, ± 6-10 mm long, free parts triquetrous, papillate, with some bigger papillae along margins, mucro recurved.Capsules: typical of genus; expanding sheets rising to form keels.Typical of species: pseudo-sheaths formed by one leaf pair consisting of almost free leaves, but turning papery for the dry season like connate sheaths in other species; these sheaths are about as long as broad in A. subtruncata.Sim ilar in general appearence to A. emarcescens and A. stayneri, both with slender leaves.92.Antimima triquetra (L.Bolus) H.E.K.Hart mann, comb.nov.
Plants compact with long shoots, erect at first, later decumbent, heterophyllous.Leaves: one type of leal pair with a papery protective cover of two parts ± 8 mm long, connate for only ± 2 mm, apical part subulate, with low papillae; second leaf pair ± 6-8 mm long, triquetrous, with long papillae along keel and margins, mucro erect.Capsules: typical of genus.Typical of species: pseudo sheaths formed by one leaf pair consisting of almost free leaves, but turning papery for the dry season like connate sheaths in other species-in this feature, the species resembles A. subtruncata.Plants compact in habitat, developing some long shoots in cultivation, isophyllous.Leaves: triquetrous with convex sides.Capsules: typical of genus; radial pro trusions ± wing-shaped, disintegrating later, forming rows of long teeth; valve wings at first as broad as expanding keels, disintegrating later quickly.Similar to A. dualis, both compact in habitat, but the latter never changing shape.95.Antimima vanzvlii (L.Bolus) H.E.K.Hart mann, comb.nov.Mesembnanthemum and allied genera, part 2: 209 (1930).Type: Van Zyl in Fuller 94 (BOL.holo.!).
Plants compact, heterophyllous.Leaves: one type of leaf pair with a long sheath connate for almost half its length; second with a free part; free parts trigo nous and papillate in both types, but more densely so in younger, less connate leaves, 40-80(-120) mm long.13-17 mm broad, 10-13 mm diam.Capsules: without valve wings, expanding keels distant.4-6-loeuled, in completely known.The low number of locules excludes the species from Cheiridopsis.where it might otherwise belong.Low shrublets.heterophyllous.Leaves: sheath of one leaf pair long, smooth with a short, papillate, free part; other leaf pair with a short sheath and longer papillae.Capsules: with a small closing body, partly destroyed.The 'type plan f is very similar to A. stokoei, which has larger flowers.Ruschia obtusifoha L. Bolus: 298 (1962) Type: Hall 2467 (BOL, holo.!).
Plants caespitose, isophyllous.Leaves: trigonous to roundish.Capsules: typical of genus; extremely narrow valve wings at areas where expanding keel meets edge of valve.
The type material of all species included here is very similar to that of A. klaverensis in growth pattern, leaf shape, and fruit morphology, differing only in the absence of the fishy smell reported for the latter species.Plants with erect, thin branches from a denser centre, often in other bushes.Leaves: triquetrous, hardly con nate.pair at its base leaving a triangle of stem visible between them.Capsule: base elongate, funnel-to bell shaped.top almost semi-orbicular from high valve rims; covering membranes forming a roof, ridge being the con tact line, closing ledges sharp and long: closing body nearly hook-shaped, hollow; expanding keels distant at bases.Material matching the collection of Bortenschlag, which was seen by Haworth, possesses fruits of the Ruschia type, with deep locules and small, hook-shaped closing bodies.The species is therefore placed back in Ruschia.
FIGURE 1.-A-D , Antimima dualis: A, l.s.through flower showing flat ovary surface and free stigmata, in contrast to Argyroderma; B, flowering branch with persisting old leaves below; C, part of open capsule with big closing bodies (CB), no valve wings, expanding keels (EK) diverging moderately and covering membranes (CM), with distinct closing ledge (CL) below near opening; D, side view of fruiting branch of herbarium material.A, B, figs of Brown 1930; C, Hartmann & Ihlenfeldt 4157 (HBG); D, Peers NBG 370/33 (BOL!), holotype of A. villetii.E, A. pumila, Hartmann 30325 (HBG!), side view of fruiting heterophyllous branch: long connate sheath is papery and smooth, short erect free parts papillate.F, A. dolomitica, Dinter 3782 (BOL, iso.!), radial l.s.through capsule in middle of locule: straight covering membrane (CM) lying distally on big closing body (CB), which is also touched by closing ledge (CL).G, A. fenestrata, Hartmann < 5c Dehn 15626 (HBG!), radial l.s.through capsule in middle of locule: covering membranes sloping down from a high centre ending in a distal recurving pressed against big white closing body (CB); erect wing in nearly radial position stabilizing covering membrane.H, A. eendomensis, Dinter 5207 (B, holo!), part of open capsule with basally distant moderately diverging expanding keels (EK), small valve wings (VW) broadest in middle, very big closing bodies (CB) and covering membranes (CM) pressed against them; covering membranes with radial wings on top lowering towards centre I, A. distans, Hartmann et al. 20682: heterophyllous long shoot with several leaf pairs with short connate sheaths and free long parts (FLP) and connate subterminal leaf pair (CLP) forming long sheath with short free parts enclosing terminal leaf pair.ES, expanding sheet; P, placenta; V, valve; VR, valve rim; W, wing on top of cover ing membrane.

Plants caespitose. heterophyllous. Leaves: both pairs forming sheaths, one pair with pronounced bulges above tannin idioblasts and low papillae; second pair with lower bulges but longer papillae. Capsules: typical of genus. After the First description in English in September 1928 (Bolus 1928b), Bolus published a Latin description of the species as well in November 1928 (Bolus 1928d: 7), where she cited the type. Nevertheless, the descrip tion in September fulfills all requirements for the new combination to be validly published, at that time.
R gracillima L Bolus in Notes on Mesembrianthemum and allied genera, part 2: 203 (1930) Type: Frames BOL 19208 (BOL, holo.!).
H.E.K.Hart-jng tjssueand short at base of valve, rising to inner mann, comb.nov.side,but not forming a distinct expanding keel.
Antim im a stayneri (L.Bolus) H.E.K.Hart  mann, comb.nov.Shrubs with stout stems, heterophyllous.Leaves: papery protective cover of two parts derived from one type of leaf pair, these leaves elongate, free, to 10 mm long, tips slender, pointed, dark when dry; second pair almost free, finely papillate, ± 8 mm long.Capsules: typ ical of genus.In leaves and stems A. stayneri resembles A. emarcescens and A. subtruncata, but the latter species grow as climbers in other bushes.