Plinthus Fenzl

Plinthus Fenzl, Nov. Stirp. Mus. Vindob. Dec.: 51 (1839); Ver-doorn, Bothalia 4: 177-179 (1941); Adamson, J. S. Afr. Bot. 27:147-151 (1961).

1. Gunniopsis Pax in Engler&Prantl

Gunniopsis Pax in Engler&Prantl, Nat. Pfl. fam. Ill, lb; 44

(1894); Chinnock, J. Adel. Bot. Gard. 6:133-179 (1983). A iz ooraL. subg. Gunniopsis(Pax&Hoffmann) Adamson (1959).

Herbs to woody shrubs. Leaves opposite, linear-terete, flat or oblanceolate, papillate to smooth, with multiseriate hairs ending in a conspicuous apical cell.

Procumbent perennial subshrubs, younger plants with scale-hairs. Leaves tiny, imbricate. Inflorescence of 1-3 sessile flowers; flowers perigynous; 4-5-merous; 4-5 alternisepalous stamens; ovary 2-4 locules, with a pendulous ovule each, styles short or wanting. Fruits with reduced expanding tissue. Seeds flat, reniform. About four spp., Namibia and South Africa. In need of revision.

Galenia L., Sp. PI.: 359 (1753): Adamson, J. S. Afr. Bot. 22: 87127 (1956).

Perennial shrublets, rarely herbs, mostly branch-shed-ders, erect, often with secund branches. Leaves small, mostly obovate. Inflorescence often clearly separated from a well developed vegetative base; perigynous flowers; the distal ones often with adnate bracts and bracteoles; 4-5-merous; stamens twice the number of perianth lobes, in alternisepalous pairs; ovary 2-5 locules with a singular pendulous ovule each. Fruits weakly hygrochastic or xerochastic. About 15 spp. in two subgenera distinguished by differences in the opening of the fruit, Namibia and Western part of South Africa. It seems promising to re-consider the reestablishment of two genera from Galenia.

6. Acrosanthes Ecklon & Zeyher

Acrosanthes Ecklon & Zeyher, Enum.: 328 (1837); Adamson, R.S., J. S. Afr. Bot. 25:23-28 (1959).

Prostrate to sprawling perennial, branching dichasial, but one branch predominant. Leaves opposite, glabrous. Inflorescences on suppressed dichasial branches, therefore seemingly lateral, probably on a relatively well developed vegetative base; perigynous flowers with 5 perianth lobes with dorsal appendages; 8 to many stamens sometimes in groups; ovary weakly inferior, divided incompletely by a basal septum into two locules with a basal ovule each. Fruit a xerochastic capsule persisting in the perianth tube. Seeds reniform to orbicular, testa rugose to tubercu-late. Four spp., southwestern part of Africa. The genus urgently needs revision based on more material than hitherto.

7. Sesuvium L.

Sesuvium L„ Syst. nat., ed. 10,2 Vegetabilia: 1058 (1759); Bogle, J. Arnold Arbor. 51: 431-462 (1970).

Branches rarely rooting. Leaves exstipulate but with membranes at the base of the petiole. Flowers with 2 basal bracteoles; perianth-lobes with long unifacial dorsal appendages; stamens 5 to many. Fruit 2-5-lo-cular, + conical top. About 12 spp., worldwide in the tropics and subtropics.

8. Cypselea Turpin

Cypselea Turpin, Ann. Mus. Hist. Nat. Paris 7: 219, pi. 12, Figs. 10-11 (1806); Bogle, J. Arnold Arbor. 51:431-462 (1970).

Annual procumbent herb forming mats. Leaves small, stipules scarious, laciniate. Flowers solitary with 2 basal bracteoles; perianth lobes without dorsal appendages; stamens 1-5; locules 2(-3). Septum in ripe fruit incomplete; fruit a subglobose, thin walled capsule. One sp., West Indies, S North America and N South America.

9. Trianthema L.

Trianthema L., Sp. PI.: 223 (1753); Bogle L., J. Arnold Arbor.

Leaves with membranes at the base of the petioles; surface + glabrous, papillose or hairy. Flowers in clusters with 2 bracteoles each; perianth markedly connate; stamens 5-°°; ovary with a single style, 1 carpel, ovules 2 to many, operculum undivided with 1-2 seeds inside. About 17 spp., worldwide in the tropics and subtropics.

II. Subfam. Sesuvioideae Lindley (1853)

Leaves often stipulate; flowers bracteate, perigynous; perianth internally petaloid, externally sepaloid, perianth segments mostly apiculate on the back towards the apex; capsule circumscissile; seed completely covered by an aril; x = 8. Four genera, worldwide in the tropics and subtropics.

Key to the Genera of the Sesuvioideae

1. Style 1 9. Trianthema

2. Septa complete in ripe fruits; seeds brown to blackish 3

- Septa incomplete or absent in ripe fruits; seeds light-coloured

  1. Cypselea
  2. Operculum of capsule breaking into 2 parts; 4 seeds per fruit

10. Zaleya

- Operculum of capsule in one piece; numerous seeds per fruit

  1. Sesuvium
  2. Zaleya Burm. f.

Zaleya Burm. f., Fl. Ind.: 110, t. 31 (1768); Jeffrey, Kew Bull. 14: 235-238 (1960).

Prostrate to ascending perennials. Leaves with a membranaceous sheath at the base of the petiole. Flowers in clusters of 5-20, sessile; perianth connate for 1/2 length; stamens 5-15. Fruit with 2 locules; 2 to many ovules, 2 seeds in each locule, operculum splitting septicidal sooner or later, each unit containing one seed. About six spp., NE and E Africa, India, Australia.

DDL Subfam. Tetragonioideae Lindley (1853)

Inflorescence often andromonoecious; perianth peta-loid, externally sepaloid; flower (semi-)epigynous; fruit mostly horn-shaped or a winged nut; seeds one per locule, with a short funicle; x = 8. Two genera.

Key to the Genera of the Tetragonioideae

I. Fruit derived from a single ovary, winged, ridged or horned, perianth with free or only basally connate lobes


- Fruit compound, derived from a complete inflorescence including the bracts and bracteoles; spiny; perianth connate for ± half its length 12. Tribulocarpus

II. Tetragonia L.

Tetragonia L., Sp. PL: 480 (1753); Adamson, J. S. Afr. Bot. 21: 109-154 (1955).

Erect to prostrate, rarely with a tuber and annual shoots. Leaves often rhomboid. Flowers 1 to many, mostly axillary, rarely terminal; perianth free, rarely connate at base. Fruit a nut, consisting of a single ovary. About 85 spp., in three to four subgenera, tropics and subtropics of the S hemisphere with a centre in S Africa, also in N America. One species grown as a leafy vegetable in suitable conditions.

12. Tribulocarpus S. Moore

Tribulocarpus S. Moore, J. Bot. 59: 228 (1921); Adamson, I. S. Afr. Bot. 21: 109-154 (1955); Verdcourt, Kew Bull. 12: 348349 (1957).

Low shrublets. Flowers bracteolate in terminal dense groups; perianth connate for ± 1/2 length. Fruit compound, spiny, stalked. One sp., Namibia and Ethiopia. The genus is separated from Tetragonia because of the compound nut fruit including the sclerotinised bracts, but the horns of some species of Tetragonia may be of comparable origin.

IV. Subfam. Mesembyranthemoideae Ihlenfeldt, Schwantes & Straka (1962)

Aptenioideae Schwantes ex Bittrich & H. E. K. Hartmann

Leaves ephemeral to persistent; with or without bladder cells; the inflorescence mostly very rich ± representing the entire plant, rarely reduced to solitary flowers; bracts and bracteoles mostly frondose, rarely hypsophyllous; calyx (4-5) and petals each mostly basally connate into tubes; nectaries 4-5, koilomorphic; petals often transgrading into filamentous stami-nodes and further into stamens; ovary semi-inferior to inferior, septate palcentation central; fruits with 4-5 locules, Mesembryanthemum-type, loculicidal hygrochastic capsule with septal expanding keels extending from the columella to the tip of the valve, valves opening often into a vertical position only. Nine genera, mainly in S Africa, N Africa and the Mediterrranean into Asia as far as Iran and Iraq, west coast of N and S America, Australia; widely introduced into suitable climates, e.g. S of England and the Channel Islands, often escaped.

Key to the Genera of the mesembryanthemoideae

  1. Plants compact, intemodes completely enclosed by the leaves and invisible, leaves fat-digitiform, alternate
  2. Dactylopsis

- Plants shrubby, herbaceous or straggling, leaves flat or cylindrical, if the latter, much smaller than above 2

2. Leaves opposite and basally free, but with tubular sheaths enclosing each other, seeds large (> 2 mm), flowers always solitary and large (0 to 45 mm) 17. Aspazoma

- Leaves alternate, if opposite, not free and not sheathing each other, seeds smaller (< 2 mm), flowers in dichasia, if solitary, smaller 3

3. Plants with articulate assimilating stems with a persisting succulent primary cortex 4

- Plants with a continuous stem, only the youngest stages green, primary cortex not persistent 7

4. Seeds 1.3-1.5 mm long, blackish-brown 16. Aptenia

- Seeds small, 0.6-1.3 mm long (if longer, enclosed'in a nut) 5

5. Stems and leaves with similar, ± reduced bladder cells, seeds less than 1 mm long, placenta between the bases of the expanding keels; inner staminodes scarious 19. Psilocaulon

- Stems with densely arranged xeromorphic bladder cells, leaves with distant ± mesomorphic ones, seeds 1-1.5 mm long, placenta not dividing the expanding keels or expanding tissue absent; inner staminodes not scarious 6

6. Fruit a woody nut 2L Pseudobrownanthus

- Fruit a hygrochastic capsule 20. Brownanthus

7. Seeds small, 0.5-1 mm long, plants annual or somewhat longer-lived 8

- Seeds medium to large, 1.2-2 mm long, plants perennial sub-shrubs or shrubs 15. Phyllobolus

  1. Leaves only opposite, basal ones united broadly into a funnel or tube, epidermis with finely pleated cuticle
  2. Synaptophyttum

- Leaves alternate or opposite, if so, never united into a dish or tube, but possibly connate, epidermis with a smooth cuticle

  1. Mesembryanthemum
  2. Mesembryanthemum L.

Mesembryanthemum L., Sp. PI.: 480 (1753), nom. cons.; Bittrich, Mitt. Inst. Allg. Bot. Hamb. 21: 5-116 (1987); Hart-mann&Bittrich, Bothalia20:153-157 (1990). Cryophytum N.E.Br. (1925). Derenbergiella Schwantes (1928). Eurystigma L. Bolus (1930). Halenbergia Dinter (1937). Hydrodea N. E. Br. (1925). Opophytum N.E.Br. (1925).

Lush erect, ascending or procumbent annuals, often facultatively perennial, often rosette. Leaves flat, rarely terete, mosdy only basally opposite, often with undulate margins; bladder cells uniform on stems and leaves, often elongated into hairs. Inflorescences very rich, rarely solitary; flowers often large; calyx 5-lobed; petals white, pink or yellow, narrow; nectaries 5; pollen punctuate to reticulate, smooth to microspinulose. Fruit 5-locular. Seeds smooth or papillate. About 25 spp. in four subgenera. Western and central S Africa, Namibia to S Angola; also in N Africa and in the Mediterranean, California and along the western coast of South America and in Australia. Widely introduced in suitable climates.

14.Synaptophyllum N.E.Br.

Synaptophylium N.E.Br., Gardeners' Chronicle III, 78: 412 (1925).

Procumbent to ascending annual herb, representing an inflorescence. Leaves spherical or ovoid, basally connate forming a dish or tubular sheath, upper leaves free; bladder-cells reduced, confined mainly to margins. Flowers in dichasia; calyx 4-lobed, unequal; androecial elements free. Fruit 4-locular, Mesembryanthemum-type. Seeds 0.6 mm long, very smooth, white. One sp., Southwestern part of Namibia.

15. Phyllobolus N.E.Br.

Phyllobolus N.E.Br., Gardeners' Chronicle III, 78: 413 (1925); Bittrich, Mitt. Inst. Allg. Bot. Hamb. 21: 5-116 (1987); Hart-mann&Bittrich, Bothalia 20:153-157 (1990). Amoebophyllum N.E. Br. (1925). Aridaria N.E.Br. (1925).

Nycteranthus Necker ex Rothmaler (1941) nom. illeg. Prenia N.E.Br. (1925). Sceletium N.E.Br. (1925). Sphalmanthus N.E.Br. (1925).

Procumbent to erect shrublets or subshrubs; often with thickened roots; sometimes geophytes with annual shoots; stems often thickened, succulent-soft or woody. Leaves often deciduous, alternate or opposite throughout or inflorescence with alternate leaves; bladder cells on leaves and stems uniform, prominent or reduced. Flowers solitary or in + rich inflorescences; petals white, yellow, greenish, purple, or red; pollen punctate to punctate-foveolate, sometimes + rugulose. Fruit sometimes without valve wings. Seeds D-shaped or horseshoe-shaped, brown, testa cells in rows, ± papillate. About 25 spp., in five subgenera, mainly distinguished by characters of leaves and seeds. S Namibia, western and central S Africa.

Aptenia N.E.Br., Gardeners' Chronicle III, 78: 412 (1925);

Bittrich, Mitt. Inst. Allg. Bot. Hamb. 21: 5-116 (1987). Platythyra N.E.Br. (1925). Tetracoilanthus Rappa & Cammarone (1953).

Erect, sometimes procumbent or climbing shrubs with green succulent cortex and slightly xeromorphic bladder cells in the epidermis of the stems. Leaves rarely terete; without a distinct central water tissue, epidermis with distant + low bladder cells, sometimes alternate in the inflorescences. Flowers sometimes solitary; calyx basally connate; nectaries 4; petals white, pink, cream to yellowish; pollen sometimes rugulose. Fruit 4-locular, with or without valve wings. Seeds D-shaped, papillae in rows, blackish brown, with epicuticular fine sculptures. About four spp., naturally in the summer rainfall areas of Namibia and South Africa, but worldwide used as an ornamental in suitable climates.

Yl.Aspazoma N.E.Br.

Aspazoma N.E.Br., Gardeners' Chronicle III, 78: 413 (1925).

Low shrubs with styloids in medullae of stems, stem epidermis with slightly xeromorphic bladder-cells. Leaves ± terete, basally free but enveloping each other forming a prominent sheath; chlorenchyma with water cells, epidermis with distant, low bladder cells. Flowers solitary; petals white to pale yellow; pollen punctate-foveolate, microspinulose. Locules mostly with 2 seeds only. Seeds 2.2 mm long, light brown, almost smooth, epicuticular sculpture very finely verrucose. One sp., western part of S Africa.

Dactylopsis N. E. Br., Gardeners' Chronicle III, 78:413 (1925).

Compact perennial; stems soft with slightly xeromorphic bladder cells. Leaves alternate, fat, finger-shaped with long sheaths completely enveloping the internodes; epidermis with distant, ± flat reduced bladder-cells. Flowers solitary, white, continuously open; calyx equally 5-lobed; petals stiff; nectaries 5; pollen punctuate to punctuate-foveolate, microspinulose. Fruit 5-locular. Seeds completely smooth, brown. One to two spp., western part of S Africa.

Psilocaulon N.E.Br., Gardeners' Chronicle III, 78: 433 (1925); Ihlenfeldt & Bittrich, Bot. Jahrb. Syst. 105: 289-322 (1985).

Procumbent to erect herbs, subshrubs or shrubs (0-3 m); stems articulate with a persistent, green succulent cortex. Leaves opposite, sometimes alternate in inflorescences, terete, semi-terete or triangular, sometimes mucronate, ephemeral; without a central water tissue, epidermis of leaves and stems similar, bladder cells sometimes elongate or ± reduced, sto-mata sometimes sunken. Flowers in dichasial inflorescences, rarely solitary; petals white, yellow, pink, or purple, rarely reduced, basally not or little connate with the stamens; inner staminodes scarious. Locules often very deep. Seeds 0.6-1 mm long, testa smooth or papillate. About 12 spp., S and W South Africa, S Namibia to S Angola.

20. Brownanthus Schwantes

Brownanthus Schwantes, Z. Sukkulentenk. 3: 14 (1927); Bittrich, Mitt. Inst. Allg. Bot. Hamb. 21:5-116 (1987).

Densely branched shrubs with xeromorphic epidermal cells. Leaves ± terete, ephemerous, sometimes spinescent; bladder cells at base and edges often elongated into hairs. Flowers solitary or in dichasia, white or cream; calyx lobes ± equal, stiff during an-thesis; petals basally free; pollen punctuate micro-spinulose. Fruit sometimes with a basal seed pocket, expanding keels not separated by placenta. Seeds 11.3 nlm long, D- or pear-shaped, + smooth, hardly any fine sculpture. About eight spp., western part of Namibia, S Africa and S Angola.

21. Pseudobrownanthus Ihlenf. & Bittrich

Pseudobrownanthus Ihlenf.&Bittrich, Bot. Jahrb. Syst. 105: 319 (1985); Bittrich, Mitt. Inst. Allg. Bot. Hamb. 21: 5-116 (1987).

Dense shrubs with stems like Psilocaubn but with slightly xeromorphic bladder cells. Leaves basally free. Flowers in dichasia; calyx 3-5-lobed; petals white, stiff. Fruit 3-4-locular, woody nut, expanding keels reduced, connate with the fruit, one seed per locule. Seeds ovoid, length = 1.5 mm, almost smooth, light brown. One sp., SW part of S Africa. Very closely allied to Brownanthus and perhaps better united with that genus.

V. Subfam. Ruschioideae Schwantes in Ihlenf., Schwantes &Straka emend. Bittrich &Hartmann (1988).

Leaves often sheathing; epidermis with bladder cells or uniformly xeromorphic; inflorescence mostly distinct; petals mostly free; filaments papillose at the base or very rarely naked; nectaries nearly always crest-like, free or connate into a ring; ovary inferior, placenta basal or parietal; fruit very frequently a hy-grochastic capsule with opening keels on the valves and extending into the locule, most often with covering membranes and additional closing devices; x = 9. Genera 106, S Africa, E to NE Africa, S Arabia, coasts of the Mediterranean, California, Australia.

Groupings within the subfamily Ruschioideae are tentative and presented here as a base to promote further critical studies.

1. Apatesia Group

Annuals to perennials. Leaves ± flat, only some bladder cells along the leaf margins. Flowers with a flat broad nectariferous ring. Capsules with reduced expanding tissue; often breaking into mericarps; often with seed pockets derived from false septa, rarely nuts. Genera 22-28.

  1. Cleretum Group
  2. Leaves flat, with prominent bladder cells. Flowers with a flat, broad nectary ring. Capsules with or without covering membranes; with or without spongy sills (as closing bodies) at the outer rim of the distal opening; expanding sheets prédominent, their extension related directly to the size of the covering membranes. Genera 29-31.
  3. Mitrophyllum Group
  4. Mostly heterophyllous, mostly two leaf-pairs per season, rarely anisophyllous; with prominent bladder cells. Flowers solitary, with a nectary ring, rarely reduced. Capsules with complete or reduced covering membranes qf constant shape; expanding sheets merging into expanding keels, rarely distinct; with or without spongy closing sills or bodies; with broad rectangular valve wings. Genera 32-41.
  5. Delosperma Group
  6. Leaves terete to trigonous, rarely flat, with ± prominent bladder cells and/or hairs. Flowers with five separate nectaries; with filamentous staminodes, rarely without. Capsules with or without covering membranes; expandingkeels and sheets separated, rarely merging; closing bodies absent; valve wings present, broad, rectangular; often with basal seed pockets. Genera 42-51.
  7. Stomatium Group

Plants perennial, rhizomatous. Leaves two to three pairs to a branch; homogeneous xeromorphic, crystals in irregular zones, surface rough or warty, from elevations above subhypodermal tannin idioblasts, often denticulate. Flowers with ftve separate nectaries. Capsule with or without covering membranes; without closing bodies; valve wings broad to reduced. Genera 52-60.

6. Utanopsis Group

Plants perennial; with a thickened hypocotyl and fleshy tap root, rarely with fibrous roots, but often with a fleshy rhizome. Leaves in rosettes or only one pair per branch; surface homogenous xeromorphic, crystal layers in certain areas often prédominent, smooth or warty. Flowers solitary with a nectary ring. Capsules without covering membranes or with them, these bending later into the empty locule along a performed line; with broad valve wings tapering towards the tip of the valve; without or with very small closing bodies. Genera 61-70.

7. Dracophilus Group

Plants perennial. Leaves xeromorphic with distinct differences in cell sizes, surface rarely sticky. Flowers mostly solitary; with a nectary ring, rarely five separate glands; without, rarely with filamentous staminodes. Capsules mostly 5-locular; with or without covering membranes; without closing bodies; with broad valve wings. Genera 71-81.

S. Bergeranthus Group

Plants perennial. Leaves homogeneous, xeromorphic, ± free. Rowers mostly in exserted inflorescences; with five separate nectaries. Capsules mostly 5-locular; with firm, straight, complete covering membranes; closing bodies present or absent; valve wings reduced. Genera 82-88.

9. Lampranthus Group

Plants perennial. Leaves free; homogeneous, mesomorphic to xeromorphic, crystal layer thin to absent Flowers with five separate nectaries, sometimes united into a ring. Capsule mostly 5-locular, deep; with firm complete covering membranes with additional closing devices at their distal recurved ends; opening blocked by sterile funicles; fruit rarely a berry. Genera 89-101.

10. Ruschia Group

Plants perennial. Leaves free to connate; surfaces homogeneous, xeromorphic, crystal layer thick. Flowers with filamentous staminodes and a nectary ring. Capsules with firm, complete, ± convex covering membranes with closing rodlets at their distal recurving; with closing bodies up to medium size; without, more rarely with valve wings; rarely nutlets. Genera 102-112.

1L Leipoldtia Group

Plants perennial. Leaves free to connate, homogeneous, xeromorphic, crystal layer of fine granules, often as sand. Flowers with or without filamentous staminodes; with a lophomorphic nectary ring. Capsules mostly multilocular; with firm, ± concave covering membranes with closing ledges, bulges or rodlets at the ± prominent distal recurving, rarely without; large, rarely smaller closing bodies. Genera 113-127.

Key to the Genera of Subfamily Ruschioideae.

1. Fruit a fleshy berry 91. Carpobrotus

2. Fruit a nut, finally disintegrating into nutlets or breaking into mericarps with previous loculicidal opening 3

- Fruit a loculicidal capsule 5

3. Fruit woody, dispersal unit nut or nutlets 4

- Fruit leathery, breaking into one-seeded mericarps as dispersal units 26. Hymenogytte

4. Fruit > 10 mm in 0, finally breaking completely into mericarps; leaves flat 28. Caryotophora

- Fruit < 7 mm in 0; the sclerotinised bundles of the fruit remaining on the plant; leaves ± club-shaped to falcate

  1. Ruschianthemum
  2. Capsule xerochastic, expanding tissue absent 6

- Capsule hygrochastic, expanding tissue present 8

6. Locules 5 7

- Locules > 8 25. Conicosia

7. Flat base of bottom of fruit widened, containing seeds, formed like a ring-shaped pocket, septa remaining connate with valves 24. Saphesia

Fruits with separate seed pockets at half the height of the capsule, septa free from valves 23. Skiatophytum

Expanding tissue reduced, consisting of expanding sheets or expanding keels only 9

Expanding tissue of expanding sheets and keels 12

Expanding keels only 10

Expanding sheets only 11

Locules open, seeds visible, seed pockets at outer base of locules; pedicels glabrous; leaf margins with bladder cells only 22. Apatesia

Locules covered by septa bending over them, seeds hidden, placenta raised, but no seed pockets; pedicel and calyx with long hairs; leaf margins with long hairs 27. Carpanthea Leaves lyrate 31. Aethephyllum

Leaves entire 30. Dorotheanthus

Capsule opening only once, expanding keels breaking after first opening 13

Capsule opening repeatedly, expanding keels remaining functional 14

Locules 5, valves free to base, valve wings broad, hard, erect; seeds wind-dispersed 103. Stoeberia

Locules 8, valves connate laterally at base, valve wings absent; seeds rain-dispersed 107. Stayneria Covering membranes present 15 Covering membranes absent (sometimes narrow rims) 16 Covering membranes of constant shape 44 Covering membranes declining into empty locules 45 Expanding keels distinctly separated from expanding sheets, keels parallel, high, ± sharply pleated 19 Expanding keels merging into expanding sheets (if expanding keels very low, see 11), keels widely diverging 17 Heterophyllous shrubs 34. Mitrophyllum Isophyllous plants 18 Annual herbs 29. Cleretum Perennial plants 42. Delosperma Calyx-like leaves sticky, sand adhering to it

74 Psammophora

Calyx-like leaves not sticky 20

Valve wings absent or as awns 21

Valve wings present, rectangular 24

Capsule opening incompletely (valves at most erect); leaves ± club-shaped with apical windows 56. Frithia Capsule opening completely (valves spreading); leaves without apical windows 22

Pedicel very long (5-12 cm), flowers mostly ternate; leaves keeled, toothed; plants compact 85. Carruanthus

Pedicel shorter (-2.5 cm), flowers solitary; leaves not keeled, not toothed; plants shrubby, caespitose to decumbent or creeping 23 Valves with long ribbon-shaped appendages at tips; flowers opening midday; shrubs 77. Jensenobotrya Valves without appendages at tips; flowers opening at night; plants creeping or caespitose 57. Neohemicia Valve wings at most as broad as expanding keels 25 Valve win¡p broader than expanding keels 26 Plants creeping 55. Mossia Plants not creeping 106 Bracteoles p resent (different from foliage leaves) 33 Bracteoles absent (indistinguishable from foliage leaves) 27 Base of capsule with very long soft hairs; leaves hairy

50. Muiria

Base of capsule and leaves with bladder cells, glabrous, or velvety 28

28. Base of capsule and leaves with bladder cells; plants heterophyllous 36. Oophytum

- Base of capsule and leaves xeromorphic, glabrous or velvety; plants homophyllous 29

29. Leaves obliquely triquetrous 68. Sckwantesia

- Leaves isometric 30

30. Plants forming clumps above the ground 66. Dinteranthus

- Plants sunken in the ground 31

31. Predominant number of locules per fruit in a population 5-6 65. Lithops

- Predominant number of locules per fruit 7 and above 32

32. Seeds tuberculate, tubercles distant 66. Dinteranthus

- Seeds smooth to rugose 65. Lithops

33. Flowere in ± rich dichasia 34

- Flowers solitary 36

34. Complete plant representing the inflorescence; annual vegetative regeneration from base regular; leaves mesomorphic with bladder cells 42. Delosperma

- Inflorescence perennial, well raised above and separated from basal vegetative clump; no annual regeneration of parts; leaves xeromorphic 35

35. Flowers remaining open during anthesis, petals stiff

75 .Nelia

- Flowers closing over night, petals ± lax 42. Delosperma

36. Nectary as five separate glands 37

- Nectary as a lophomorphic ring 39

37 Flowers with a short hypanthium, leaves mostly with teeth

54. Stomatium

- Flowers with free elements from the top of the ovary; leaves always without teeth 38

38. Leaf surface fine velvety, whitish grey 49. Corpuscularia

- Leaf surface rough, granular, dark green with white dots

  1. Rhinephyllum
  2. Locules <7 41

40. Base of capsule and leaves finely papillate, pedicel persistent on fruit; plants above ground 73. Namibia

- Base of capsule and leaves glabrous, capsule breaking off, dispersal as tumble fruit; plants at least partly sunken in the ground

41. Petals and stamens united into a tube

- Petals and stamens free from the base

42. Leaves warty

- Leaves smooth or papillate

43. Leaves soft, mesomorphic

- Leaves ± hard, xeromorphic

64. Nananthus

81. Herreanthus 78. Berrisfordia

  1. Ophthalmophyllum 79. Conophytum
  2. Covering membranes without distal recurving and always without any additional closing devices like rodlets, ledges or bulges at their distal ends 45

- Covering membranes with a ± distinct distal recurving, with or without additional closing devices like rodlets, ledges or bulges; if straight, always with some additional closing device 91

45. Covering membranes declining into the empty locule along a preformed edge 82

- Covering membranes straight or only slightly and gradually declining into empty locules 46

46. Plants with distinct prominent bladder cells 47

- Plants glabrous or hairy, surfaces ± homogeneous 63

47. Leaves flat, petiolate; plants annual 48

- Leaves terete to trigonous or semiglobose; plants perennial 51

48. Leaves lyrate 49

- Leaves entire 50

Top of capsule flat, expanding sheets only 31. Aethephyllum

Top of capsule with ribs, expanding sheets merging into expanding keels 29. Cleretum Valve wings ± broad, rectangular 29. Cleretum Valve wings tapering into awns 30. Dorotheanthus Plants heterophyllous 52 Plants homophyllous 57 Expanding keels divergent, emerging gradually from expanding sheets 53 Expanding keels parallel, sheets separate from expanding keels 36. Oophytum Plants compact and low (height < 4 cm) 37. Diplosoma Plants shrubby, if compact, height > 5 cm 54 Basal parts of old leaf sheaths sclerotic, completely enveloping the stem 35. Monilaria Old leaf sheaths papery-pergamentaceous 55 Flowers basally enclosed by connate hypsophylls forming a cup 32. Dicrocaulon Flowers exserted above leaves, no cups 56 Both leaf pairs of a season connate for > 30 %

33. Meyerophytum

One leaf pair of a season connate for <25%, spreading, the other for > 50 %, erect 34. Mitrophyllum

Leaves with an apical diadem 44 Trichodiadema

Leaves without an apical diadem 58

Bladder cells on leaves thin-walled, collapsing when dry, surface glittering 59

Bladder cells thick-walled, constant in shape, surface velvety 62 Surfaces of primary internodes with prominent bladder cells 43. Drosanthemum Surfaces of primary internodes glabrous 60 Leaves fat-digitiform, length > 4 cm; 38. Jacobsenia Leaves subterete to trigonous, length < 3 cm 61

Locules 5 Locules S5 10 Locules 8 Locules 6

Leaves hairy-velvety Leaves glabrous Flowers with a hypanthium Flowers with free petals and stamens Plants compact to caespitose Plants shrubby Calyx 4-lobed Calyx > 5-lobed Leaves soft, mesomorphic Leaves ± hard, firm, xeromorphic Locules 5 Locules 3s 10

Closing body absent; shrubs Closing body bipartite; creepers Flowers yellow

Flowers white, pale pink, or purple Petals stiff Petals lax

Closing body present, covering membranes complete

109. Polymita Closing body absent, covering membranes incomplete

75. Nelia

. Top of capsule raised, old leaves dark brown 74

Top of capsule flat, old leaves parchment-like, whitish

  1. Delosperma
  2. Mestoklema
  3. Malephora 39. Drosanthemopsis
  4. Drosanthemum

64 66

51. Imitaria

48. Gibbaeum


70 69

47. Malephora

40. Disphyma

Leaves digitiform, more than three times as long as broad, 95. plants caespitose 92. Antegibbaeum

Leaves ± trigonous, with convex sides, plants compact 75 Valve wings narrow to absent 87. Cerochlamys

Valve wings broad 48. Gibbaeum

Closing bodies present, sometimes as sills 77 96.

Closing bodies absent 80

Leaves sharply triquetrous, acuminate 84. Bergeranthus 97. Leaves trigonous to subterete, hunched 78

Leaves markedly oblique and asymmetric, plant compact, 98. enclosing the short-stalked fruits 88. Bijlia

Leaves symmetrical, plants shrubby, if compact, fruits ex- 99. serted well above the leaves 79 -

Closing body bipartite, often resembling two plates 100.

83. Rhombophyllum Closing body single, small 82.Hereroa 101.

Valve wings broad (> expanding keels) -

53. Chasmatophyllum 102. Valve wings narrow or absent 82. Hereroa -

Flowers solitary, ebracteolate 89. Lamp ran thus 103.

Flowers in rich inflorescences, bracteolate 98

Valve wings absent, rims of valves high 112. Ebracteola -Valve wings present, at least basally very broad 83 104.

Top of capsule flat or conical, pedicel persistent, bracteolate 84 Top of capsule rounded-retuse (with a central depression), 105. pedicel often breaking above the insertion of the bracte-oles thus forming a tumble-fruit 62. A loinopsis 106. Closing body absent 86 -Closing body present, of endocarpal origin, but very small 107.

Base of capsule papillate; leaves papillate, keeled, toothed 108.

69 a. Ihlenfeldtia Base of capsule smooth; leaves smooth, not keeled but fat- 109. digitiform, blunt, never toothed 69. Tanquana

Locules 5-7, rarely in single fruits 8 87 110.

Locules 8 in all fruits 89

Locules very flat, covering membranes therefore held up by the funicles; flowers appearing after the leaf-pairs thus seemingly lateral 70. Didymaotus 111.

Locules deeper, covering membranes bending into empty -locule; flowers distinctly terminal 88 112.

Calyx lobes truncate and warty, like the leaves; upper leaf surface exposed, leaves never keeled, blunt, spathulate; 113. plants mostly sunken in the ground 61. Titanopsis

Calyx lobes trigonous to pointed, smooth, like the ± deltoid, keeled, gibbose leaves; plants clumpy above ground

67. Lapidaria 114. Capsule whitish grey, top ± flat or conical, base funnel-shaped; leaves broadly trigonous, grey, smooth 90

Capsule ± dark brown, top rounded to semiglobose; 115. leaves acuminate, dark green, with raised dots 58. Rabiea Locules 8 123. Octopoma

Locules 10-11 144 116.

Closing body absent 92

Closing body present, sometimes as a broad sill 109 117. Valve wings absent or narrow (< width of expanding keels) 99

Valve wings, at least at base, much broader than expanding keels 93 118.

Valve wings rectangular 95

Valve wings tapering into awns towards the tip of 119.

the valve 94

Capsules multilocular, plants compact 126. Pieiospilos Capsules 5-locular, if multilocular, plants shrubby

98. Erepsia

Valve wings in open capsules stiff, stretching over the locules and not moving outwards with the valves

59. Faucaria

Valve wings in open capsules flexible, moving outwards with the valve 96

Seeds echinate 90. Braunsia

Seeds with low papillae 97

Plants compact 60. Orthopterum

Plants shrubby, rarely caespitose or creeping 98

Filaments broadened and connate at base 100. Circandra Filaments filiform, free 89. Lampranthus

Leaves xeromorphic 102

Leaves mesomorphic 100

Leaves flat, petiolate; plants annual 27. Carpanthea Leaves terete to trigonous; plants perennial 101

Flowers yellow 101. Scopelogena

Flowers white to purple 46. Oscularia

Seeds echinate 94. Namaquanthus

Seeds smooth to moderately papillate 103

Surface of ovary (in flower) concave, forming a hypan-thium 98. Erepsia

Surface of ovary (in flower) flat to raised 104

Leaves connate into a sheath, transgrading into the cortex

Leaves ± free 106

Free parts of leaves < 1 cm long 96. Smicrostigma

Free parts of leaves 2-3 cm long 97. Zeuktophyllum Plants compact 76. Ruschianthus

Plants shrubby 107

Low, ± creeping shrub (-15 cm) 99. Esterhuysenia Higher, ± erect shrub (> 30 cm) 108

Flowers very bright purple 93. Enarganthe

Flowers white to pink 95. Wooleya

Closing body bipartite 110

Closing body not divided 111

Closing body u-shaped; leaves mesomorphic; plants creeping 40. Dispkyma Closing body of two ± flat plates; leaves xeromorphic; plants compact to shrubby 83. Rhombophyllum Leaves soft, mesomorphic 112 Leaves hard, xeromorphic 114 Plants annual, leaves flat 30. Dorotheanthus Plants perennial 113 Covering membranes ± straight; plants heterophyllous 34. Mitrophyllum

Covering membranes undulate; leaves anisophyllous in a pair 41. Glottiphyllum

Expanding keel clearly separate from expanding sheet 115

Expanding keel merging into expanding sheet 88. Bijlia Valve wings broad (at least at base) 116

Valve wings absent or narrow (< width of expanding keels) 127

Valve wings tapering from a broad base 125

Valve wings ± rectangular 117

Closing bodies not larger than seeds, leaving space at the sides 118

Closing bodies larger than seeds, ± blocking the distal. opening of the locule 120

Capsule pentamerous 102. Eberlanzia

Capsule multilocular 119

Capsules dispersed as tumble-fruits; leaves with apical window; plants under ground 122. Fenestraria Capsules persistent on pedicels; leaves fusiform without a window; plants above ground 121. Jordaaniella

120. Leaves with a sticky surface 108. Arenífera

- Leaves without a sticky surface 121

  1. Base of capsule velvety or rough from papillae
  2. Cheiridopsis

- Base of capsule glabrous 122

  1. First leaves of side branches connate > 50%
  2. Vanzijlia

- First leaves of side branches hardly connate like all others (seedlings occasionally with connate leaf pairs, however)

Capsules with closing bulges 114. Cephalophyllum

Capsules with closing rodlets 124

Top of capsule turret-shaped; procumbent branches from compact centre, not rooting 117. Hallianthus

Top of capsule rounded, but not turret-shaped; erect to ascending, or saltatory and rooting plants 116. Leipoldtia Flowers white 109. Polymita

Flowers yellow 126

Leaves sharply triquetrous, grey 86. Machairophyllum Leaves trigonous to semiovate, gibbose, dark green

126. Pleiospilos

Closing body large, completely blocking the locule 128 Closing body smaller, leaving some space at the sides 133






















Capsule pentamerous 129

Capsule multilocular 130

Flowers yellow 84. Bergeranthus

Flowers purple 113. A ntimima

Leaves glabrous 132

Leaves velvety, hairy or rough 131

Leaves and base of capsule velvety 120. Odontophorus Leaves and base of capsule rough 119. Cheiridopsis Flowers with a hypanthium, capsule 12-24-locular

  1. Argywderma Flowers without a hypanthium, capsule 8-locular
  2. Octopoma

Closing rodlets, ledges or bulges present 138

Additional closing devices absent 134

Leaves triquetrous-acuminate 136

Leaves trigonous to terete 135

Flowers white to cream, with ± 2000 elements

127. Cylindrophyllum Flowers yellow, with <300 elements 82.Hereroa

Flowers yellow 86. Machairophyllum

Flowers white to purple 137

Petals with a darker purplish longitudinal stripe, pedicels longer than leaves, locules 5 110. Acrodon

Petals uniformly coloured, white to purple, pedicels at most as long as leaves, locules 6-10 111. Khadta

Flowers very large, 0 > 8 cm, > 2000 elements

127. Cylindrophyllum Flowers smaller, 0 < 6 cm, < 100 elements 139

Closing rodlets present 140

Closing bulge or ledge present 125. Schlechteranthus Rims of valves very high 141

Rims of valves rather low 142

Seeds echinate, at least in parts 106. Astridia

Seeds smooth to papillate 105. Ruschia

Capsules 5-locular, very small, 0 2-4 mm 105. Ruschia Capsules 6-8-locular, bigger, 0 > 6 mm 143

Foliage leaves > 5 cm long 124. Ottosonderia

Foliage leaves < 4 cm long 123. Octopoma

Flowers yellow 63. Vanheerdea

Flowers pink to white 71. Dracophilus

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