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Bothalia 1 2,3 : 377-393 (1978)
New taxa and new combinations in Cotyledon and allied genera
H. R. TÓLKEN*
ABSTRACT
As a result of a re-evaluation of Cotyledon and Adromisehus, it was found necessary to amend the delimi
tation of Cotyledon and to describe a new genus Tylecodon. A summarized revision of Adromisehus with keys to
sections, species and subspecific taxa is provided. The following new names and new combinations are effected:
Adromisehus sect. Boreali Toelken, A. cristatus var. clavifolius (Haw.) Toelken,—var. schonlandii (Phill.)
Toelken,—var. zeyheri (Harv.) Toelken, A. fallax Toelken, A. fiticaulis subsp. marlothii (Schonl.) Toelken, A.
inamoenus Toelken, A. marianae var. hallii (P. C. Hutch.) Toelken,—var. kubusensis (Uitew.) Toelken, A.
schuldtianus subsp. juttae (V. Poelln.) Toelken, A. subviridis Toelken, A. umbraticola subsp. raniosa Toelken,
Tylecodon buchholzianus (Schuldt & Steph.) Toelken, T. cacalioides (L.f.) Toelken, T. decipiens Toelken, T.
fragilis (Dyer) Toelken, T. faucium (V. Poelln.) Toelken, T. grandiflorus (Burm.f.) Toelken, T. hallii (Toelken) Toel
ken, T. hirtifolium (W. F. Barker) Toelken, T. leucothrix (C. A. Smith) Toelken, T. occultans (Toelken) Toelken, T.
paniculatus (L.f.) Toelken, T. pearsonii (Schonl.) Toelken, T. pygmaeus (W. F. Barker) Toelken,—var. tenuis
(Toelken) Toelken, T. racemosus (Harv.) Toelken, T. reticulatus (L.f.) Toelken,—subsp. phyllopodium Toelken,
T. rubrovenosus (Dinter) Toelken, T. schaeferanus (Dinter) Toelken, T. similis (Toelken) Toelken, T. singularis
(Dyer) Toelken, T. striatus (P. C. Hutch.) Toelken, T. suffultus Bruyns ex Toelken, T. sulphureus (Toelken)
Toelken, T. torulosus Toelken, T. tuberosus Toelken, T. ventricosus (Burm.f.) Toelken, T. viridiflorus (Toelken)
Toelken, T. wallichii (Harv.) Toelken,—subsp. ecklonianus (Harv.) Toelken.
RÉSUMÉ
NOUVEAUX TAXA ET NOUVELLES COMBINAISONS DANS COTYLEDON ET GENRES APPARENTÉS
Suite á une réévaluation des genres Cotyledon et Adromisehus, il s'est avéré nécessaire de rectifier la dilimi
tation de Cotyledon et de décrire un genre nouveau: Tylecodon. Une revision abrégée ^’Adromisehus est fournie,
avec des clés pour les sections, espêces et taxa sub-spécifiques. Les noms et combinaisons nouveaux qui out été
établis sont les suivants: Adromisehus sect. Boreali Toelken, A. cristatus var. clavifolius (Haw.) Toelken,—var.
schonlandii (Phill.) Toelken,—var. zeyheri (Harv.) Toelken, A. fallax Toelken, A. filicaulis subsp. marlothii
(Schonl.) Toelken, A. inamoenus Toelken, A. marianae var. hallii (P. C. Hutch.) Toelken,—var. kubusensis
(Uitew.) Toelken, A. schuldtianus subsp. juttae (V. Poelln.) Toelken, A. subviridis Toelken, A. umbraticola
subsp. ramosa Toelken, Tylecodon buchholzianus (Schuldt & Steph.) Toelken, T. cacalioides (L.f) Toelken,
T. decipiens Toelken, T. fragilis (Dver) Toelken, T. faucium (V. Poelln.) Toelken, T. grandi^orus (Burm.f.) Toelken
T. hallii (Toelken) , T. hirtifolium (W.F. Barker) Toelken, T. leucothrix (C. A. Smith) Toelken, T. occultans
(Toelken) Toelken, T. paniculatus (L.f) Toelken, T. pearsonii (Schonl.) Toelken, T. pygmaeus (W. F. Barker)
Toelken,—var. tenuis (Toelken) Toelken, T. racemosus (Harv.) Toelken, T. reticulatus (L.f.) Toelken,—subsp.
phyllopodium Toelken, T. rubrovenosus (Dinter) Toelken, T. schaeferanus (Dinter) (Toelken), T. similis (Toelken)
Toelken, T. singularis (Dyer) Toelken, T. striatus (P. C. Hutch.) Toelken, T. suffultus Bruyns ex Toelken, T.
sulphureus (Toelken) Toelken, T. torulosus Toelken, T. tuberosus Toelken, T. ventricosus (Burm.f) Toelken,
T. viridiflorus (Toelken) Toelken, T. wallichii (Harv.) Toelken,—subsp. ecklonianus (Harv.) Toelken.
NOTES ON COTYLEDON, TYLECODON AND
ADROMISCHUS
Linnaeus included a number of different elements
in his genus Cotyledon and in time these were placed
into separate genera such as Kalanchoe Adans. (1763),
Umbilicus DC. (1801), Orostachys Fisch. (1808) and
Pistorina DC. (1828). In 1852 Lemaire described the
genus Adromisehus on the basis of its spike-like inflo
rescence and the long corolla tube. However, only
since 1930 when Berger used the name, has the genus
Adromisehus become generally accepted and the con
cept of the genus Cotyledon remained unchanged
except for a few species which were at first too little
known to be placed into their correct genus. Berger
accepted the genus Adromisehus, but followed Schon-
land (1915) in leaving the second group of species with
spirally arranged leaves within Cotyledon with no
rank at all. De Candolle (1828) recognised the fol
lowing three groups in Cotyledon: “(a) Foliis, op-
positis; (b) Foliis alternis, marcescentibus; (c) Foliis
alternis, persistentibus.” It is remarkable that he
conceived such a simple and profound system seeing
that he had knowledge of the much more complex
subdivision proposed by Salm-Dyck, but only
published in 1834. De Candolle’s groups agree well
with the proposed concepts of Cotyledon sensu strieto,
Tylecodon and Adromisehus as set out in Table 1 in
which the more important characteristics of the three
genera are summarized.
TABLE 1.—Comparison of the main characteristics of the genera Adromisehus, Cotyledon and Tylecodon
ADROMISCHUS COTYLEDON TYLECODON
Leaves
.........................
Leaf epidermis
............
Inflorescence
...............
Flowers
.......................
Penducle......................
Sepals, adaxial surface
Petals
..........................
Filaments
....................
Nectary scales
.............
spiral, stiff, persistent
....................
opposite, stiff, persistent
................
isodiametric cells with straight isodiametric cells with straight
anticlinal walls anticlinal walls
usually a spike-like thyrse
.............
thyrse with 1-6 dichasia, each en
ding in a monochasium
erect or spreading (pendulous in pendulous
A. phillipsiae)
with abrupt change from leaves to with abrupt change from leaves to
bracts bracts
with club-shaped trichomes........... glabrous
fused beyond the tube
..................
fused for less than the full length of
the tube
smooth or papillose
......................
usually hairy towards the base___
disc-like, free
................................. laterally fused to carpel to form a
cup
spiral, soft, herbaceous, deciduous,
elongate cells with sinuate anti
clinal walls,
thyrse with 1-8 monochasia.
erect or spreading (pendulous in
T. pearsonii).
with gradual transition from leaves
to bracts,
often with club-shaped trichomes.
fused for less than the full length
of the tube,
usually hairy towards the base,
disc-like, free.
* Botanical Research Institute, Department of Agricultural
Technical Services, Private Bag X101, Pretoria.
378 NEW TAXA AN D NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
The inflorescence in Cotyledon is a thyrse with one to reports by C. A. Smith (1939) who keyed out a
to several dichasia in which each branch terminates number of species of Adromischus by their opposite
in a monochasium with two bracts below each flower. leaves. The leaves are often so closely packed that the
The sympodial branch system from the second bract spiral arrangement becomes only clearly visible in
is suppressed. Species of Cotyledon have an opposite etiolated branches in which the internodes elongate
leaf arrangement, so that a dichasium similar to that somewhat. The question remains whether the two
in Crassula could be expected. bracts below each flower on the monochasia of
Tylecodon and Adromischus have spirally arranged Tylecodon and Adromischus are comparable to those
i i j ■ i .i
_
Í on the terminal monochasia of Cotyledon and thus
leaves and, accordingly, the thyrse develops a number whether fhese monoch asia are derived from dichasia
of monochasia spirally arranged along the central Í Sf • , ™°n0(:{Jas,a are derived from dichasia,
axis of the inflorescence and they reach maturity and ,he sP,ral from the °PPos,te leaf arrangement.
acropetally. This basic pattern, which is found for No direct affinities were found between any of the
instance in T. grandiflora, could theoretically undergo two genera and there appear to be about as many
two extreme developments: characters shared among any two of the genera as
(a) A basitonous development (Troll, 1964), which ‘here, a,re dissimilarities. Thus, while Tylecodon and
results in a shortening of the central axis of the Cotyledon agree in then karyotype (Ogawa, unpublish-
inflorescence together with a pronounced development ed) a"d >n their relatively short corolla tube, they differ
of the lateral part-inflorescences (monochasia) to in these respects from Adromischus The leaves are
produce a corymb-like thyrse as in T. cacalioides. persistent and their isodiametnc epidermis cells have
In many plants of T. wallichii the central axis is even st[a 'gh‘ anticlinal walls in Cotyledon and Adromischus,
further reduced, so that 2-4 monochasia seem to whde ln Tylecodon the leaves are deciduous and their
branch from the same node (pleiochasia in Troll, epidermis cells have anticlinal walls. The leaves are
1964), a condition which should not be confused spirally arrangedand the nectaries disc-like in Tyleco-
with the basic dichasial pattern of Cotyledon. don and Adromischus, while in Cotyledon the leaves
.are opposite and the nectaries cup-shaped. Not one
(b) An acrotonous development (Troll, 1964), in 0f genera can apparently be derived from one of
which the central axis elongates while the lateral the others. None of them represents merely a group
part-inflorescences are reduced to sessile flowers, or Gf species more highly developed than related ones in
almost so, and thus form a spike-like thyrse. Extreme regard to a particular character as was found to be
forms of this development lead to the single flower the case in groups like Rochea, Rhopalota, Pagella
per part-inflorescence in Adromischus, which can be or Globulea, which are now again included in the genus
deduced from the two bracts on the pedicel of each Crassula. Within each one of the three genera at
flower on the inflorescence. least one ultimate development in regard to a certain
A similar spike-like inflorescence is found in character such as pointed out in the above discussion
Crassula capitella subsp. capitella and subsp. nodulosa, of the inflorescence is found, but they do not merely
which belong to the sect. Rosulares of which other consist of species showing this highly developed
members such as C. vaginata and C. alba show a characteristic.
basitonous development. In Tylecodon, however, only three genera are obviously very similar to one
the latter development has produced extreme forms, another, but no indication of direct or close relation-
while in Adromischus the acrotonous development is ship between any two of them was found. It is sig-
predominant except for sect. Brevipedunculati in nificant, and particularly in the Crassulaceae where
which the thyrse may consist of 1-4 monochasia each numerous hybrids between genera have been produced
with one to eight flowers. (Uhl, personal communication), that no natural
The inflorescence alone cannot be used to separate intergeneric hybrids between any of the three genera
the genus Adromischus from Cotyledon and Tylecodon. have been recorded, although plants often grow near
All representatives of Tylecodon and Adromischus one another. Yet numerous natural hybrids between
investigated have spirally arranged leaves, contrary species of each of the genera are known.
KEY TO GENERA
Leaves deciduous, with elongate epidermis cells with sinuate anticlinal cell walls.............................2. Tylecodon
Leaves persistent with isodiametric cells with straight anticlinal cross walls:
Leaves opposite (or in whorls of 3); petals fused for less than the full length of tube, nectary cup
shaped ...............................................................................................................................1. Cotyledon
Leaves spirally arranged; petals fused beyond the tube; nectary disc-like
................................
3. Adromischus
1. COTYLEDON to red. Filaments hairy towards the base where they
Cotyledon L., Sp. PI. ed. 1: 429 (1753), pro parte; are ^used to the corolla tube. Squamae laterally
DC., Prodr. 3: 396 (1828), pro parte; Harv., FI. fused t0 the carPel t0 form a CUP in which the n?ctar
Cap.2:370 (1862), pro parte; Schonl. in Rec. Albany collects. Ovary with (50-) 100 or more ovules in at
Mus. 3:130(1915), pro parte; V. Poelln. in Reprium least two rows on each placenta,
nov. Spec. Regni veg. 42: 15 (1937); Friedr., in Prodr.
FI. S.W.Afr. 52: 5 (1968), pro parte; Dyer, Gen. 1: 195 2. TYLECODON
(1975), pro parte, emend. Type species: C. orbiculata L.
Shrubs, rarely suffrutices with somewhat woody Tylecodon Toelken, gen. nov. a Cotyledone foliis
branches usually with flaking bark, without tuberous spiralibus, deciduis et squamis disciformibus, ab
base. Leaves opposite, persistent, stiff, with smooth Adromischo folns deciduis quikus gradatim breve-
waxy surface. Inflorescence a thyrse with 1-6 dichasia scentibus bracteis in pedunculo differt.
each ending in a monochasium, usually with many Frutices vel plantae parvissimae vix altiores 30 mm,
pendulous flowers; bracts on peduncle few and much ramis succulentis rare cartilagineis et plerumque
shorter than leaves below. Calyx glabrous or some- cortice chartaceo, saepe tuberibus multiramosis. Folia
times with glandular hairs on abaxial surface. Corolla spiralia, decidua, molliter herbacea sine strato ceraceo
fused for less than the full length of the tube, glabrous et saepe epidermidis cellulosis protuberantibus. Inflo-
or rarely with glandular hairs on the outside, yellow rescent ia thyrsus 1-8 monochasiis quoque l-oo flori-
H. R. TOLKEN 379
bus erectis vel efíusis. Calyx glaber vel pilis glanduli-
feris trichomatibus clavatis supra et/vel subtus. Corolla
connata non per totam longitudinem tubi, glabra vel
pilis glanduliferis extus, viridis, fusca vel alba et saepe
suffusa rosea. Filamenta tomentosa ad basem. Squamae
librae disciformae flavae vel tangerinae. Ovarium 4-30
(-90) ovulis plerumque una serie in quoque placenta.
Type species: T. cacalioides (L.f.) Toelken.
Shrubs to very small plants scarcely higher than
30 mm with succulent branches rarely cartilaginous
and usually with peeling bark, often with much-
branched tubers. Leaves spirally arranged, deciduous,
soft herbaceous and without waxy layer and often
with bulging epidermis cells. Inflorescence a thyrse
with 1-8 monochasia each with one to many erect
or spreading flowers; bracts on peduncle becoming
gradually shorter from the leaves to the flowers.
Calyx glabrous or with glandular hairs or with club-
shaped trichomes on the abaxial and/or the adaxial
surface. Corolla fused for less than the full length of
the tube, glabrous or with glandular hairs on the
outside, green, brown or white and often tinged pink
rarely orange. Filaments hairy towards the base where
they are fused to the corolla tube. Squamae free and
disc-like, yellow to orange. Ovary with 4-30(-90)
ovules usually in one row on each placenta.
Occurring in an area between southern South West
Africa to south-western Cape and eastern parts of the
Little Karoo, but also with one outlier in the moun
tains around Graaff Reinet.
The name Tylecodon is an anagram of Cotyledon.
T. cacaliodes was selected as the type species as it
is the only species enumerated by De Candolle (1828)
of which the name has not been changed in one or
other way since that time.
T. buchholzianus (Schuldt & Steph.) Toelken,
comb. nov.
Cotyledon buchholziana Schuldt & Steph. in Kakteenkunde
(1937) 111, fig.; V. Poelln. in Kakteenkunde (1938) 111, fig.;
Friedr. in Prodr. FI. S. W. Afr. 52: 8 (1968); Jacobsen, Handb.
Succ. PI. 1: 279, fig. 274 (1960); Sukk. Lex. 132, t.38,2 (1970).
Type: Cape, 200 km from Port Nolloth, M. Schlechter s.n. (Bf).
T. cacalioides (L.f.) Toelken, comb. nov.
Cotyledon cacalioides L.f., Suppl. 242 (1781); Thunb., Prodr.
3: 397 (1800); FI. Cap. ed. Schultes 397 (1823); DC., Prodr. 3:
397 (1828); Harv., FI. Cap. 2: 274 (1862); Schonl. in Rec.
Albany Mus. 3: 147 (1915); V. Poelln. in Reprium nov. Spec.
Regni veg. 42: 18 (1937). Type: Cape, near Olifant's Bad,
Thunberg in Herb. Thunberg 10 998 (UPS, holo!).
T. decipiens Toelken, sp. nov. a T. schaeferano
ramis latis (6-10 mm in diametro), foliis cymbiformi-
bus et ovariis pilis paucibus; a T. toruloso corolla
plerumque glabra tubo 9-11 mm longo et lobis roseis,
foliis cymbiformibus differt.
Plantae perennes ramis multiramosis 6-10 mm in
diametro usque ad 0,2m longis sed plerumque brevis-
simis et tegetes densae formantes super basim tumidam
multiramosam, phyllopodiis torulosis, cortice ramium
veterium alba et confrigenti et caduco. Folia oblan-
ceolata vel elliptica 4-12 (-15) mm longa, 3-8 mm
lata, apicibus obtusis vel rotundatis, cuneata, cymbi-
formia, pilis glanduliferis paucis vel glabrescentia,
viridia vel griseo-viridia. Inflorescentia thyrsus 1,2(3)
monochasiis quoque 1,2 floribus sed 1 flore in natura;
pedunculus 8-20 mm longus, filiformis; pilis glanduli
feris; pedicellus 6-15 mm longus. Calycis lohi anguste
triangulares, 2 ,5-3,5 mm Iongi, acuti, pilis glandulo-
sis. Corolla tubo 9-11 mm longo, glabro, pallide flavo-
viride; lobi 4—5 mm longi, lanceolati, acuti, glabri,
rosei vel paene albi. Stamina 10—12 mm longa, antheris
aequalibus 1,2-1 ,4 mm longis, filamentis rectis pilis
tenuibus patentibus ubi connatis corollae tubo.
Squamae oblongae, 1 ,2 -1 ,4 xO ,3-0,4 mm, non con-
stnctae ad bases, emarginatae, leviter succulentae,
pallide flavae. Carpella ovariis gracilibus pilis patent
ibus et gradatim constrictis ad stylos erectos stig-
matibus terminalibus; ovarium 26-28 ovulis quoque
circiter duplo longiore quam lato leviter latis factis
ad funiculum et cristis verticalibus.
Type: Cape, near Grootmis, Tolken 5252 (PRE,
holo!).
Perennials with much-branched stems 6-10 mm
in diameter up and to 0,2 m long but usually very
short and forming mats so dense that the branches
can scarcely be distinguished from the much branched
swollen base, with leaf bases giving stems a slightly
knobby appearance, with bark on older branches
white and flaking. Leaves oblanceolate to elliptic,
4-12 (-15) mm long, 3-8 mm broad, with obtuse or
rounded apices, cuneate, cymbiform, covered with few
glandular hairs to glabrous, green to grey-green.
Inflorescence a thyrse with 1 or 2 (3) monochasia each
with one flower (but in cultivation with one or two
flowers); peduncles 8-20 mm long, thread-like, covered
with glandular hairs; pedicels 6-15 mm long. Calyx
lobes narrowly triangular 2,5-3,5 mm long, sharply
acute, covered with glandular hairs. Corolla with
tube 9-11 mm long, glabrous, pale yellowish-green;
lobes 4-5 mm long, lanceolate, acute, glabrous, pink
rarely almost white. Stamens 10-12 mm long, with
anthers equally long 1,3 -1,4 mm, with straight
filaments covered with fine spreading hairs where
fused to the petal tube. Squamae oblong, l,2 - l,4 x
0,3-0 ,4 mm, not constricted downwards, emarginate,
slightly fleshy, pale yellow. Carpels with slender
ovaries with spreading hairs and gradually constricted
into erect styles with terminal stigmas; ovary with 26-
28 ovules each about twice as long as broad and
slightly broadened at the funicle and with vertical
ridges.
In the field the branches do not elongate much
and, together with the branched swollen bases, they
form dense mats similar to those found in the form of
T. schaeferanus which was described as Cotyledon
sinus-alexandri, and with which the species has hitherto
been confused.
The confusion was only cleared up when, after
three years in cultivation, erect branches had
developed and flowers could be investigated.
T. fragilis {Dyer) Toelken, comb. nov.
Cotyledon fragilis Dyer in Flower. PI. Afr. 41, pi. 1631 (1971).
Type: Cape, near Strandfontein, Hall 3426 (PRE, holo!).
T. faucium ( V. Poelln.) Toelken, comb. nov.
Cotyledon faucium V. Poelln. in Reprium nov. Spec. Regni veg.
50: 323 (1941). Type: Cape, Verlatenkloof, Herre in SUG 6841
(BOL, clono!).
T. grandiflorus (Burm.f.) Toelken, comb. nov.
Cotyledon grandiflora Burm.f., Prodr. FI. Cap. 13 (1768);
Schonl. & Bak.f. in J. Bot., Lond. 40: 23 (1902); Schonl. in Rec.
Albany Mus. 3: 147 (1915); V. Poelln. in Reprium nov. Spec.
Regni veg. 42: 20 (1937); Adams., FI. Cape Penins. 439 (1950);
Kidd, Wild Flow. Cape Penins. pi. 2, 3 (1950). Iconotype: J.
Burm., Rar. Afr. PI. t.20, fig.l (1738).
T. hallii (Toelken) Toelken, comb. nov.
Cotyledon hallii Toelken in Bothalia 12: 193 (1977). Type:
Cape, De Hoop, Hall 1300 (NBG, holo!).
T. hirtifolium (fV. F. Barker) Toelken, comb. nov.
Cotyledon hirtifolium W. F. Barker in Flower. PI. Afr. 18, t.
690 (1938). Type: Cape, near Komaggas, Herre in BOL 22 165
(BOL, holo!).
T. leucothrix (C. A. Smith) Toelken, comb. nov.
Adromisehus leucothrix C. A. Smith in Bothalia 3: 637, pl.l
(April, 1939). Iconotype: Bothalia 3: 637, pi. 1 (lecto!).
Cotyledon swartbergensis V. Poelln. in Reprium nov. Spec.
Regni veg. 47: 1 (October, 1939). Type: Cape, Klein Swartberg,
Herre in SUG 6897 (BOL, clono!). C. leucothrix (C. A. Smith)
Fourcade in Mem. bot. Surv. S. Afr. 20: 34 (1941).
380 NEW TAXA AN D NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
C. A. Smith described the species only from vegeta
tive material and apparently did not make a specimen
as he had probably waited in vain for the plants to
flower. The only record of the original material is
the photograph published together with the original
description. This was therefore chosen as lectotype.
T. occultans (Toelken) Toelken, comb. nov.
Cotyledon occultans Toelken in Bothalia 12: 191 (1977). Type:
Cape, near Bitterfontein, Hall4289 (PRE, holo!; NBG!).
T. paniculatus (L.f.) Toelken, comb. nov.
Cotyledon paniculata L.f., Suppl. 242 (1781); Thunb., Prodr.
83 (1794); FI. Cap. ed. Schultes 396 (1823); Marl., FI. S. Afr. 2:
14, pi. 4 (1925); Schonl. in Rec. Albany Mus. 3: 147 (1915); V.
Poelln. in Reprium nov. Spec. Regni veg. 42: 21 (1937); Adam
son, FI. Cape Penins. 439 (1950); Henderson in Flower. PI. Afr.
1142 (1953); Friedr. in Prodr. FI. S. W. Afr. 52: 9 (1968);
Jacobsen, Handb. Succ. PI. 1: 288, fig. 285 (1960); Sukk. Lex.
134, t. 40.1 (1970). Type, near Hartekwas Kloof, Thunberg in
Herb. Thunb. 11 010 (UPS, holo; microfiche!).
T. pearsonii (Schonl.) Toelken, comb. nov.
Cotyledonpearsonii Schonl. in Ann. S. Afr. Mus. 9: 55 (1912);
in Rec. Albany Mus. 3: 148 (1915); V. Poelln. in Reprium nov.
Spec. Regni veg. 42: 22 (1937); Friedr. in Prodr. FI. S. W. Afr.
52: 9 (1968). Type: Cape, between Annenous and Chubiesis,
Pearson 5981 (GRA, lecto!). C. luteosquamata V. Poelln. in
Desert PI. Life 11: 65, fig. (1939); in Reprium nov. Spec. Regni
veg. 46: 78 (1939); Jacobsen, Sukk. Lex. 134, t. 39,2 (1970).
Type: Cape, Bushmanland, Triebner s.n. (B|).
T. pygmaeus (W. F. Barker) Toelken, comb. nov.
Cotyledon pygmaea W. F. Barker in Flower. PI. Afr. 10, pi.
396 (1930). Type: Cape, Vanrhynsdorp, Vigne in NBG 2267/29
(BOL, holo!).
var. tenuis (Toelken) Toelken, comb. nov.
Cotyledon pygmaea var. tenuis Toelken in Bothalia
12: 192 (1977). Type: Cape, near Holrivier Station,
Hall 3925 (PRE, holo!; NBG!).
T. racemosus (Harv.) Toelken, comb. nov.
Cotyledon racemosa Harv., FI. Cap. 2: 375 (1862), pro parte,
excl. specimen b; N.E.Br. in Gdnrs’ Chron. ser. 3, 51: 348
(1912); Schonl. in Rec. Albany Mus. 3: 149 (1915); V. Poelln. in
Reprium nov. Spec. Regni veg. 42: 22 (1937). Type: Cape,
between Kaus, Natvoet and Doornpoort, Drêge s.n. (S, lecto!;
BM!; K!). C. choroleuca Dinter ex Friedr., Mitt. Bot. Staats-
samml. Munchen 3: 597, fig. (1960); Prodr. FI. S.W.Afr. 52:
8 (1968). Type: South West Africa, 20 km N of Sendelingsdrift,
Herre in SUG 20 039 (M, holo!).
Drege’s specimen from near Verleptpram, which
E. Meyer named Cotyledon racemosa b, must be
identified as T. Iiallii. However, Harvey included this
collection in his citation of specimens under C. race
mosa. The specimen of C. racemosa a at Stockholm
Herbarium was thus chosen as lectotype.
T. reticulatus (L.f.) Toelken, comb. nov.
Cotyledon reticulata L.f., Suppl. 242 (1781); Thunb., Prodr.
83 (1794); FI. Cap. ed. Schultes 393 (1823); DC., Prodr. 3: 398
(1828); Harv., FI. Cap. 2: 376 (1862); Schonl. in Rec. Albany
Mus. 3: 148 (1915); Marl., FI. S. Afr. 2,1: 17, pi. 7 (1926); V.
Poelln. in Reprium nov. Spec. Regni veg. 42: 23 (1937); Jacob
sen, Handb. Succ. PI. 1: 290, fig. 287 (1960); Sukk. Lex. 135, t.
40,2 (1970). Type: Cape, beyond Hartequas Kloof, Thunberg in
Herb. Thunberg 11013 (UPS, holo; michrofiche!).
subsp. reticulatus.
Cotyledon reticulata L.f., Suppl. 242 (1781).
Young branches not less than 10 mm in diameter
but usually these are so short that they are difficult to
distinguish from the apex of the main stem which has
several growth points and is smooth or covered with
crescent-shaped leaf scars which disappear after one
year when the peeling bark is produced. Inflorescence
rigid, with flowers usually spreading; pedicels with
sessile or short glands.
Occurring on lower gravelly slopes in most parts of
Namaqualand as far north as the Port Nolloth area,
but also in Bushmanland, the Tanqua Karoo and the
southern parts of the Great Karoo as far east as
Willowmore and occasionally in the north-eastern
Little Karoo.
subsp. phyllopodium Toelken, subsp. nov.
Rami juvenes usque ad 6 mm in diametro et saepe
15 mm longiores, phyllopodiis tecti quoque folii
cicatricibus plus minusve rotundis et visibilibus annis
aliquot. Inflorescentia fragilis, floribus plus minusve
ascendibus; pedicellum pilis saltern duplo longioribus
quam apicibus glanduliferis latis.
Type: South West Africa, Numais, Dinter 8092
(BOL, holo!).
Young branches up to 6 mm in diameter and often
longer than 15 mm, covered with phyllopodia each
with a more or less round leaf scar and these remain
visible for several years. Inflorescence brittle, with
flowers more or less erect; glandular hairs on pedicels
at least twice as long as the diameter of their glandular
apex.
Found on rock outcrops or often on rock faces from
just south of Komaggas to south-western South
West Africa. Wherever the two subspecies occur
close together, as for instance near Komaggas, they
are ecologically well separated.
T. rubrovenosus (Dinter) Toelken, comb. nov.
Cotyledon rubrovenosa Dinter in Reprium nov. Spec. Regn.
veg. 30: 194 (1932); V. Poelln. in Reprium nov. Spec. Regni vegi
42: 24 (1937); Friedr. in Prodr. FI. S. W. Afr. 52: 10 (1968).
Type: South West Africa, south of Warmbad, Dinter s.n. (B,
holo!).
T. schaeferanus (Dinter) Toelken, comb. nov.
Cotyledon schaeferana Dinter in Reprium nov. Spec. Regni
veg. 19: 145 (1923); W. F. Barker in Flower. PI. Afr. 10, pi. 394
(1930); Friedr. in Prodr. FI. S. W. Afr. 52: 10 (1968); Jacobsen
in Sukk. Lex. 135, t. 40,3 (1970).
Type: South West Africa, Luderitz Bay, Dinter
4449 (B, t).
T. similis (Toelken), Toelken, comb. nov.
Cotyledon similis Toelken in Bothalia 12: 192 (1977). Type:
Cape, N of Grootmis, Wisura 1303 (NBG, holo!).
T. singularis (Dyer) Toelken, comb. nov.
Cotyledon singularis Dyer in Flower. PI. Afr. 41, pi. 1606
(1970). Type: South West Africa, near Rosh Pinah, Hardy 2632,
(PRE, holo!).
T. striatus (P. C. Hutch.) Toelken, comb. nov.
Cotyledon striata P. C. Hutch, in Cact. Succ. J., Los Ang. 36:
16, figs. (1964). Type: Cape, near Garies, Rodin 1405 (Univ.
Calif. Bot. Gard. 49. 1794-1: BOL. holo!; PRE!).
T. sufFultus Bruyns ex Toelken, sp. nov. a T.
reticulato ramis gracilibus laevibus (3-6 mm in dia
metro) et petalorum lobis roseis pilis patentibus intra;
a T. pygmaeo et T. simili pilis glanduliferis gracilis
in petalis et sepalis externis et pilis brevibus paucis
loborum partibus interioribus differt.
Plantae perennes ramis gracilibus procumbentibus
3-6 (-8) mm in diametro et usque ad 0,3 m longis,
radicibus tuberosis saepe ramosis. Folia lineari-
elliptica (8-) 10-20 (-25) mm longa, 2,5-4 ,5 mm lata,
acuta, plana vel canaliculata supra, convexissima
subtus, glabra atro-viridia. Inflorescentia thyrsus 3-5
monochasiis quoque 1-3 floribus, pedunculis rigidis
et pedicellis pilis glanduliferis. Calycis lobi anguste
triangulares, 4-5 mm longi, acuti pilis glanduliferis
paucis, succulenti, flavo-virides. Corolla tubo 5-6 mm
longo, flavo-viride, pilis glanduliferis extus; lobi 4-5
mm longi, lanceolati, acuti vel pungentes, piles glandu
liferis extus et pilis gracilibus eglanduliferis intra,
rosei. Stamina 5,5-6,5 mm longa, antheris gracilibus
inaequalibus circiter 1,3 mm et 1,6 mm longis,
filamentis rectis pilis tenuibus patentibus ubi connatis
corollae tubo. Squamae oblongae, 1-1 ,2 xc. 0,8 mm,
H. R. TÓLKEN 381
emarginatae, constrictae ad bases, flavae. Carpella
ovariis gracilibus gradatim ad stylos stigmatibus
terminalibus constrictis; ovarium 16-20 ovulis leviter
latis factis ad bases et cristis verticalibus.
Type: Cape, below Vanrhyn’s Pass, Bruyns 1091
(PRE, holo!).
Scrambler with slender procumbent branches 3-6
(-8) mm in diameter and up to 0 ,3 m long, with
tuberous roots often branched. Leaves linear-elliptic
(8-) 10-20(-25) mm long, 2,5-4,5 mm broad, acute,
flat or with longitudinal groove above, very convex
below, glabrous, dark green. Inflorescence a thyrse
with 3-5 monochasia each with 1-3 flowers, with
stiff peduncles and pedicles covered with glandular
hairs. Calyx lobes narrowly triangular, 4-5 mm long,
acute, with few scattered glandular hairs, fleshy,
yellowish-green. Corolla with tube 5-6 mm long,
with glandular hairs outside, yellowish-green; lobes
4-5 mm long, acute or drawn into a sharp point, with
glandular hairs outside and with fine eglandular hairs
inside, pink. Stamens 5 ,5-6,5 mm long, with slender
anthers unequal c. 1,3 mm and 1,6 mm long, with
straight filaments with a tuft of hairs where fused
to the corolla tube. Squamae oblong, 1-1,2x c. 0,8
mm, emarginate, slightly constricted towards the base,
yellow. Carpels with slender ovaries gradually con
stricted into the styles and with terminal stigmas;
ovary with 16-20 ovules slightly broadened towards
the base and with vertical ridges.
At present the plant is known only from one locality
but it may well be more wide-spread. As the species
scrambles among other succulent shrubs, it is almost
impossible to find unless in flower. The calyx seems
to form an abscission layer at its base, so that a loose
star-shaped disc remains on the pedicels of old inflo
rescences similar to those found in T. reticulatus.
T. sulphureus (Toelken) Toelken, comb. nov.
Cotyledonsulphurea Toelken in Bothalia 12: 191 (1977). Type:
Cape, near Pofadder, Tolken 3676A (BOL, holo!).
T. torulosus Toelken, sp. nov. a T. decipienti
ramis robustis (8—20 mm in diametro) foliis et petalis
dense tectis pilis glanduliferis; a T. viridifloro ramis
robustis torulosis et foliis ovato-spathulatis differt.
Plantae perennes ramis erectis multis 8-20 mm in
diametro et usque ad 0,3 m longis incrassatis ad
bases torulosis ob phyllopodia multa, cortice in ramis
juvenibus bruneo et veteribus albo. Folia ovato-
spathulata, 20-35(-40) mm longa, 5-15 (-21) mm
lata, apicibus obtusis vel rotundatis et recurvata,
cuneata vel petiolata, interdum canaliculata, pilis
glanduliferis dense tecta, griseo-viridia vel prasina.
Inflorescentia thyrsus 2-5 monochasiis quoque 1-3
floribus; pedunculus 5-8 mm longus, (1-) 2-3 mm
latus, pilis glanduliferis. Calycis lobi triangulares
2,5-3,5 mm longi, acuti vel pungentes, pilis glanduli
feris. Corolla tubo 18-23 mm longo pilis glanduliferis
praecipue in cristis inter petala flavo-viridi; lobi 4-5
mm longi, lanceolati, obtusi, pilis glanduliferis extus,
pallide flavi. Stamina 23-24 mm longa, antheris
aequalibus c. 1,8 mm longis, filamentis infra antheras
recurvatis et pilis gracilibus patentibus ad basim ubi
corollae tubo connatis. Squamae paene quadratae
1-1,2x0,9-1 mm, vix ad basim constrictae, emar
ginatae leviter succulentae, albae. Carpella ovariis gra
cilibus gradatim in stylos erectos stigmatibus ter
minalibus constrictis; ovarium 18-24 ovulis quoque
circiter duplo longiore quam lato leviter lato facto
ad funiculum et ciistis verticalibus.
Type: Cape, Karruchab Poort, Tolken 5317
(PRE, holo!).
Perennial with many erect branches 8-20 mm in
diameter and up to 0,3 m long becoming thicker
towards the base, with stems torulose due to the
numerous leaf bases, with bark on young branches
dark brown later white. Leaves ovate-spathulate,
20-35 (-40) mm long, 5—15(—21) mm broad, with
obtuse or rounded apices usually recurved, cuneate
to petiolate, sometimes longitudinally grooved, den
sely covered with glandular hairs, grey-green to
blue-green. Inflorescence a thyrse with 2-5 mono
chasia each with 1-3 flowers; peduncle 5-8 mm long,
(1—)2—3 mm thick, with glandular hairs; pedicels
2—5(—8) mm long, with glandular hairs. Calyx lobes
triangular 2,5-3 ,5 mm long, sharply acute, with
glandular hairs. Corolla with tube 18-23 mm long
with glandular hairs mainly on the ridges between
petals, yellowish-green; lobes 4-5 mm long, lanceo
late, obtuse, with glandular hairs on outside, pale
yellow. Stamens 23-24 mm long, with equal anthers
c. 1,8 mm long, with filaments somewhat recurved
below anthers and with fine spreading hairs towards
the base where fused to the petal tube. Squamae
almost square 1-1,2 x0 ,9-1 mm, scarcely constricted
towards the base, emarginate, slightly fleshy, white.
Carpels with slender ovaries gradually constricted
into erect styles with terminal stigmas; ovary with
18-24 ovules each about twice as long as broad,
slightly broadened towards the base and with vertical
ridges.
T. tuberosus Toelken, sp. nov a T. ventricoso
ramis 5-8 mm in diametro, foliis pilis glanduliferis
2-3 mm longis, petalis 14-17 mm longis; a T. striato
ramis flavis levibus usque ad 50 mm longes, foliis
dorsoventraliter compressis et pilis glanduliferis 2-3
mm longis differt.
Plantae perennes ramis multiramosis 5-8 mm in
diametro, usque ad 50 mm longis supra terram sed
tegetes multiramosae densae formantes infra, cortice
flava levi et phyllopodiis in ramiis aeriis. Folia
elliptica vel lanceolato-spathulata, (15-) 20-50(-70)
mm longa, (6-) 8-16 mm lata, acuta vel obtusa,
cuneata vel petiolata, dorsiventraliter compressa et
recurva, pilis glanduliferis 1,5-3 mm longis tecta,
griseo-viridia vel veneta. Inflorescentia thyrsus (1)
2-8 monochasiis quoque 1-5 floribus; pedunculi
(0,15-) 0 ,2-0,3 m longi rigidi, pilis glanduliferis;
pedicelli 6—10(—15) mm longi. Calycis lobi triangulares
3-4 mm longi, acuti, dense pilis glanduliferis tecti.
Corolla tubo 8-10 mm longo pilis glanduliferis extus
et pilis tenuibus paucis intra viridi suffuso fusco;
lobi 5-7 mm longi, lanceolati vel acuminati, pilis
glanduliferis extus, fusci. Stamina 9-13 mm longa,
antheris aequalibus c. 1 mm longis et oblongis etiam
post polline exutum est, filamentis flexis centrifuge
et pilis glanduliferis ubi corollae tubo connatis.
Squamae oblongae, 1-1 ,2 x 0 ,2-0,3 mm, non con
strictae ad bases, vix emarginatae, leviter succulentae,
pallide flavae. Carpella ovariis gracilibus gradatim
constrictis in stylos erectos stigmatibus terminalibus;
ovarium 36-40 ovulis quoque leviter latis factis ad
funiculum et cristis verticalibus.
Type: Cape, near Steinkopf, Marloth 13229
(PRE, holo!).
Perennials with much-branched stems 5-9 mm in
diameter up to 50 mm long above ground level but
forming a dense much-branched mat underground,
with smooth yellow bark and phyllopodia on aerial
branches. Leaves elliptic to lanceolate-spathulate,
(15-) 29-50 (-70) mm long, (6-) 8-16 mm broad,
acute to obtuse, cuneate to petiolate, dorsiventrally
compressed and recurved, covered with long glandular
hairs c. 1,5-3 mm long, grey-green to blue-green.
Inflorescence a thyrse with (1) 2-8 monochasia each
with 1-5 flowers; peduncles (0,15-) 0 ,2-0 ,3 m long,
stiff, covered with glandular hairs; pedicels 6-10
382 NEW TAXA A ND NEW COMBINATIONS IN COTYLEDON AN D ALLIED GENERA
(-15) mm long. Calyx lobes triangular 3-4 mm long,
acute, densely covered with glandular hairs. Corolla
with tube 8-10 mm long with glandular hairs outside,
with few fine hairs inside, green tinged brown; lobes
5-7 mm long, lanceolate, acute to acuminate, glandu
lar hairy outside, brown. Stamens 9-13 mm long,
with anthers equal, c. 1 mm long and oblong even
after the pollen is shed, with filaments bent outwards
below anthers and with fine spreading hairs where
fused to the petal tube. Squamae oblong, 1-1,2 x 0,2 -
0,3 mm, not constricted towards the base, scarcely
emarginate, slightly fleshy, pale yellow. Carpels with
slender ovaries gradually constricted into erect styles
with terminal stigmas; ovary with 36-40 ovules each
slightly broadened towards the funicle and with
vertical ridges.
T. ventricosus (Burm.f.) Toelken, comb. nov.
Cotyledon ventricosa Burm.f., Prodr. FI. Cap. 13 (1768);
DC. Prodr. 3: 397 (1828); Harv., FI. Cap. 2: 375 (1862); Schonl.
& Bak.f. in J. Bot., Lond. 40: 90 (1902); Schonl. in Rec. Albany
Mus. 3: 150(1915). Iconotype:J. Burm., Rar. Afr. PI. t.21, fig. 1
(1732). — var. alpina Harv., FI. Cap. 2: 376 (1862); Schonl. &
Bak.f. in J. Bot., Lond. 40: 90 (1902). Type: Cape, Elandsberg,
Wallich s.n. (K, lecto!).
T. ventricosus is an extremely variable species which
is distinguished from a number of very similar species
by its thick stems [13-20 (-30) mm in diameter] with
yellow bark and phyllopodia even when young and
long brown petals 20-28 mm long. Numerous local
forms occur throughout its distribution range often
in isolated mountains in and around the Great
Karoo. Whether some of these forms should be recog
nized as subspecies can not be finally decided at present
because too little is known about the range of variation
of the vegetative parts as these are poorly represented
in herbaria. The type specimen of the var. alpina
is a depauperate specimen of typical T. ventricosus.
T. viriditlorus (Toelken) Toelken, comb. nov.
Cotyledon viridiflora Toelken in Bothalia 12: 193 (1977).
Cape, near Modderfontein, Tolken 5327 (PRE, holo!).
T. wallichii (Harv.) Toelken, comb. nov.
Cotyledon wallichii Harv., FI. Cap. 2: 374 (1862); Schonl. in
Rec. Albany Mus. 3 :148 (1915). Type: Cape, Snowy Mountains,
Wallich s.n. (K, lecto!).
subsp. wallichii.
Cotyledon wallichii Harv., FI. Cap. 2: 374 (1862); Schonl. in
Rec. Albany Mus. 2: 152 (1904); 3: 148 (1915); Marl, in FI. S.
Afr. 2: 4 (1925); V. Poelln. in Reprium nov. Spec. Regni veg.
42: 25 (1937).
Peduncle and pedicels usually covered with glandu
lar hairs. Calyx usually densely covered with glandular
hairs. Petals densely covered with glandular hairs on
the lobes and along the ridges on the petals but also
a few smaller ones between ridges. Corolla tube
(8-) 10-12 mm long.
Occuring on lower slopes or on gravelly or sandy
soils in depressions from south-west and north-east
of Kamieskroon mainly along the eastern parts of
south-western Cape mountains to near Oudtshoorn.
subsp. ecklonianus (Harv.) Toelken, stat. nov.
Cotyledon eckloniana Harv., FI. Cap. 2: 374 (1862); Schonl.
in Rec. Albany Mus. 3: 148 (1915); V. Poelln. in Reprium nov.
Spec. Regni veg. 42: 20 (1937). Type: Cape, Kamiesberg,
Ecklon & Zeyher 1967 (S. holo!). C. dinteri Bak.f. in Bull. Herb.
Boissier ser. 2, 3: 818 (1903); V. Poelln. in Reprium nov. Spec.
Regni veg. 42: 19 (1937); Jacobsen, Handb. Succ. PI. 1: 281,
pi. 276 (1960); Sukk. Lex. 133, t. 38.4 (1970); Friedr. in Prodr.
FI. S. W. Afr. 52: 8 (1968). Type: So ith West Africa, Great
Namaland, Dinter 931 (Bf). C. cacalioides sensu Eckl. & Zeyh.,
Enum. 307 (1837), non Thunb.
Peduncles and pedicels usually glabrous. Calyx
glabrous or with few glandular hairs when in bud.
Petals glabrous except for a few glandular hairs on
the margins of the lobes and on the area where the
petals are fused to one another; corolla tube 5-8 mm
long.
Occurring on gravelly slopes from north of Kamies
kroon to Aus in south-western South West Africa.
3. ADROMISCHUS
A revived interest in the genus Adromischus between
1952-1960 led to the description of 15 new species,
but no monograph of the genus has yet been published.
A revision of the genus was attempted by C. A. Smith
(1939) and von Poellnitz (1940), but due to a lack of
authentic specimens and especially of adequate ranges
of material it was impossible for these authors to
evaluate the amplitude of the variation adequately.
Therefore, a more detailed account of the genus is
presented here.
The history of the nomenclature of the sections is
varied, but typically shows the neglect and superficial
treatment given to this genus. In the publication of
the original description of the genus, Lemaire (1852)
distinguished two groups of species and indicated
in the text that he had envisaged sectional level
for them. Since Jacobsen (1966) chose A. hemisphaeri-
cus as the type species of the genus, sect. Suffruticuli
Lem. must be replaced by sect. Adromischus. The
name of Lemaire’s second section cannot be accepted,
because it is a polynomial. Schonland (1915) recog
nized two groups in that part of Cotyledon which
agrees with the present Adromischus, but the names
were never taken up at sectional level. C. A. Smith
(1939) proposed to divide the genus into a section
with opposite leaves and one with spirally arranged
leaves. These characters, however, are very difficult
to recognize and von Poellnitz (1940a) already pointed
out that he could not place many of his plants clearly
into either of the categories. The names proposed by
Smith were, however, not validly published, because
he did not add Latin diagnoses of these sections.
The sections Brevipedunculati V. Poelln. and Longipe-
dunculati V. Poelln. are also not based on floral
morphology. Uitewaal (1952, 1963) again ignored
all previous proposals when he divided the genus
into sections Inscisilobati and Connatilobati, but
these sections were for the first time based on floral
morphology. Finally, Jacobsen (1966, 1970) in his
review of the genus did not validly publish his
subsect. Pendenti, because he omitted to add a Latin
diagnosis.
The importance of floral characters in the classifica
tion of this genus was only recently recognized so
that many interpretations in older literature cannot
be evaluated. The absence of voucher specimens for
the species of Haworth and de Candolle makes it
impossible to check their identifications. For instance,
the name A. mammillaris L.f. had been accepted for
a long time for plants with decumbent stems and
terete leaves. It was only now found that the type of
this species represents an extremely rare species from
the vicinity of Calitzdorp, while Thunberg apparently
never collected the very common species which was
known by this name for about two hundred years.
The criticism has been raised that types of flowers
considered to be typical of a certain section are
merely flower types with a specific pollination syn
drome. This can easily be refuted by pointing out that
although species with the same flower type may occur
in different sections of Adromischus, they are in other
respects typical of their section. The salver-shaped
flowers of A. leucophyllus, for instance, superficially
resemble those found in sect. Brevipedunculati, but the
whole plant is densely covered with a bloom, club-
shaped trichomes are restricted to the throat and the
H. R. TÓLKEN 383
squamae are not oblong. All these characters place A.
leucophyllusrather into the sect. Longipedunculati. Even
more obvious are the red pendulous flowers of
A. phillipsiae, which seem to simulate those of Cotyle
don orbiculata.
Most flowers in Adromisehus are, however, super
ficially similar and are pollinated according to the
Thunbergia-grandiflora-syndrome (Faegri & van der
Pijl, 1971). Although the flowers of Adromisehus
are morphologically different and much less showy
than T. grandiflora, they are visited, and most probably
pollinated by bees and wasps. The extra-floral nectar
attracts ants and their presence prevents the pollina
tors from piercing the corolla at the base in order to
get more readily at the nectar inside. This was often
observed in the field and there seems to be no specifi
city in the type of bee or wasp that is attracted to
the flowers. The ants move continuously from one
extra-floral nectary to another, but they have never
been observed to enter the flowers or even to be
attracted to the nectar inside them, as was often seen
in species of Cotyledon.
Adromisehus Lem., Jard. Fleur. 2, Misc. 60 (1852);
Berger in Pflanzenfam. ed. 2, 18a: 415 (1930); C. A.
Smith in Bothalia 3: 613 (1939); V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 89 (1940); Jacobsen in
Kakteen, Berl. 17: 189 (1966); Friedr. in Prodr. FI.
S. W. Afr. 52: 1 (1968); Dyer, Gen. 1: 198 (1975).
Type species: A. hemisphaericus (L.) Lem.
Shrublets rarely up to 0,25 m high without inflores
cence, with cartilaginous branches usually with
peeling bark on very old stems, often with much
branched tuberous base. Leaves spirally arranged,
persistent, stiff or rarely softly herbaceous and with
smooth waxy surface. Inflorescence usually a spike
like thyrse with 1 — numerous monochasia each
with 1 (-5) erect or spreading flowers; bracts on
peduncle much shorter than and not grading gra
dually into the leaves below. Calyx glabrous on
abaxial surface (often hairy in A. cristatus) and usually
with club-shaped trichomes on adaxial surface. Corolla
fused beyond the tube, glabrous (rarely hairy in
A. cristatus), with green to white lobes often tinged
pink to deep mauve (orange in A. phillipsiae). Filaments
glabrous or papillose along the whole length. Squamae
a free disc, white or pale yellow. Ovary with 20-30
(-50) ovules in one row along each placenta.
Fig. 1.— Diagrams of floral characteristics of the sections o f Adromisehus: 1, A.filicaulis subsp. marlothii, sect.
Adromisehus (Tolken 55 04); 2, A. trigynus, sect. Boreali (Tolken 54 16); 3, A. humilis, sect. Brevipeduneulati
{Marloth 46 89); 4, A. maculatus, sect. Inseisilobati (Tolken 5493); 5, A. subviridis, sect. Longipedunculati
(Tolken 5349). l- 5a , openin g flower fro m a bove ( x 2); 1-5 b, mature flower in side view ( x 4); 1—5 c, m ature
bud (x 1); 1- 5 d, nectary scale ( x 1).
Key to Sections (see also Fig. 1)
Anthers protruding above the corolla tube; corolla lobes at least as broad as long:
Corolla lobes abruptly constricted into apical point, undulate and frilled
.....................
1. sect. Adromisehus
Corolla lobes gradually constricted into acute apex, undulate............................................2. sect. Boreali
Anthers included in corolla tube; corolla lobes up to 3 times longer than broad:
Corolla tube obconical in flowers; squamae about twice as long as broad
.................
3. sect. Brevipeduneulati
Corolla tube cylindrical in flowers; squamae usually broader than long, up to as long as broad, rarely
slightly longer:
Buds distinctly grooved between petals and until flowering adpressed to central axis; club-shaped
trichomes only in throat of corolla tube
............................................................ 5. sect. Longipedunculati
Buds cylindrical or slightly angular and spreading; club-shaped trichomes on lower parts of lobes and
in throat of corolla tube........................................................................................4. sect. Inseisilobati
384 NEW TAXA AND NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
1. Sect. Adromischus. Jacobsen in Kakteen, Berl.
17: 189 (1966); Sukk. Lex. 27 (1970).
Adromischus Lem., Jard. Fleur. 2, Misc. 60 (1852):
Berger in Pflanzenfam. ed. 2, 18a: 415 (1930); Friedr.
in Prodr. FI. S.W. Afr. 52: 1 (1968). A. sect.
Suffruticuli Lem., Jard. Fleur. 2, Misc. 60 (1852).
Type species: A. robustus Lem. A. sect. Connatilobati
Uitew. in Natn. Cact. Succ. J. 7: 70 (1952). Type
species; A. filicaulis (Eckl. & Zeyh.) C. A. Smith
[=A. mammillaris sensu Uitew., non (L.f.) Lem.].
A. [sect. Longipedunculati V. Poelln. in Reprium nov.
Spec. Regni veg. 48: 89 (1940)] subsect. Hemisphaeri-
cus (Schonl.) V. Poelln. in Reprium nov. Spec. Regni
veg. 48: 89 (1940); Jacobsen in Kakteen, Berl. 17:
190 (1966); in Sukk. Lex. 27 (1970). Type species:
A. hemisphaericus (L.) Lem.
Cotyledon [sect. Spicatae Harv., FI. Cap. 2: 370
(1862)] group Hemisphaerica Schonl. in Rec. Albany
Mus. 3: 151 (1915).
Inflorescence straight, with mature buds and flowers
usually spreading at about right angles and with
extra-floral nectaries present on pedicels. Buds with
almost cylindrical tube and abruptly constricted into
a short but acute apex. Corolla with tube green often
tinged red, glabrous; lobes with very broad base
abruptly constricted into often short apical points
and usually reflexed against the tube, with undulate
and frilled margins, rough but usually without
trichomes, white, pale pink rarely red except for the
midrib and continued into the acumen. Stamens
unequally long, with anthers of outer whorl slightly
longer and just protruding above the throat or rarely
enclosed, with filaments usually papillose. Squamae
about square, broadest at about the middle.
Represented by a number of species in the area
between south-western South West Africa and the
South-Western Cape, but A. bicolor has been recorded
a few times east of Steytlerville.
Key to Species and Subspecies
Leaves linear-elliptic rarely linear-lanceolate, terete or almost so:
Stems erect or decumbent and rarely with fibrous adventitious roots.........................A. filicaulis subsp. filicaulis
Stems prostrate or decumbent and with stilt roots unbranched for about 10-20 mm A. filicaulis subsp. marlothii
Leaves oblanceolate, obovate to orbicular and more or less dorsiventrally compressed:
Leaves petiolate:
Leaf blade deltoid to obtriangular............................................................................................. A. liebenbergii
Leaf blade oblanceolate to obovate:
Stems with leaves (3-) 4-6 mm in diameter and usually somewhat zig-zagging; leaf blade oblanceolate
to obovate............................................................................................................................A. roaneanus
Stems with leaves (6-) 7 -10 mm in diameter and straight; leaf blade oblong-oblanceolate
.................
A. filicaulis subsp. filicaulis
T pavps
Leaves (30-) 40-80 (-110) mm long:
Stems with leaves (3-) 4-6 mm in diameter and usually somewhat zig-zagging.........................A. roaneanus
Stems with leaves (6-) 8-12 mm in diameter and more or less straight but sometimes very short and
much branched:
Namaqualand to SW South West Africa; stems becoming woody and little branched..........A. alstonii
Eastern Cape, east of Steytlerville; stems much branched and tough cartilaginous
.............
A. bicolor
Leaves (8-) 20-30 (-35) mm long:
Inflorescence (40-) 80-100 (-150) mm long; northern Richtersveld and SW South West Africa
..........
A. montium-klinghardtii
Inflorescence (150—) 200-400 (-450) mm long; South-Western Cape mountains up to Nieuwoudtville
and Eastern Cape:
Leaves densely covered with red spots and wax which on old leaves become powdery; eastern
Cape, east of Steytlerville...................................................................................................A. bicolor
Leaves rarely with purple spots and wax which on old leaves breaks into angular pieces larger than
2 mm square; South-western Cape:
Leaves acute and with a sharply-edged margin; mountains between Clanwilliam and Nieuwoudt
ville ................................................................................................................................A. roaneanus
Leaves usually rounded, rarely bluntly obtuse or emarginate; mountains between Worcester and
Cape Town.........................................................................................................A. hemisphaericus
A. alstonii (Schonl. & Bak.f.) C. A. Smith in
Bothalia 3: 638 (1939). Type: Cape, Namaqualand
Alston s.n. (GRA, ho lo!; SAM !).
Cotyledon alstonii Schonl. & Bak.f. in J. Bot., Lond. 40: 93
(1902).
Adromischus triebneri V. Poelln. in Beitr. Sukkulentenk. (1939)
18; Friedr. in Prodr. FI. S. W. Afr. 52: 5 (1968.) Type: Cape,
Springbok, Triebner 1331 (Bf). A. subrubellus V. Poelln. in
Reprium nov. Spec. Regni veg. 50: 319 (1941). Type: Cape, 30
km NW Vanrhynsdorp, Triebner 1351 (Bf). A. pulchellus P. C.
Hutch, in Cact. Succ. J., Los Ang. 31: 118, figs 52-4 (1959).
Type: Cape, Bowesdorp, Hall in NBG 761/53 (Univ. Calif. Bot.
Gard. 54.110-1: BOL, holo!; PRE!).
The illustration of the flower of A. robustus Lem.
with its original description indicates that this species
belongs into this section, but neither is the description
specific enough nor has a type specimen been pre
served, so that it is impossible to identify the species.
The name infers a robust habit and, as A. alstonii
is the most common species north of Garies, it is
likely that A. robustus, being an older name, may
have to replace the present name if more evidence can
be found.
Similarly, no isotypes of A. triebneri and A. subru
bellus could be traced, so that these species are included
here on the strength of their fairly large leaves and on
the evidence of specimens found at the type localities.
A. bicolor P. C. Hutch, in Cact. Succ. J., Los
Ang. 29: 15 (1957). Type: Cape, near Steytlerville,
Hall in NBG 914/47 (Univ. Calif. Bot. Gard. 53.1109-
1: BOL,holo!; PRE!; UC!).
This species is only known from a few localities
east of Steytlerville. The specimen Flanagan 1113
from near Komga has unusually long cuneate leaf
bases but no plants of that taxon have been found in
recent years in that area.
A. filicaulis (Eckl. & Zeyh.) C. A. Smith in Botha
lia 3: 630 (1939). Type: Cape, Kamiesberg, Ecklon &
Zeyher 1975 (S!).
Cotyledon filicaulis Eckl. & Zeyh., Enum. 307 (1837).
The range of variation in size, shape and colour
of the leaves, in the habit and in the colour of the
petals observed in specimens from the area between
Springbok, Kamieskroon and Komaggas is so wide
that it is impossible to distinguish the species described
within the present circumscription of subsp. filicaulis.
Yet at no single locality within this area is an
extensive range found which would suggest a hybrid
swarm.
H. R. TÓLKEN 385
subsp. filicaulis.
Cotyledon filicaulis Eckl. & Zeyh., Enum. 307 (1837). C-
fusiformis Rolfe in Kew Bull. (1916) 229. Type: Cape, sine loc.»
Pearson 5585 (K, holo!).
Adromisehus filicaulis (Eckl. & Zeyh.) C. A. Smith in Bothalia
3: 630 (1939). A. fusiformis (Rolfe) Berger in Pflanzenfam. ed.
2 18a: 416 (1930). A. mammillaris var. rubra V. Poelln. in
Desert PI. Life 10: 112, fig. (1938); in Reprium nov. Spec.
Regni veg. 40: 109 (1940). Iconotype: Bot. Mag. 99, t.6020
(1873).—var. filicaulis (Eckl. & Zeyh.) Jacobsen, Sukk. Lex.
29 (1970), nomen non rite publicatum quoad sine typo. —var.
fusiformis (Rolfe) Jacobsen, Sukk. Lex. 29 (1970), nomen non
rite publicatum quoad sine type.—var. marlothii (Schonl.)
Jacobsen, Sukk. Lex. 29 (1970), non rite publicatum quoad sine
typo. A. kleinioides C. A. Smith in Bothalia 4: 631 (1939).
Iconotype: Bot. Mag. 99, t.6020 (1873). A. fragilis P. C. Hutch.
inCact. Succ. J., Los Ang. 31: 167 (1959). Type: Cape, Richters-
veld, Hellsberg, Rodin 1620 (Univ. Calif. Bot. Gard. 50.1180:
BOL, holo!).—var. numeesensis P. C. Hutch, in Cact. Succ. J.,
Los Ang. 31: 169 (1959). Type: Cape, Richtersveld, Numees,
Hall in NBG 692/53 (Univ. Calif. Bot. Gard. 54.113-1: BOL,
holo!; PRE!).
subsp. marlothii (Schonl.) Toelken, stat. nov.
Cotyledon marlothii Schonl. in Rec. Albany Mus. 1: 59(1903);
3: 153 (1915). Type: Cape, near Laingsburg, Marloth 2520
(GRA, holo!). C mammillaris sensu Harv., FI. Cap. 2: 377
(1862), pro parte, excl. C. filicaulis; sensu Schonl. in Rec.
Albany Mus. 3: 153 (1915), pro parte, non L.f.
Adromisehus marlothii (Schonl.) Berger in Pflanzenfam. ed. 2,
18a: 416(1930). A. tricolorC. A. Smith in Bothalia 4: 632(1939).
Type: Cape, Brandvlei, Schlechter 9933 (PRE, holo!; K!).
A. mammillaris sensu Berger in Pflanzenfam. ed. 2, 18a: 416
(1930); sensu V. Poelln. in Desert PI. Life 10: 112 (1938); in
Reprium nov. Spec. Regni veg. 48: 109 (1940); sensu C. A.
Smith in Bothalia 3: 631 (1939), non (L.f.) Lem.
The type specimen of Adromisehus mammillaris
L.f. clearly shows that this species must be placed
into the sect. Inseisilobati.
A. hemisphaericus (L.) Lem., Jard. Fleur. 2,
Misc. 60 (1852); C. A. Smith in Bothalia 3: 625
(1939); V. Poelln. in Reprium nov. Spec. Regni veg.
48: 105 (1940). Iconotype: Dill., Hort. Eltham. t.95.
fig. Ill (1732).
Cotyledon hemisphaerica L., Sp. PI. ed. 1, 429 (1753); DC.*
PI. Hist. Succ. t.87 (1802); Prodr. 3: 398 (1828); Eckl. & Zeyh.»
Enum 307 (1837), pro parte; Harv., FI. Cap. 2: 376 (1862)»
pro parte; Schonl. in Rec. Albany Mus. 3: 152(1915), pro parte»
excl. C. triflora. C. crassifolia Salisb., Prodr. 307 (1796). Type:
unknown. C. rotundifolia Haw. in Phil. Mag. (1827) 273;
Schonl. & Bak.f. in J. Bot., Lond. 40: 91, t.435 (1902); Dyer in
Bot. Mag. 157, t.9368 (1934). Type: sine loc., Haworth s.n.
(OXF, holo!).
Adromisehus rotundifolius (Haw.) C. A. Smith in Bothalia
3: 627 (1939); V. Poelln. in Kakteenkunde (1940) 17, fig.
A characteristic of this species is that the wax
layer covering the epidermis of the leaves breaks
into large isodiametric pieces. Similar leaves may also
be found in A. roaneanus and a few other species of
this section when grown under adverse conditions,
but the phenomenon is then usually restricted to a
few leaves.
Plants from the Cape Peninsula tend to have smaller
leaves often with an obtuse apex, and club-shaped
trichomes are absent on the corolla lobes. Plants
from the mountains to the east of the Cape Peninsula,
and especially in the vicinity of Worcester, however,
tend to have larger leaves with rounded or emarginate
apices and, in contrast to all other species in the
section, a few club-shaped trichomes are present on
the throat of the corolla. Formal rank was not given
to the latter form because of a range of intermediates
in collections such as Esterhuysen 23 898 and 27 599.
A. liebenbergii P. C. Hutch, in Cact. Succ. J.,
Los Ang. 31: 81, fig. 40 (1959). Type: Cape, Vars-
bokkraal near Laingsburg, Liebenberg 6186 (BOL,
holo!).
Except for the very markedly convex base of the
expanded leaf blade, vegetative material of A. lieben
bergii cannot be distinguished from the local form of
A. triflorus (sect. Inseisilobati), which tends to have
petiolate leaves in the eastern parts of its distribution
range.
A. montium-klinghardtii (Dinter) Berger in Pflan
zenfam. ed. 2, 18a: 416 (1930); Friedr. in Prodr.
FI. S.W. Afr. 52: 4 (1968). Type: South West
Africa, Klinghardt mountains, Dinter 4265 (Bf).
Cotyledon montium-klinghardtii Dinter in Reprium nov. Spec.
Regni veg. 19: 147 (1923).
This species has often been linked with A. hemis
phaericus on account of its similar small leaves, but
apart from the different habitat and distribution, it
is distinguished by the absence of the cracked wax
layer on the leaves and its very short inflorescence.
The latter character may also be induced by adverse
conditions in plants of A. hemisphaericus, but the
inflorescence of A. montium-klinghardtii is always
markedly shorter.
A. roaneanus Uitew. in Natn. Cact. Succ. J. 7:
69, figs (1952). Type: Cape, Vanrhyn’s Pass, Herre
in SUG 6058 (AVU 10011, holo).
P. C. Hutchinson has annotated herbarium speci
mens of this species as A. grasbergensis, or, in the
case of more vigorous plants as A. violaceus. Neither
of these names has, however, been validly published.
2. Sect. Boreali Toelken, sect. nov. a sect. Adromischo
corollae lobis late triangularibus et trichomatibus
claviformibus differt.
Inflorescentia recta alabastris semper plus minusve
adpressis rami dum ante florentibus, plerumque sine
nectariis extrafloralibus in pedicellis. Alabastra tubis
plus minusve cylindricis sed aliquantum sulcatis inter
petala et gradatim constricta ad apices acutos. Corolla
tubo glauco vel rubro; lobi ovato-triangulares et
gradatim constricti ad apices acutos et plerumque
reflexi et plus minusve adpressi tubem, asperi et
trichomatibus claviformibus praecipue in fauce, albi
vel rosei et atro-rosei in parte aperta exterior. Stamina
paene simile longa, antheris papillosis et circiter
simile longis et plerumque leviter protrusis super
tubem corollae, filamentis glabris. Squamae plerumque
quadratae vel oblongae et latissimae ad bases.
Type species: A. umbraticola C. A. Smith
Inflorescence straight with buds always more or
less adpressed to the stem until just before flowering,
usually without extra-floral nectaries on pedicels.
Buds with more or less cylindrical tube but somewhat
grooved between petals and gradually constricted into
acute apices. Corolla with tube glaucous-pink or red;
lobes ovate-triangular and gradually tapering into
an acute apex, usually reflexed against the tube which
thus appears to have an undulate fringe without
lobes, rough and usually with some club-shaped
trichomes mainly in the throat, white or pink and dark
pink on the exposed part of the outside. Stamens
almost equally long, with anthers papillose and about
equally long, usually slightly protruding above the
reflexed petals, with filaments glabrous. Squamae
square to oblong and broadest towards the base.
Occurring in dryer mountainous areas to the east,
south and west of the Kalahari basin. All species
are found in the summer rainfall area to the north
of all the other species of Adromisehus.
386 NEW TA X A AND NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
Key to Species and Subspecies
Leaves elliptic to orbicular with a marked marginal ridge extending to its base................................................A. trigynus
Leaves oblanceolate to obovate, with marginal ridge rarely extending beyond the middle, or if so then narrow and
not horny:
Corolla with club-shaped trichomes in throat and on lower parts of lobes
.............
A. nmbraticola subsp. umbraticola
Corolla without club-shaped trichomes in throat or on lobes:
Stems up to 30 mm long, much branched...........................................................A. schuldtianus subsp. schuldtianus
Stems 40-120 mm long, little branched:
Branches sinuous; northern Transvaal...............................................................A. umbraticola subsp. ramosus
Branches spreading; Karas Mountains in South West Africa
................................
A. schuldtianus subsp. juttae
A. schuldtianus (V. Poelln.) V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 110(1940); Friedr. in Prodr.
FI. S.W. Afr. 52: 4 (1968). Type: sine loc. et leg.
(Bf).
A. schuldtianus is found widespread in the moun
tains of central South West Africa. In cultivation the
longer branches of the subsp. juttae remain distinctive,
similar to those in A. umbraticola subsp. ramosus.
A particularly narrow-leafed form of the subsp.
schuldtianus was described by Nordenstam (1974)
from the upper Brandberg, but even the few records
of the species known at present show considerable
range of variation in the shape and size of the leaves.
subsp. schuldtianus.
Cotyledon schuldtiana V. Poelln. in Jb. dt. KaktGes. 1: 95
(1936). C. trigynus sensu Schonl. in Ann. Bolus Herb. 1: 15
(1914).
Adromischus schuldtianus (V. Poelln.) V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 110 (1940); Friedr. in Prodr. FI. S. W.
Afr. 52:4(1968); Nordenstam in Dinteria 11: 15, fig. 1 (1974).
subsp. juttae (V. Poelln.) Toelken, stat. nov.
Adromischus juttae V. Poelln. in Kakteenkunde (1939) 52, fig.
Syntypes: South West Africa, Garub, Triebner 1305 (Bf);
Norachus, Triebner 1315 (Bf).
A. trigynus (Burch.) V. Poelln. in Reprium nov.
Spec. Regni veg. 44: 61 (1938); 48: 111 (1940), pro
parte; C. A. Smith in Bothalia 3: 642 (1939). Type:
Cape, near Griqua Town, Burchell 1898 (K, holo!).
Cotyledon trigynus Burch., Trav. 2: 226 (1824); DC., Prodr. 3:
398 (1828); Schonl. & Bak.f. in J. Bot., Lond. 40: 91 (1902);
Schonl. in Rec. Albany Mus. 3: 153 (1915). C. rhombifolia var.
spathulata N.E. Br. in Marloth, FI. S. Afr. 2,1: 15, t. 2D (1925),
nom nud.
Adromischus subcompressus V. Poelln. in Reprium nov.
Spec. Regni veg. 44: 62 (1938), pro parte quoad Triebner 1330.
A. rupicola C. A. Smith in Bothalia 3: 642 (1939). Type: Cape,
Fauresmith, Smith 5603 (PRE, holo!). A. nanus sensu C. A.
Smith in Bothalia 3: 640(1939), non (N.E. Br.) V. Poelln.
Widespread, but never common in rock outcrops
in the northern Karoo and in Bushmanland. The
leaves are tough and remarkably uniform in shape and
size on the same plant in contrast to A. umbraticola
and A. schuldtianus.
C. A. Smith described A. rupicola, because he could
not clearly interpret Burchell’s A. trigynus. However,
the fragmentary type specimen together with a collec
tion from the type locality (Liebenberg 5955) leave no
doubt about the identity of the species.
A. umbraticola C. A. Smith in Onderstepoort J.
vet. Sci. Anim. Ind. 1: 174 (1933); in Bothalia 3:
643 (1939). Type: Transvaal, Silkaatsnek, Smith 3432
(PRE, holo!).
The species usually grows in somewhat shaded
localities but it may also be found on exposed rock
faces with a south-eastern aspect. A. saxicola grades
into A. umbraticola as can be observed when studying
the wide range of variation in the size and shape of
leaves of Tolken 5435 from the Magaliesberg. A.
saxicola is thus placed into the synonomy of A.
umbraticola.
It stands to reason that plants in a shaded position
will produce longer stems but the absence of club
shaped trichomes on the throat of the corolla tube
will always identify plants of the subsp. ramosa.
subsp. umbraticola.
Adromischus umbraticola C. A. Smith in Onderstepoort J. vet.
Sci. Anim. Ind. 1: 174 (1933); in Bothalia 3: 643 (1939); Letty,
Wild Flow. Transv. fig. 75,4 (1962). A. saxicola C. A. Smith in
Bothalia 3: 647 (1939). Type: Transvaal, Baviaanspoort, Smith
3424 (PRE, lecto!).
Cotyledon trigyna sensu Burtt Davy, FI. Transv. 143 (1926),
non Burch.
Branches 20-40 (-60) mm long, much branched.
Leaves grey-green or glaucous often tinged brown and
sometimes faint purplish spots. Corolla with many
club-shaped trichomes.
subsp. ramosa Toelken, subsp. nov.
Rami 60-120 mm long, pauciramosi. Folia viridia
saepe suffusa fusca, rare viridi-grisea, immaculata.
Corolla sine trichomatibus clavatis.
Type: Transvaal, Chunies Poort, Tolken 1215 (PRE,
holo!).
Branches 60-120 mm long, little branched. Leaves
green often tinged brown, rarely grey-green and without
spots. Corolla without club-shaped trichomes.
Occurring in the mountains from Middelburg to the
Zoutpansberg.
3. Sect. Brevipedunculati V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 89 (1940), pro parte;
Jacobsen in Kakteen, Berl. 17: 189 (1966), pro parte;
in Sukk. Lex. 27 (1970), pro parte. Type species:
A. humilis (M arl.) V. Poelln.
Cotyledon [sect. Paniculatae Harv., FI. Cap. 2:
370 (1862)] group Caryophyllacea Schonl. in Rec.
Albany Mus. 3: 150(1915). Type species: C. caryophyl
lacea Burm. f.
Adromischus [sect. Adromischus. Jacobsen in Kak
teen, Berl. 17: 190 (1966)] subsect. Pendenti Jacobsen
in Kakteen, Berl. 17: 191 (1966); in Sukk. Lex. 27
(1970), nomen non rite publicatum. Type species:
A. phillipsiae (M arl.) V. Poelln.
Inflorescence more or less straight and often
branched, with spreading, rarely pendulous flowers,
usually without extra-floral nectaries on the pedicels.
Buds cylindrical, grooved between the upper parts
of the petals and gradually constricted into blunt
apices. Corolla with tube glaucous-green rarely tinged
pink, glabrous; lobes ovate to lanceolate but usually
with a broad base, spreading to recurved, rough and
usually with club-shaped trichomes on the lobes and
in the throat, white to deep mauve or purple (rarely
orange), or mauve along the middle of the petals.
Stamens unequal, longer ones with anthers up to
twice as long as shorter ones, enclosed in corolla
tube, with filaments usually glabrous. Squamae oblong,
at least twice as long as broad, not bulging.
Recorded mainly from areas just south of the Great
Karoo except for A. humilis, which occurs in the
Nieuweveld Mountains, and A. nanus which is found
near Steinkopf. Most species have a very restricted
distribution.
H. R. TÓLKEN 387
Species in this section often have soft herbaceous
leaves without a horny marginal ridge, and are thus
reminiscent of the leaves of species in Tylecodon,
but they are not seasonal although their number is
usually drastically reduced during the dry period.
Most of the species have tuberous roots similar to
A. marianae (sect. Longipedunculati).
The subsect. Pendenti was not validly described
with a Latin diagnosis in either of the publications
in which Jacobsen expounded his classification of
the genus.
Key to Species
Leaves tough, dorsiventrally compressed.................................................................................A. caryophyllaceus
Leaves soft herbaceous, grooved, concave to cymbiform above:
Flowers pendulous, orange-red......................................................................................................A. phillipsiae
Flowers spreading, white tinged pink to purple:
Inflorescence with 1 (-3) flowers..........................................................................................................A. nanus
Inflorescence with 5—12 flowers:
Stems short (up to 40 mm long), much branched; corolla with few or no club-shaped
trichomes............................................................................................................................A. humilis
Stems long (up to 150 mm), little branched; corolla with many club-shaped trichomes
.........
A. fallax
A. caryophyllaceus (Burm.f.) Lem., Jard. Fleur. Suffrutex with decumbent branches 5-15 mm Oi
2, Misc. 60 (1852); C. A. Smith in Bothalia 3: 629 diameter. Leaves oblanceolate to elliptic, 20-40 (-5)n
(1939), pro parte excl. Bolus 758; V. Poelln. in mm long, 8-15 (-20) mm broad, acute to obtuse,
Reprium nov. Spec. Regni veg. 48: 107 (1940). cuneate, each without horny marginal ridge. lnflores-
Iconotype: J. Burm., Rar. Afr. PI. 1.17 (1738). cence a spike-like thyrse with 1-3 (-5) flowers, with
Cotyledon caryophyllacea Burm. f., Prodr. FI. Cap. 13 (1768); pedicels 10—15 mm long with fleshy bracts widely
DC., Prodr. 3: 398 (1828); Schonl. & Bak.f. in J. Bot., Lond. spaced. Sepals triangular, 4-5 mm long, acute, green.
40: 93 (1902), pro parte; in Rec. Albany Mus. 3: 151 (1915), Petals 14-17 mm long, fused into a tube 10-13 mm
DC™'Prodr 3*398 lo ng; lobes o vate-triangular, 3 -4 ,5 mm long, acute,
(1828). Type: sine loc., Salm-Dyck sub Haworth (OXF, lecto!). with club-shaped trichomes around the throat. Sta-
C. bolusii Schonl. in Rec. Albany Mus. 1: 59 (1903); 3: 154 mens 12-14 mm long, with anthers of longer outer
(1915). Type: Cape, Mosselbaai, Bolus 8648 (BOL, holo!). whorl 1,8-2 mm long while those of inner whorl are
1-1.' 1°"S> all included. Squamae oblong,
18a: 416 (1930). A. grandiflorus Uitew. in Succulenta (1953) 8. 1 >° 2 x0 ,6-0 ,8 mm, usually deeply emarginate. Car-
Type: Cape, Bonnievale, sine leg. in SUG 6879 (AVU 10013, pels with slender ovaries gradually constricted into
holo). erect styles with terminal stigmas; ovary with 35
The leaves of this species are usually spathulate- ovules at least three times longer than broad and
oblanceolate but under adverse conditions their bases covered with vertical ridges.
become abruptly cuneate. A. grandiflorus is an extreme Bolus mentioned that this plant was very rare and
form with all its leaves abruptly cuneate and the restricted to the one locality, all attempts to redis
plays have a more erect habit. Intermediates between cover the species have failed,
this form and typical A. caryophyllaceus have been .
recorded in collections from Touwsberg, Esterhuysen A. humilis (Marl.) V. Poelln. in Reprium nov.
25 879, Bonnievale, Van der Merwe 97; Hermanus, Spec. Regni veg. 48: 91 (J940)._Type: Cape, near
Kriege in SUG 5360 and others. Judging by specimens - Beaufort West, Marloth 4689 (PRE, holo.).
from these localities it seems that plants from the Cotyledon humilis Marl, in Schonl. in Rec. Albany Mus. 3:
western parts of the distribution range of A. caryophyl- í 5^ 1915^ FIS- Aír AV w sensu Marl‘
, . j . , i_ * in Trans. Roy. Soc. S. Afr. 2 :33 (1910), non N.E. Br.
laceus show a tendency to have more abruptly cuneate „ . , , , „ , ,
leaf bases ^his plant has been collected only tw ice by M arloth
in the Nieuweveld Mountains and it could not be
A. fallax Toelken, sp. nov. ab A. humili plantis relocated,
grandioribus et trichomatibus clavatis multis in fauce A nanus £ Br ) y p m ; Desert p]
torolUe; ah A caryophyUaceo stammibus longioribus u f 10: 222 (1938). Type: Cape, sine loc., MacOwan
antheris duplo longionb us stammibus brevibus differt. s n (K h olo!)
i C° !y ie^ c°ry?Phyllacea sensu Schonl & Bak f. in J Bot Cotyledon nana N.E. Br. in Gdnr's Chron. 30: 270 (1901):
^ • 4 0 : 93 (1902), pro parte; Schonl. in Rec. Albany Mus. 3: Schonl & Bak f in j Bot Lond 40; 93 (1902).
151 (1915), pro parte quoad Bolus 758. Adromisehus pauciflorus P. C. Hutch, in Cact. Succ. J., Los
Adromisehus caryophyllaceus sensu C. A. Smith in Bothalia 3: Ang. 32: 63, fig. 32,33 (1960). Type: Cape, near Steinkopf, Hall
629(1939), pro parte quoad Bolus 758. in NBG 205/56 (Univ. Calif. Bot. Gard. 56.711-2: BOL, holo!;
UC!).
Suffrutex ramis decumbentibus 5-15 mm in dia- , . . . , ~
metro. Folia oblanceolata vel elliptica, 2(M 0 (-50) mm . Thls miniature species is known only from the
longa, 8-15 (-20) mm lata, acuta vel obtusa, cuneata, viclnl'>' °f S<el"k° pf, but th!i restricted distribution
quoque sine crista marginali. Inflorescentia thrysus "«V ube attributable to insufficient collecting in the
spiciformis 1-3 (-5) floribus, pedicellis 10-15 mm northern Cape.
longis. Sepala triangularia, 4-5 mm longa, acuta, A. phillipsiae (Marl.) V. Poelln. in Reprium nov.
vindia. Petala 14-17 mm longa, connata et formantia Spec. Regni veg. 48: 88 (1940). Type: Cape, southern
tubum 10-13 mm longum; lobi ovato-triangulares, Roggeveld, Marloth 3912 (PRE, lecto!; BOL!; G RA!;
3-4,5 mm longi, acuti, asperi cum trichomatibus SAM!).
clavatis in fauce. Stamina 12-14 mm longa, antheris _ . , ..... . T c Af D. .. ~ 1C
1 -U • 1 • C Cotyledon phillipsiae Marl, in Trans. S. Afr. Phil. Soc. 18.
lnaeque longis, filamentis laevibus inclusis. Squamae 46 (19J 7); F, s f a 2,\: 17, t.3C, fig. 5 (1925); Schonl. in Rec;
oblongae, 1 ,8-2 x0,6-0 ,8 mm, plerumque pennitus Albany Mus. 3: 151 (1915).
emarginatae. Carpella ovariis tenuibus gradatim in The dulous red flowers simuiate those of species
stylos et shgmatibus termina ibus constncta; ovarium of CotZedo and the similarity is so striking that early
35 ovuhbus quoque 3-plo longiore latum et cristis taxonomists piaced this species rather hesitantly into
e icalibus. the genus Adromisehus. A lectotype was selected,
Type: Cape, near Graaff Reinet, Bolus 758 (BOL, because the sheet of the holotype contains a few
holo!; K!; SAM!). flowers of Cotyledon orbiculata.
388 NEW TAX A AND NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
4. Sect. Inscisilobati Uitew. in Natn. Cact. Succ. length, enclosed in tube, with filaments usually papil-
J. 7: 70 (1952), pro parte. Type species: A. maculatus lose. Squamae about square, broadest usually below
(Salm-Dyck) Lem. the middle.
Inflorescence straight, with mature buds and flowers Occurring from the Eastern Cape to the South-
erect and with extra-floral nectaries on the pedicels. Westem Cape as far norlh as the Qifb but rare|
Buds with almost cylindrical tube gradually con- west of the mouniains of that area.
stncted into an acute apex. Corolla with tube pale
green, glabrous; lobes narrowly lanceolate but scarcely The flowers of species of this section are super
constricted at the base, spreading to recurved, rough ficially similar to those of the sect. Longipedunculati
and with club-shaped trichomes on lower lobes and except that the club-shaped trichomes are not
in throat, white or pale pink and usually deep mauve restricted to the throat. In live material the dense
on the exposed part on the outside. Stamens unequally waxy bloom on the flowers and the pronounced
long, with anthers of shorter inner ones longer than grooves between the petals immediately distinguish
those of the longer outer stamens, rarely of equal flowers of the sect. Longipedunculati.
Key to Species
Leaves terete or almost so, fusiform...................................................................................................A. mammillaris
Leaves dorsiventrally compressed, oblanceolate to obovate:
Marginal ridge of leaves winged and continued to the base (auriculate):
Leaves evenly and gradually constricted towards the base, i.e. oblanceolate, never with spots:
Corolla tube 8-9 mm long; mountains between Clanwilliam and Vanrhynsdorp..................A. maximus
Corolla tube 10-11 (-12) mm long; karroid vegetation between Graaff Reinet, Steytlerville and
East London
...............
...................................................................................................A. sphenophyllus
Leaves abruptly constricted and subpetiolate, i.e. obovate-spathulate to oblanceolate-spathulate, often
with purple spots.......................................................................................................................A. maculatus
Marginal ridge of leaves keeled and rarely continued beyond the middle of the leaf:
Mature buds curved outward towards the apex................................................................................A. triflorus
Mature buds straight:
Leaves 20-30 mm long; between Grahamstown, Alicedale and Port Elizabeth
.....................
A. inamoenus
Leaves 40-130 (-160) mm long; between Clanwilliam and Vanrhynsdorp
.................................A. maximus
A. inamoenus Toelken, sp. nov. ab A. sphenophyllo on the outside. Stamens 9-10 mm long, with anthers
foliis parvioribus cristis marginalibus brevibus ad of about equal length, with filaments slightly papillose,
apices; ab A. trifloro alabastris rectis et foliis non Squamae about square, 1-1,2 mm long and broad,
angulato-oblanceolatis, immaculatis differt. emarginate, broadest at the middle or below. Carpels
Cotyledon rhombifolia sensu Schonl. & Bak.f. in J. Bot., Lond. with slender ovaries gradually constricted into styles
40:92(1902), partly, as for live plant. with terminal stigmas; ovary with 22-28 ovules with
Sufifrutex ramis decumbentibus vel prostratis (4-) vertical ridges.
6-10 mm in diametro. Folia oblanceolato-spathulata, Occurring on somewhat sheltered rock outcrops
20-28 (-32) mm longa, 15-25 mm lata, obtusa vel usually associated with False Macchia in the vicinity
rotundata interdum mucronata, cuneata vel sub- of Grahamstown.
petiolata, cristis marginalibus ad apices, griseo-viridia
interdum suffusa fusca et immaculata. Inflorescentia A. maculatus (Salm-Dyck) Lem., Jard. Fleur. 2,
thyrsus spiciformis 1 (2) floribus in quoque cyma, pedi- Misc. 60 (1852); Illustr. Hort. 7, Misc. 70 (1870);
cellis (1-) 2-3 (-4) mm longa. Sepala triangularia, C. A. Smith in Bothalia 3: 622 (1939); V. Poelln. in
1,5-2 mm longa, aceria, griseo-viridia. Petala 11-15 Reprium nov. Spec. Regni veg. 48: 99 (1940): Uitew.
mm longa, connata et formantia tubos 8-10 mm in Succulenta (1949) 37. Type: sine loc., Salm-Dyck
longos et virides; lobi elliptico-oblongi, 2-3,5 mm sub Haworth s.n. (OXF, lecto!).
longi, acuti, asperi cum trichomatibus clavatibus corol- Cotyledon maculata Salm-Dyck, Obs. 5 (1820); Haw., Rev. PI.
lae expansae in partibus demissis et in fauce, nivei Succ. 21 (1821); DC., Prodr. 3: 398 (1828); Schonl. & Bak.f. in
suffusi rosei vel malvini praecipue in partibus apertis L°wd'ui4x 92 ^19.02^ c• °hernans Haw., Suppl. 26
„ , •
_
• e .
__
. n irv 1 .i • .. (1819), non Vahl. Type: unknown.
exterions. Stamina 9 10 mm longa, antheris circiter Adromischus rhombifolius sensu C. A. Smith in Bothalia 4:
aequantibus longitudine; filamentis leviter papillosis. 625 (1939).
Squamae plus minusve quadratae, 1-1,2 mm longae et ^ poorly recorded species which seems to occur
latae, emarginatae et latissimae ad medium vel onjy on rocjc outcrops in Mountain Renosterbosveld
infernae. Carpel/a ovarns tenuibus gradatim in stylos an(j jn drier parts of Cape Macchia from near Wor-
stigmatibus terminahbus constrictis; ovarium 22-28 cester eastwards mainly along the mountains south
ovulis cristis verticalibus. 0f the jjttie Karoo to near Uniondale.
Type: Cape, 1 km north of Salem, Tolken 5508 A,., , . .. . .. .
(PRE holo!) Although species in this section are often not
_ ’ A . , , , , easy to distinguish, the type specimen, and in particu-
/^n\U^rUi^X Wlth. decumbent to_ Prostrate branches jar the illustration prepared for Haworth, which is
(40) 6-10 mm in diameter. Leaves oblanceolate- now housed in Kew Herbarium, leave no doubt about
spathulate, 20-28 (-32) mm long; 15-25 mm broad, identity of the species. The undulate apices of
obtuse or rounded and sometimes mucronate at the young leaves are characteristic of that species.
apex, cuneate to subpetiolate, with horny margin at .
the apex and rarely beyond the middle, grey-green mammillaris (L.f.) Lem., Jard. Fleur. 2, Misc.
sometimes tinged brown and without purple spots. 60 (1852). Type: Cape, near Olifantsbad, Thunberg
Inflorescence a spike-like thyrse with 1 (2) flowers *n Herb. Thunberg 11006 (UPS, holo!).
in each cyme, with pedicels (1—) 2—3 (—4) mm long. Cotyledon mammillaris hS., Suppl. 242 (1781); Thunb., Prodr.
Sepals triangular 1,5-2 mm long, sharply acute, 83 (1800); FI. Cap. ed. Schultes 397 (1823); Haw., Suppl. 22
grey-green. Petals 11-15 mm long, fused into a tube (1819); Rev. PI. Succ. 21 (1821).
8-10 mm long and green; lobes elliptic-oblong, 2 -3,5 Thunberg’s type specimen does not seem to have
mm long, acute, rough and with club-shaped trichomes been consulted since the younger Linnaeus published
on lower parts of expanded corolla and in throat, the original description. It is not possible to establish
white tinged pink or mauve especially on exposed part when the much more common but also terete-leaved
H. R. TOLKEN 389
A. filicaulis reached Europe and the confusion be
tween the two species started. In all the earlier publica
tions specimens were distinguished by their leaves
only.
The hybrid A. filicaulis var. marlothii x A. macula
tus is superficially very simlar to A. mammillaris
but can be distinguished by its somewhat dorsiven-
trally compressed leaves, by the anthers protruding
above the corolla tube and by the petal lobes
having a broader expanding base between them.
A. maximus P. C. Hutch, in Cact. Succ. J., Los Ang.
31: 133, fig. 59,60 (1959). Type: Cape, Gif berg,
Hall in NBG 475/53 (Univ. Calif. Bot. Gard. 53.
1106-1: BOL, holo!).
This very robust species is found on lower mountain
slopes between Clanwilliam and Vanrhynsdorp, so
that it cannot be confused with the very similar
A. sphenophyllus from the eastern Cape if the locality
is known.
A. sphenophyllus C. A. Smith in Bothalia 3:624 (1939).
Type: Cape, sine loc., Cooper 2338 (K, lecto!).
Cotyledon rhombifolia sensu Bak.f. in Bot. Refug. 1, t.36
(1869); sensu Schonl. & Bak.f. in J. Bot., Lond. 40, 92 (1902),
pro parte; sensu Schonl. in Rec. Albany Mus. 3: 154 (1915), non
Haw.
Adromischus rhombifolius var. bakeri V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 102, 110 (1940). Type: same as for
A. sphenophyllus. —var. sphenophyllus (C. A. Smith) Jacobsen,
Sukk. Lex. 30 (1970), nomen non rite publicatum quoad sine
typo.
This species usually occurs in dry karroid vegetation
between Steytlerville, Graaflf Reinet and East London,
but it is often also found on exposed rock outcrops
in or near False Macchia. It is in these transitional
areas that it hybridizes with A. inamoenus, which
accounts for the intermediate plants which are occa
sionally found. However, the leaves of A. inamoenus
are usually much shorter, so that the two species can
be readily distinguished.
A. triflorus (L.f.) Berger in Pflanzenfam. ed. 2’
18a: 416 (1930); C. A. Smith in Bothalia 3: 623
(1939); V. Poelln. in Reprium nov. Spec. Regni veg.
48: 103 (1940). Type: Cape, Zeko River, Thunberg
in Herb. Thunberg 11016 (UPS, holo!].
Cotyledon triflora L.f., Suppl. 242 (1781); Thunb., Prodr. 83
(1800); FI. Cap. ed. Schultes 396 (1823); Salm-Dyck, Obs. 6
(1820); Haw., Rev. PI. Succ. 19 (1821); Schonl. & Bak.f. in J.
Bot., Lond. 40: 91 (1902). C. bolusii Schonl. var. karroensis
Schonl. in Rec. Albany Mus. 1:1 19 (1904); 3: 154 (1915). Type:
Cape, Laingsburg, Marloth 2519 (GRA, holo!). C. proeurva N.E.
Br. in Kew Bull. (1912) 276; Schonl. in Rec. Albany Mus. 3:
154(1915). Type: South Africa, sine loc. et leg. (K, holo!).
Adromischus subcompressus V. Poelln. in Reprium nov. Spec-
Regni veg. 44: 62 (1938), pro parte quoad Lauder s.n. A-
subpetiolatus V. Poelln. in Reprium nov. Spec. Regni veg. 44:
61 (1938). Type: unknown. A. procurvus (N.E. Br.) C. A. Smith
in Bothalia 3: 641 (1939); V. Poelln. in Reprium nov. Spec.
Regni veg. 48: 109 (1940).
This species occurs in arid vegetation in the Little
Karoo and in areas further north and east from near
Ceres to east of Oudtshoorn.
5. Sect. Longipedunculati V. Poelln. in Reprium
nov. Spec. Regni veg. 48: 89, 95 (1940). Type
species: A. crist at us (Haw.) Lem.
Cotyledon [sect. Spicatae Harv, FI. Cap. 2: 370
(1862)] group Crist at a Schonl. in Rec. Albany Mus.
3: 152 (1915). Type species: A. cristatus (Haw.) Lem.
Adromischus [sect. Longipedunculati V. Poelln. in
Reprium nov. Spec. Regni veg. 48: 89 (1940)] subsect.
Cristati (Schonl.) V. Poelln. in Reprium nov. Spec.
Regni veg. 48: 89 (1940).
Inflorescence more or less zig-zagging at least when
young, with buds at first adpressed to the stem but
later erect to somewhat spreading, with extra-floral
nectaries usually absent. Buds almost cylindrical but
with a distinctly grooved tube, gradually constricted
into bluntly acute apices. Corolla with tube glaucous-
green, glabrous or hairy; lobes lanceolate, often
slightly constricted at the base, erect, spreading or
recurved, rough and with club-shaped trichomes re
stricted to the throat, white and usually deep red
on the exposed part on the outside. Stamens of
different length, with anthers more or less equally
long, with filaments rarely slightly papillose. Squamae
usually transversely oblong to almost square and
broadest at about the middle.
One group of species occurs from just east of the
south-western Cape to the eastern Cape and another
group is found mainly in Namaqualand and extends
into south-western South West Africa, with a few
populations occurring along the Roggeveld escarp
ment.
The taxa in this section appear to be rather hetero
genous when compared with other sections of
Adromischus. A. leucophyllus from the south-western
Little Karoo has large flowers strongly resembling
those found in sect. Brevipedunculati. It appears,
however, that the species merely simulates the flower
type of that section. The following characters indicate
that it should be rather placed in the sect. Longipdun-
culati: the corolla tube is scarcely widened towards
the apex, a thick bloom is found on the flowers, and
the club-shaped trichomes are restricted to the throat
of the corolla. Furthermore, the closely related A.
subviridis has flowers which are typical of sect.
Longipedunculati.
The flowers of species from the north-western Cape
are usually more rigidly succulent than those of species
from the eastern Cape.
Key to Species and Varieties
Leaves dorsiventrally compressed, i. e. at least 3 times broader than thick at the middle of the leaf:
Corolla lobes (3-) 4-5 mm long; leaves round or almost so..........................................................A. leucophyllus
Corolla lobes 1,5-3 mm long; leaves oblanceolate to obovate but with a distinctly cuneate base:
Leaves oblanceolate and with long cuneate base; apical gland on anthers sessile
........................
A. subviridis
Leaves obovate abruptly constricted to subpetiolate; apical gland on anthers raised about 0,5 mm
above pollen sacs........................................................................................A. marianae var. hallii
Leaves distinctly convex on both surfaces i.e. about as broad as thick, rarely up to twice as broad as thick at
the middle of the leaf:
Leaves usually acute, convex towards the apex on both surfaces or canaliculate above; apical gland on
each anther raised above pollen sacs:
Roots tuberous:
Leaves smooth with marginal ridge horny and usually undulate
.....................A. marianae var. hallii
Leaves verrucose (sometimes only visible under 10 times lens) with marginal ridge slightly raised but
indistinct, not horny and undulate...................................................... A. marianae var. immaculatus
390 NEW TAXA AND NEW COMBINATIONS IN COTYLEDON AN D ALLIED GENERA
Roots fibrous:
Leaves concave above, marginal ridge horny and raised
............................
A. marianae var. marianae
Leaves convex above (rarely somewhat concave towards the apex) and marginal ridge never
horny and raised.......................................................................................A. marianae var. kubusensis
Leaves obtuse, truncate and/or dorsiventrally compressed towards the apex; apical gland on each anther
sessile:
Club-shaped trichomes in throat and on corolla lobes; brown aerial adventitious roots absent on
glabrous stems (rarely with adventitious roots which grow into the soil); leaves glabrous... .A. cooperi
Club-shaped trichomes usually only in the corolla throat; brown aerial adventitious roots densely
covering the stems (if absent then stems hairy); leaves usually with glandular hairs:
Stems 40-80 mm long, without adventitious roots, covered with glandular hairs
......................
A. cristatus var. zeyheri
Stems 20-40 mm long, covered with aerial adventitious roots:
Ridge at apex of leaves narrower than broadest point on leaf; inflorescence and petals with
glandular hairs..................................................................................A. cristatus var. schoenlandii
Ridge at apex of leaf constitutes the broadest point on leaf; inflorescence and petals glabrous,
rarely with a few hairs when young:
Leaf blade 1-1^ times longer than the apical ridge is broad; leaves obtriangular-spathulate
usually with distinct petiole....................................................................A. cristatus var. cristatus
Leaves 2-5 times longer than the apical ridge is broad, narrowly oblanceolate to narrowly
obtriangular, glabrous, rarely with few hairs
................................
A. cristatus var. clavifolius
A. cooperi (Bak.) Berger in Pflanzenfam. ed. 2, Adromisehus cristatus (Haw.) Lem., Jard. Fleur. 2, Misc. 60
18a: 416 (1930); C. A. Smith in Bothalia 3: 632 Í,1852.); c S. Smith in Bothalia 3: 635 (1939); V. Poelln. in
Aim \j r> n n c n • ~ Reprium nov. Spec. Regni veg. 49:60 (1940).
(1939); V. Poelln. in Reprium nov. Spec. Regni veg. K K 6 6 v
49: 60 (1940). Iconotype: Saund. Refug. Bot. t.72 Occurring in widely scattered localities between
(1869). P°rt Elizabeth, Uniondale and Graaff Reinet.
Cotyledon cooperi Bak. in Saund. Refug. Bot. t.72 (1869); var. zeyheri (H arv) Toelken, stat. nov.
Schonl. & Bak.f. in J. Bot., Lond. 40: 91 (1902); Schonl. in Rec. Cotyledon zeyheri Harv., FI. Cap. 2: 377 (1862); Schonl. &
Albany Mus. 3: 153 (1915).—var. immaculata Schonl. & Bak.f, Bak.f. in J. Bot., Lond. 40: 91 (1902); Schonl. in Rec. Albany
in J. Bot., Lond. 40: 91 (1902). Type: Cape, Graaff Reinet, Mus 3: 155 ( 1915). Type: Cape, Kenko River, Zeyher 2571
Rattray s.n. (GRA, holo!). (K, lecto!; S!; SAM!).
Adromisehus pachylophus C. A. Smith in Bothalia 3: 633 Adromisehus zeyheri (Harv.) V. Poelln. in Cact. J. 6: 68
(1939). Type: same as C. cooperi var. immaculata. A. festivus (1938); in Reprium nov. Spec. Regni veg. 48: 98 (1940); C. A.
C.A. Smith in Bothalia 3: 633, fig. 3 (1939); V. Poelln. in Smith in Bothalia 3: 635 (1939).
oSSeK IS iHKk -fSLSïï. K r ay is kiTn Honiy fT 'Si ,ypfcoltoio? ?" ,h?
Poelln. in Reprium nov. Spec. Regni veg. 48: 102 (1940», non repreSenla“VC °f
(Thunb.) Lem. Type: Cape, Halesowen, Herre in SUG 6866 the species solar recorded west ot Uniondale
(Bf). A. halesowensis Uitew. in Desert PI. Life 20: 142 (1948); var. clavifolius (Haw.) Toelken, stat. nov.
in Succulcnta (1950) 36. Type: same as for A. cmeatus V. Poelln. Cotyledon clavifolia Haw. in Phil. Mag. (1827) 274; DC.,
Plants from near Graaff Reinet (Tolken 5534) show Prodr. 3: 399 (1828); Schonl. & Bak.f. in J. Bot., Lond. 40:
some va riation in the shape o f the leaves and the 92 (1902). Iconotype: Haworth plate (K, lecto!). C. nussbaume-
presence or absence of spots on them. However, all sfnelrc et°le%Bt)Jt> d* KaktGes‘ 1: 95 (1936)-TyPe: CaPe*
plants have scattered club-shaped trichomes on the In®, ' . e,s , . , . , _ , . ...
v ,, , , , • • .. . j Adromisehus clavifolius (Haw.) Lem., Jard. Fleur. 2, Misc.
corolla lobes, and even on the specimens investigated 50 (1852); V. Poelln. in Reprium nov. Spec. Regni veg. 49:
by C. A. Smith the reported absence could not be 60 (1940); Uitew. in Natn. Cact. Succ. J. 7: 33 (1952). A.
confirmed. A. pachylophus is thus relegated to the poellnitzianus Werderm. in Reprium nov. Spec. Regni veg. 39:
synonomy of A. cooperi. 270(1936); V. Poelln. in Cactus J. 6: 68, fig. (1936); in Reprium
J J ^ nov. Spec. Regni veg. 48: 97 (1940). Type: Cape, near East
Typical A. cooperi in which the apical ridge is some- London, Kluth s.n. (Bt). A. nussbaumeranus (V. Poelln.) V.
what wider than the remainder of the leaf, has been Poelln. in Reprium nov. Spec. Regni veg. 48: 109 (1940).
recorded from near Aberdeen (Pringle in NBG 779/59) Kakteenkunde (1940) 64’ fig‘
but, except for that character, which is also somewhat ype* P®» . , Al , .
variable in the plants from Graaff Reinet (Tolken Occurring on rocky outcrops between Alexandria,
5534), the plant cannot be distinguished from het Grahamstown and hast London. Near Grahamstown
rest of the species t"6 *eaves become more hairy, but the shape remains
. * . , » , ^ . .. unlike that of var. cristata (Liebenberg 6007).
A. cristatus (Haw.) Lem., Jard. Fleur. 2, Misc. 60 , ■ . „ , . r
(1852); C. A. Smith in Bothalia 3: 635 (1939); The combination A. nussbaumeranus, that of
V. Poelln. in Reprium nov. Spec. Regni veg. 49: 60 £ schuldtianus was not validly published until 1940,
(1940). Type: sine loc., Haworth s.n. (OXF, holo!). becausem 1936 von Poellnitz merely cited the combina-
_ , . tion in the synonomy of Cotyledon nussbaumerana.
Cotyledon cristata Haw. in Phil. Mag. (1827) 274; DC., J
Prodr. 3: 399 (1828); Harv., FI. Cap. 2: 376 (1862); Schonl. in var. schonlandii (P hill.) Toelken, stat. nov.
Rec. Albany Mus. 3. 155(1915). Cotyledon schonlandii Phill. in Flower. PI. S. Afr. 9, t.328
A. cristatus is widespread in the eastern Cape Pro- (1929). Type: sine loc. et leg. in PRE 7944 (PRE, holo!).
vince and is often locally common, but these popula- Adromisehus schonlandii (Phill.) V. Poelln. in Cactus J. 6:
tions are far apart and not easy to find. A continuous 68 (1938); in Reprium nov. Spec. Regni veg. 48: 97 (1940);
variation could not be found. In contrast to A. Uitew. in Natn. Cact. Succ. J. 7: 33 (1952).
marianae, the variation is restricted to local popula- Found on sheltered and shaded rock outcrops in
tions. Nevertheless, the variation is considerable ravines in the Langkloof.
depending on environmental conditions and the age . , . „ WJ.^ . XT A ^ . 0 ¥ n
of the plants as has been recorded for var. clavifolia I in»^a // // ^’m dp
by Dyer 5438 from near Bathurst, and for va r. cristata |8 fig. (1954) Type: C ape, M ontagu, Hall in N BG
by Tolken 5407, 5408 and 5522 from near Graaff »55/33 (AVU 10015, holo.)
Reinet very distinct species k now n from the western
parts of the Little Karoo. The salvershaped corolla
var. cristatus. wjtj1 t^e tu^e nQt broacjenecj towards the apex and
Cotyledon cristata Haw. in Phill. Mag. 0827) 274; DC., ciub-shaped trichomes restricted to the throat
Prodr. 3: 399 (1828); Harv. FI. Cap. 2: 376 (1862); Schonl. in . .. . • , ,£■ ■ c „ n „ . Dl.„t • „
Rec. Albany Mus. 3: 155 (1915); Marl., FI. S. Afr. 2,1: 20, distinguish this species from those o f sect. Brevipe-
t.9,5 (1925). dunculati.
H. R. TÓLKEN 391
A. marianae (Marl.) Berger in Pflanzenfam. ed. 2,
18a: 416 (1930); C. A. Smith in Bothalia 3: 639
(1939). Type: Cape, Clanwilliam, Marloth 3489 (PRE,
holo!; GRA!; K!).
Cotyledon marianae Marl, in Trans. S. Afr. Phil. Soc. 18: 47
(1907); Schonl. in Rec. Albany Mus. 3: 153 (1915).
An extremely variable species occurring from near
Clanwilliam to just north of the Orange River. Recent
collections show a continuous range of variation
leading to distinct extreme forms in isolated localities.
After considerable field studies, it was found that most
of the transitional ranges between extreme forms are
restricted to areas where taxa apparently come into
close contact with one another. As whole ranges of
variation are found within these areas introgressive
hybridization is considered to be taking place. Fig. 2
shows a selection of the full range of variation and
the geographic distribution of certain characteristics,
but it also illustrates why certain species have been
relegated to the synonomy of the four varieties recog
nized. The following species, listed with the number
by which they are indicated on Fig. 2, fall within the
range of variation of the specimens cited in the cap
tions: 2, A. hallii; 5, A. geyeri; 6, A. kubusensis,
A. rodinii; 9, A. blosianus; 13, A. antidorcadum,
A. alveotatus; 14, A. herrei; 18, A. marianae var.
immaculatus; 25, A. marianae var. marianae.
var. marianae.
Cotyledon marianae Marl, in Trans. S. Afr. Phil. Soc. 18: 47
(1907); FI. S. Afr. 2,1: 17, t. 3A (1925); Schonl. in Rec. Albany
Mus. 3:153(1915).
Adromischus marianae (Marl.) Berger in Pflanzenfam. ed. 2,
18a: 416(1930); C. A. Smith in Bothalia 3: 639 (1939); V. Poelln.
in Reprium nov. Spec. Regni veg. 48:109(1940), partly.
Var. marianae is found on dry lower slopes from
Clanwilliam to just south of Vanrhynsdorp. Plants
from Clanwilliam are all spotted on the leaves, but
along the Doring River plants with spotted or
unspotted leaves are found.
var. immaculata Uitew. in Succulenta (1953)
10, fig. Type: Cape, Vredendal, sine leg. in SUG
5932 (AVU 10014, holo.).
Cotyledon herrei W. F. Barker in S. Afr. Gard. 21: 247
(1931). Type: Cape, Nutabooi, Herre in SUG 5800 (BOL,
holo!; G RA!; K!).
Adromischus herrei (W. F. Barker) V. Poelln. in Reprium nov.
Spec. Regni veg. 44: 62 (1938); 48: 90 (1940). A. antidorcadum
V. Poelln. in Reprium nov. Spec. Regni veg. 44: 61 (1938),
"anticordatum”; in Cact. J. 7: 19 (1938), “antidorcatum” ;
in Reprium nov. Spec. Regni veg. 47: 2 (1939), “antidorcatum”;
Airy Shaw in Kew Bull. 14: 310 (1960). Type: Cape, 48 km S of
Springbok, Triebner 1324 (BOL, clono!). A. aveolatus P. C.
Hutch, in Cact. Succ. J., Los Ang. 28: 183, fig. 150, 1 (1956).
Type: Cape, 14 km N of Concordia, Hall in NBG 194/52
(Univ. Calif. Bot. Gard. 54.1161: PRE!; UC!).
Widespread in southern Namaqualand and exten
ding its distribution range to the western slopes of the
Roggeveld Mountains.
Neither the verrucose surface of the leaves nor the
tuberous roots, the characters by which the taxon is
at present distinguished, are mentioned in the original
description of the var. immaculata. However, the figure
clearly shows that the roots are swollen and the leaves
are without a horny ridge, which distinguishes it
from var. marianae the only other variety known to
occur near Vredendal.
var. kubusensis (Uitew.) Toelken, stat. nov.
Adromischus kubusensis Uitew., Succulenta (1953) 7 (Jan.
1953). Type: Cape, Kubus, Herre in SUG 6104 (AVU 10012,
holo). A. rodinii P. C. Hutch, in Cact. Succ. J., Los Ang. 25:
136, fig. 106-8 (Sept./Oct. 1953); Friedr. in Prodr. FI. S. W.
Afr. 52: 4 (1968). Type: Cape, near Hellsberg, Rodin 1617
(Univ. Calif. Bot. Gard. 50.1181: UC; holo!). A. blosianus P. C.
Hutch, in Cact. Succ. J., Los Ang. 29: 35, fig. 23,4 (1957).
Type: Cape, Holgat River, Hall in NBG 723/53 (Univ. Calif.
Bot. Gard. 54.111-1: BOL, holo!; NBG!; K !; PRE!; UC!).
A. geyeri P. C. Hutch, in Cact. Succ. J., Los Ang. 32: 89, fig. 45,
47 (1960); Friedr. in Prodr. FI. S. W. Afr. 52: 3 (1968). Type:
South West Africa, Sperrgebiet S of Liideritz, Geyer in SUG
30216 (Univ. Calif. Bot. Gard. 56.826-1: BOL, holo!; UC!).
Restricted to the western slopes of the mountains
in Richtersveld and south-western South West Africa.
var. hallii (P. C. Hutch.) Toelken, stat. nov.
Adromischus hallii P. C. Hutch, in Cact. Succ. J., Los Ang.
28: 144, fig. 111-5 (1956). Type: Cape, Buchu Twins, Hall in
NBG 75/33 (Univ. Calif. Bot. Gard. 53.1115-2: BOL, holo!;
PRE!; UC!). A. nanus (N.E. Br.) V. Poelln. in Desert PI. Life 10:
228, fig. (1938), pro parte; in Reprium nov. Spec. Regni veg. 48:
92 (1940), pro parte quoad Triebner 1318. A. casmithianus V.
Poelln. in Beitr. Sukkulentenk. (1940) 64, nomen non rite
publicatum.
Occurring mainly in rock crevices of hills within
30 km from the coast of the northern Richtersveld
and south-western South West Africa.
The dorsiventrally compressed leaves and the
almost square squamae described for A. casmithianus,
indicate that the collection Triebner 1318 must be
identified as var. hallii. However, it must have been
a very depauperate plant as its leaves are very small
and single-flowered inflorescences are rare in A.
marianae. It is unfortunate that no further specimens
of this plant could be traced and that consequently
some uncertainty remains about the identity of the
species concerned. However, the collection Roux 525
(BOL) would suggest that A. casmithianus belongs to
the A. marianae-o.om\Aex rather than to A. nanus.
The detailed descriptions of A. nanus published in
1938 and 1940 were obviously based on the collection
Triebner 1318, the type of A. casmithianus. As both
publications are later than 1935 and not accompanied
by a Latin diagnosis, the new combination A.
casmithianus, which is based on a wrong identification
without a Latin diagnosis, cannot be accepted.
A. subviridis Toelken, sp. nov. ab A. leucophyllo
foliis oblanceolatis basibus longis cuneatis et tubo
corollae cylindrico; ab A. marianae var. hallio foliis
oblanceolatis basibus longis cuneatis ramis decum-
bentibus longis usque ad 0,25 m longis et in quoque
anthera glande apicali magna et sessili differt.
Suffrutex multiramosus, ramis decumbentibus vel
erectiusculis usque ad 0,25 m longis et radicibus
fibrosis. Folia oblanceolata, 30-55 (-60) mm longa,
14-20 (-25) mm lata, acuta, cuneata, crista marginale
cornea pallida, dorsoventraliter compressa sed leviter
convexa supra et subtus, subviridia vel flavo-viridia,
glauca et maculatis rubellis inordinatis vel absentibus.
Inflorescentia thyrsus spiciformis, cymis 1 (2) floribus,
pedicellis 4-6 mm longis. Sepala triangularia, 2-3 mm
longa, acuta, glauco-viridia. Petala 12-13 mm longa,
connata tubo 9-10 mm longa, glauco-viridia; lobi
lanceolati et leviter constricti ad basim, acuti, succu-
lenti, asperi cum trichomatibus clavatis fauce, albi,
pallide flavi vel leviter suffusi rosei. Stamina 7-10
mm longa, inaequalia, antheris circiter aequantibus
longitudine et glandibus terminalibus sessilibus.
Squamae transverse oblongae 1-1 ,3 X 1,3 -1,6 mm,
leviter emarginatae et latissimae ad medium. Carpella
ovariis tenuibus, gradatim in stylos stigmatibus ter
minalibus constricta; ovarium 26-30 ovulis cristis
verticalibus.
Type: Cape, Bloukrans Pass, Tolken 5349 (PRE,
holo!).
Suffrutex much branched and mainly from the base
with decumbent or scrambling branches up to 0,25 m
long and with fibrous roots. Leaves sessile, oblanceo
late, 30-55 (-60) mm long, 14-20 (-25) mm broad,
cuneate, acute, with pale marginal ridge horny,
dorsiventrally compressed but slightly convex on
both surfaces, pale green to yellowish-green with
392 NEW TAXA AN D NEW COMBINATIONS IN COTYLEDON AND ALLIED GENERA
PORT
‘SPRINGBOK
Fig. 2.—Amplitude of variation in shape, size and surface structure of leaves and the type of roots in Adromischus marianae.—var.
hallii: 1, van der Merwe in PRE 57 053; 2, 2a, Hall in NBG 75/53; 3, Tolken 5274.—var. kubusensis: 4, Tolken 5292; 5,
Tolken 5305; 6, 6a; Tolken 5313; 7, Tolken 5318; 8, Tolken 5260; 9, Tolken 5269;—var. immaculatus\ 10, Tolken 5258; 11,
Tolken 5250; 12, Tolken 5488; 13, Tolken 5200; 14, Acocks 19422; 15, Tolken 5478; 16, Tolken 5473; 17, Tolken 5452; 18, Tolken
5492; 19, Tolken 5444; 20, Bruins s.n.; 21, Bruins s.n.; 22, Hanekom 1447; 23, 23a Tolken 5353.—var. marianae: 24, 24a Tolken
5144; 25, Tolken 5136 (Qroots tuberous; o roots fibrous; A leaf surface smooth; A leaf surface verrucose).
H. R. TÓLKEN 393
faint bloom and with irregular reddish spots present
or absent. Inflorescence a spike-like thyrse with 1
(2)-flowered part-inflorescences, with pedicels 4-6 mm
long. Sepals triangular, 2-3 mm long, acute and
covered with a thick bloom. Petals 12-13 mm long
fused into a tube 9-10 mm long, green and covered
with a thick bloom; lobes lanceolate and slightly
constricted at the base, acute, slightly fleshy, rough
and with club-shaped trichomes in the throat, pale
yellow often with slight pink tinge. Stamens un
equally long, 7-10 mm, with anthers equally long and
each with a large sessile apical gland. Squamae
transversely oblong, 1-1 ,3X 1,3-1,6 mm, faintly
emarginate and broadest at about the middle, white.
Carpels with slender papillose ovaries gradually con
stricted into erect styles with terminal stigmas; ovary
with 26-30 ovules each with vertical ridges.
The shape and the pale glaucous colour of the
leaves are so distinctive in this section that the
species was described from a single collection.
Species insufficiently known
Adromischus rhombifolius (Haw.) Lem., Jard.
Fleur 2, Misc. 60 (1852).
Cotyledon rhombifolia Haw. in Phil. Mag. (1825) 33; DC.,
Prodr. 3: 398 (1828); Schonl. & Bak.f. in J. Bot., Lond. 40:
92 (1902).
No specimen or illustration was preserved of a
plant which Haworth identified as Cotyledon rhombi
folia. He had not seen flowers and the description of
the leaves could refer to those of A. alstonii, A. lieben-
bergii, A. roaneanus or A. triflora.
A. robustus Lem., Jard. Fleur. 2, Misc. 60, fig.
1-5 (1852).
Several illustrations of the flower of A. robustus
indicate that the species belongs to sect. Adromischus.
Unfortunately, no specimen could be traced and the
leaves are merely described as broad. This could
apply to A. alstonii, A. roaneanus or even forms of
A. hemisphaericus, depending on which other plants
Lemaire compared his material with.
(JITTREKSEL
As ’/i gevolg van 'n herwaardering van Cotyledon en
Adromischus is dit nodig geag om die afbakening van
Cotyledon te wysig en die nuwe genus Tylecodon te
beskryf. 'n Verkorte hersiening van Adromischus
tesame met sleutels tot die seksies, spesies en sub-
spesifieke taksons word aangebied. Die volgende nuwe
name en nuwe kombinasies word publiseer: Adromis
chus sect. Boreali Toelken, A. cristatus var. clavifolius
(Haw.) Toelken,— var. schonlandii (Phill.) Toelken, —
var. zeyheri (Harv.) Toelken, A. fallax Toelken, A. fili-
caulis subsp. marlothii (Schonl.) Toelken, A. inamoenus
Toelken, A. marianae var. hallii (P. C. Hutch.) Toelken,
var. kubusensis (Uitew.) Toelken, A. schuldtianus
subsp. juttae (V. Poelln.) Toelken, A. subviridis Toelken,
A. umbraticola subsp. ramosa Toelken, Tylecodon
buchholzianus (Schuldt & Steph.) Toelken, T. cacalioides
(L.f) Toelken, T. decipiens Toelken, T. fragilis (Dyer)
Toelken, T. faucium (V. Poelln.) Toelken, T. grandiflo-
rus (Burm.f.) Toelken, T. hallii (Toelken) Toelken,
T. hirtifolium (W. F. Barker) Toelken, T. leucothrix
(C. A. Smith) Toelken, T. occultans (Toelken) Toelken,
T. paniculatus (L.f.) Toelken, T. pearsonii (Schonl.)
Toelken, T. pygmaeus ( W. F. Barker) Toelken, — var.
tenuis (Toelken) Toelken, T. racemosus (Harv.) Toel
ken, T. reticulatus (L.f.) Toelken, —subsp. phyllopodium
Toelken, T. rubrovenosus (Dinter) Toelken, T. schae-
feranus (Dinter) Toelken, T. similis (Toelken) Toelken,
T. singularis (Dyer) Toelken, T. striatus (P. C.
Hutch.) Toelken, T. suffultus Bruyns ex Toelken,
T. sulphureus (Toelken) Toelken, T. torulosus Toelken,
T. tuberosus Toelken, T. ventricosus (Burm.f.) Toelken,
T. viridiflorus (Toelken) Toelken, T. wallichii (Harv.)
Toelken, — subsp. ecklonianus (Harv.) Toelken.
REFERENCES
Berger, A., 1930. Crassulaceae. Die Natiirlichen Pflanzen-
familien ed. 2,18a: 412-6.
De C andolle A. P., 1828. Crassulacées. Prodromus systematis
naturalis regni vegetabilis 3: 396-9.
Faegrt, K . & Van der Pijl, L., 197 1. Principles of pollination
ecology ed. 2. New York: Pergamon Press.
Jacobsen, H., 1966. Die Gattung Adromischus. Kakteen, Berl.
17:189(1966).
Jacobsen, H., 1970. Das Sukkulentenlexikon. Stuttgart: Fischer.
Lemaire, C. A., 1852. Genre nouveau de la famille des Cras-
sulacées. Jard. Fleur 2, Misc. 60.
Schonland, S., 1915. The South African species of the genus
Cotyledon. Rec. Albany Mus. 3: 130-156.
Smith, C. A., 1939. A review of the genus Adromischus Lemaire.
Bothalia 3: 613-651.
Troll, W., 1964. Die Inftorescenzen. Vol. 1. Jena: Fischer.
Uitewaal, A. J. A., 1957. Notes on Adromischus. Natn. Cact.
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Uitewaal, A. J. A., 1963. Bemerkungen zur Unterteilung der
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