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Clades of the Cape flora.
Compiled by Linder (2003:
The radiation of the Cape flora, southern
Africa. Biological Reviews 78:597-638) and reproduced here with permission from
the author.
Only “larger” clades are included.
Species numbers for the Cape Floristic Region for each genus were taken from
Goldblatt and Manning (2000a),
and total species numbers for each genus largely follow Goldblatt and Manning
(2000a), but for genera outside the CFR various sources were used,
including Mabberley (1997)
and Leistner (2000).
The arguments for inclusion in the Cape flora are briefly listed, as well as
references on the phylogeny indicating the monophyly of the clade, and also
indicating that the clade is not sister to another Cape floral clade.
|
Clade |
Family |
Total spp. |
CFR spp.
|
Evidence
for CFR root |
References |
|
Ficinia |
Cyperaceae |
c.
60 |
56 |
African
endemic, with most species in CFR. A phylogeny is needed to test whether the
basal clades are in the CFR. |
Muasya, et al. (1998). |
|
Tetratria |
Cyperaceae |
c.40 |
38 |
Western
Cape and Australia |
Muasya, et al. (1998). |
|
Irideae
p.p., including Bobartia, Dietes, Ferraria, Moraea s.l. |
Iridaceae1 |
226 |
136 |
All genera
and most species in the CFR. |
Goldblatt, (1991);
Reeves et al. (2001). |
|
Ixioideae &
Nivenioideae, excluding Patersonia and Geosiris |
Iridaceae1 |
c.900 |
516 |
Four of 26
genera not found in CFR, these are tropical Africa. Many genera extend to
tropical Africa and Madagascar. |
Goldblatt, (1991);
Reeves, et al. (2001). |
|
Disineae
(Disa,
Schizochilus) |
Orchidaceae2 |
168 |
92 |
Cladistic
analyses suggest basal elements are CFR |
Linder (1994);
Douzery, et al. (1999).
|
|
Coryciinae (Pterygodium, Corycium, etc.) |
Orchidaceae2 |
39 |
30 |
All basal
lineages and most species in CFR |
Kurzweil, et al. (1991). |
|
Ehrharta |
Poaceae |
36 |
20 |
With a few
exceptions in winter-rainfall area, some species in Australasia |
Gibbs Russell (1987);
Verboom
(2000). |
|
Pentaschistis – Pentameris - Prionanthium |
Poaceae |
81 |
55 |
Greatest
diversity in Cape, but a phylogeny is needed to estabish which clades are
basal. |
Linder &
Ellis (1990a);
Barker, et al.
(2000).
|
|
African
Restionaceae |
Restionaceae |
350 |
340 |
A
monophyletic clade, with very few, largely unrelated, species outside CFR. |
Linder, et al. (2000);
Linder (2001a).
|
|
Centella |
Araliaceae |
50 |
49 |
Most
species in the CFR, but the affinities of the genus are still unclear. |
|
|
Relhaniinae
(Disparago, Elytropappus, Metalasia, Relhania, Stoebe, and diverse
small genera) |
Asteraceae3 |
170 |
131 |
This clade
is still difficult to define, since no generic level analysis of the tribe
exists that is both complete and robust. I assume the monophyly of the Cape
Gnaphalieae |
Bremer (1976;
1994); Karis
(1989).
|
|
Arctoteae
(Arctotis, Berkheya, Cullumia) |
Asteraceae3 |
199 |
109 |
Found more
on the arid fringes, and might not be a typical Cape flora element |
Bremer (1994). |
|
Ursiniinae
(Ursinia, Athanasia) |
Asteraceae3
|
114 |
80 |
Most genera
in CFR |
Bremer (1994). |
|
Amellus grp
(Felicia, Chrysocoma) |
Asteraceae
- Astereae |
149 |
90 |
7/10 genera
in the CFR, with most species too. |
Bremer (1994). |
|
Lobostemon |
Boraginaceae |
28 |
28 |
All species
in the CFR, sister-genus is Echium in Eurasia |
Hilger &
Böhle (2000). |
|
Heliophila, Brachycarpaea,Cycloptychis, Silicularia, Thlaspeocarpa,
Schlechteria |
Brassicaceae |
77 |
60 |
Mostly
along the arid northern margins. The affinities of the clade are unclear. |
K.
Mummenhoff personal communication. |
|
Bruniaceae |
Bruniaceae |
67 |
65 |
Only two
species in the Eastern Cape and Natal. Related to Ericales s.l., possibly
sister to Sphenostemon. |
Bremer, et al.
(2001). |
|
Prismatocarpeae, Siphocodoneae, Merciereae |
Campanulaceae |
63 |
54 |
These Cape
genera may not form a clade – no generic level phylogeny available. |
Takhtajan (1997). |
|
Erica
s.l. |
Ericaceae |
860 |
658 |
The vast
majority of species are in the CFR. However, it is not clear whether (a) the
Cape species are monophyletic, or (b) basal in the genus. |
Oliver (1991; 2000). |
|
Crotalarieae p.p.
(Aspalathus,
Rafnia) |
Fabaceae4 |
297 |
291 |
Although
many genera of the rest of the tribe have species in the CFR, only
Wiborgia has most of its species there, the rest are more tropical. |
Van Wyk (1995). |
|
Podalyrieae,
Liparieae |
Fabaceae4 |
128 |
120 |
The two
tribes form a clade, the sisters of which are outside the CFR |
Van Wyk (1995). |
|
Psoraleeae
p.p. (Psoralea, Otholobium) |
Fabaceae4 |
104 |
94 |
There is no
evidence that these two genera form a clade: the rest of the tribe is in the
Northern Hemisphere or tropical. |
|
|
Pelargonium |
Geraniaceae |
250 |
148 |
80% of the
species are in the winter-rainfall area, and the basal elements are in the
CFR. |
van der Walt & Vorster (1983);
Bakker, et al. (1999a;
1999b).
|
|
Geissoloma |
Geisso-lomataceae |
1 |
1 |
Endemic
family, closests to the Northern Hemispherical Stachyuraceae, Stachyleaceae
and Crossosomataceae |
Nandi, et al. (1998). |
|
Grubbia |
Grubbiaceae |
3 |
3 |
Endemic
family, probably the sister of Curtisia, and these two the sister
group of Cornaceae. . |
Savolainen, et al. (2000);
Xiang, et al. (2002). |
|
Lampranthus |
Mesem-bryanthemacea5 |
155 |
125 |
Generic
delimitations dubious, possible the whole Lampranthus group should be
included, but here, too the genera are dubious |
Hartmann (1991). |
|
Peneaeceae |
Penaeaceae |
23 |
23 |
The family
is endemic to the Cape, the sister is Oliniaceae, which is widespread in E
and S Africa |
Conti, et al. (1997). |
|
Muraltia, Nylandia |
Polygalaceae |
117 |
108 |
This clade
is embedded within Polygala, thus maybe some of the Cape Polygala
species belong here, too. The clade is almost entirely within the CFR. |
Persson (2001). |
|
Proteae |
Proteaceae |
340 |
264 |
Greatest
diversity in Cape, all genera in the Cape. However, there is considerable
doubt about the monophyly of the African genera of Proteae. |
Johnson &
Briggs (1975);
Hoot & Douglas (1998). |
|
Phyliceae (Nesiota, Phylica, Noltia) |
Rhamnaceae |
152 |
134 |
Two of the
three genera, including the basal genus, and almost all species in the CFR. |
Richardson, et al.
(2000);
Richardson, et al.
(2001a). |
|
Cliffortia |
Rosaceae |
120 |
114 |
Very few
species outside CFR, the sister-taxa either in New Zealand or Europe. |
Helfgott, et al. (2000). |
|
Diosmeae (Agathosma, Diosma, etc.) |
Rutaceae |
276 |
268 |
The
subtribe is almost restricted to the CFR. However, the only phylogeny of the
clade could not retrieve its monophyly, and no detailed phylogenetic studies
have been carried out. |
|
|
Selago |
Scrophulariaceae6 |
190 |
101 |
The
phylogenetic situation in Scrophulariaceae is unclear. |
|
|
Stilbaceae |
Stilbaceae
(including Retziaceae) |
14 |
14 |
This
includes both Stilbaceae and Retziaceae, which are sister to the tropical
African Nuxia |
Oxelman, et al. (1999). |
|
Lachnaea |
Thymelaeaceae |
40 |
40 |
The
relationship between this genus and related genera is not known, so this
might need to be modified. |
|
|
Totals |
|
5885 |
4455 |
|
|
1. Mapping the
areas onto the phylogeny of Iridaceae proposed by Reeves, et al. (2001)
results in an ambiguous estimation of the area of origin of the family. It
remains possible that the family originated in southern Africa, and dispersed
outwards.
2. In the
earlier literature it was suggested that the tribe Diseae (inclusive of Disinae,
Coryciinae, Satyriinae, Brownleeainae and Huttoniaeinae) might be centered in
the Cape flora. However, recent cladistic analyses of DNA sequence variation (Douzery, et al.,
1999) indicated the paraphyly of both Diseae
and Coryciinae, while Disinae and Coryciinae s.s. are clearly centered in
the CFR, Satyriinae and Disperiidae have more species and possibly more
variation in tropical Africa.
3. The
Asteraceae are particularly difficult to separate into Cape and non-Cape clades.
This is partly a result of the dominance of the family in the more arid zones
which flank the CFR, and partly the result of the large number of genera, which
makes an overview difficult. In addition, many of the subtribes and tribes have
a rather wide distribution. Here, the phylogenetically based classification of
Bremer (1994)
is followed consistently, even where more recent phylogenetic hypotheses are
available.
4. The problems
with defining “Cape floral clades” in Fabaceae are rather similar to those in
Asteraceae, with the added problem that there is no recent family overview. In
several tribes, like Crotalarieae, most of the genera have a few species in the
CFR, but most outside. Other genera, like Indigofera, are similar to
Senecio in having a large number of species in the CFR, but are also very
well represented outside the Region.
5.
Mesembryanthema present very special problems. There are far too many species
recognised, and every revision results in a drastic reduction in the number of
species. The generic delimitations are dubious, many species belong to the large
genera Rushia and Lampranthus, these are most likely highly
polyphyletic. The phylogenetic relationships among the genera are unknown. It
seems therefore almost impossible to delimit Cape clades, and to get a
reasonable estimate of the number of species involved.
6.
Scrophulariaceae have largely the same problems as Asteraceae: many genera, many
with a large contingent of species in the CFR, but still inadequately known to
delimit confident Cape clades.
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