Table Of Contentuoa
PHYLOGENETIC
ANALYSIS
Peter Goldblatt^
THE SOUTH
OF AFRICAN
GENUS SPARAXIS
(INCLUDING SYNNOTIA)
(IRIDACEAE-IXIOIDEAE),
TWO NEW
WITH
SPECIES
AND REVIEW
A
THE
OF
GENUS^
Abstract
Kestricted to the western Cape Province of South Africa, Sparaxis comprises 13 species of small seasonal geophytes,
seven
belonging now
to section Sparaxis and six to section Synnotia. Until treated as a separate genus, Synnotia
IS separable from Sparaxis section Sparaxis Sparaxis sensu stricto) only by the degree of zygomorphy of their
(i.e.,
=
nowers.
Species of the two sections have similar karyotypes with x 10; specialized scarious, brown-streaked bracts;
an
unusual anatomy
leaf in which the leaves lack marginal sclerenchyma but have a specialized, heavily thickened
niargmal common
epidermis; and similar fruit and seed characteristics. Vegetatively, they have a gestalt and cannot
be
distinguished. In section Sparaxis the perianth always actinomorphic, but in two species the stamens are
is
asymmetrically
arranged with the anterior stamen opposed to the posterior tepal and the style curved behind In
it.
section Synnotia and
the stamens are unilateral and arcuate with parallel, contiguous anthers, the perianth is
zygomorphic.
Maintenance of Synnotia, based solely on this minor distinction, is unacceptable and is inconsistent
^ith
the variation example, Babiana, Gladiolus, Lapeirousia,
in several other genera of Iridaceae-Ixioideae, for
Iratsoniay
Geissorhiza, and Tritoniopsis, of which include species with either actinomorphic or medianly zygo-
all
morphic
flowers. Cladistic analysis of Sparaxis using either Tritonia or Ixia as outgroups for character polarization
produces
the same more parsimonious regard Tritonia as the immediate outgroup.
internal configuration, but to
it is
unless
characters such as karyotype are weighted. Cladistic analysis shows that the species clusters recognized as
section
Synnotia and section Sparaxis are not phylogenetically equivalent; the two sections are thus recognized for
their taxonomic
utility.
now
Restricted caryophyllacea Synnotia), there are
to the winter-rainfall region of south- (sect.
ern
Africa, Sparaxis a genus of small geophytic 13 species of Sparaxis and four subspecies,
is
perennials
with cormous
rootstocks, a fan of fairly
succulent
leaves,
few-flowered spikes of large, usu-
and Relationships
History
sUy
brightly
colored and brown-
flowers, scarious,
streaked,
now
long-cuspidate regard-
bracts. Until
ed been
as
comprising
just six species with completely -^ ^
,
when
regular 1896: 134; Lewis, 1956) and,
edged (Baker,
perianths
(Baker, 1896; Goldblatt, 1969),
Ker
1804 by John Bellenden
5/>arax/5
described in
here expanded
is to include 5x««o//a as first
section Sparaxis inrkided two of the species
Synnotia. As Synnotia (Ker-Gawler),
a genus, distin-
is
Synnotia by Robert
guished be assigned
to
Sjn-
a (1956: 138) revision of
zygomorphic In Lewis's
upper (1826).
perianth with an enlarged
m
comment
Flora Ca-
tepaL echoed Baker's
she
Including Synnotia, which Lewis (1956) notia,
in
[only]
recognized
new
five species, and adding two spe-
^
^
Although the species
perianth.
cies, Sparaxis maculosa Sparaxis) and S. in its irregular all
(sect.
ion
thi;
Compton Herbarium. Kirstenbosch Botanic
of the
. acknowledged. thank John Rourke and the staff
,
I
and Margo Branch for the .!lu.strat.on«
ardens. South Africa,
for their hospitality during ray several field trips to
used
Garden, P.O. Box 299, St. Loui. i:t
' B. A. Krukoff Curator of African Botany. Missouri Botanical
,
11.S.A.
Ann.
143-159. 1992.
Missouri
Bot. Card. 79:
T
144 Annals of the
Garden
Missouri Botanical
of Sparaxis sensu Goldblatt (1969) have an ac- and SjDara:x:/5 are considered congeneric
(Goldblatt,
tinomorphic perianth, the filaments of S. grandi- 1969). Lewis's treatment was supported by
kary-
and
flora S. bulbifera are unilateral, the abaxial ology (Goldblatt, 1971: 412), genera having
all
=
stamen is curved to lie opposite the adaxial tepal, X 10 and a similar karyotype of small chro-
and the style displaced to between stamen mosomes.
is lie this
&
and
the adaxial tepal. In Synnotia^ the perianth Studies of leaf anatomy (Rudall Goldblatt,
is zygomorphic with at least the upper tepal en- 1991) have now cast doubt on the validity of
Ixi-
larged and differentiated from the other five, and inae. Several genera of Ixioideae have a derived
stamens
the are unilateral with the lateral filaments leaf margin structure with columnar epidermal ceDs
lying close to the displaced abaxial stamen so that with very thick anticlinal walls and without sub-
the anthers are and more
parallel or less contig- epidermal sclerenchyma. These include Sparaxis
The
uous. difference in zygomorphy between Syn- (and Synnotia), Crocosmia, Chasmanthe, most
notia and the above two Sparaxis one
species species of Tritonia few have marginal epidermal
is (a
of degree and is not a valid criterion for generic cells matching those of the laminar epidermis),
separation. In other genera
of Iridaceae, for ex- Freesia, and Anapalina. Tritoniopsis (Goldblatt,
&
Babiana
ample, and
in Gladiolus, both indisput- 1990; Rudall 1991) has columnar
Goldblatt,
edly monophyletic genera, there are several species and thickened marginal epidermis, but some species
with completely actinomorphic and
flowers a great- also have subepidermal marginal sclerenchyma. Ixia
number
er with zygomorphic flowers (Lewis, 1959; and Dierama have unmodified marginal epidermal
Lewis 1972). Wats
et al., and subepidermal sclerenchyma, both consid-
cells
each have
a single actinomorphic
species that is ered basal features for Iridaceae. In addition. Die-
often segregated subgenerically (Lewis, 1958; rama has an anatomy, and
unspecialized leaf its
Goldblatt, 1989) or sectionally from zygomor-
its leaves usually lack midribs. In this paper I develop
phic congeners. There
are similar examples in Lap- a phylogeny of Sparaxis using both Ixia-Dierama
eirousia.Geissorhiza, and
Tritonia. It is clear thai and Tritonia as possible outgroups for polarizing
zygomorphy
has evolved
repeatedly in several characters. The choice of outgroup makes no dif-
monophyletic may
lines in Iridaceae-Ixioideae and
ference to the structure of the tree (although the
&
even have been some
lost in cases (HiUiard Burtt, evolution some and the im-
of characters
differs),
1991: therefore an unsound
36):
it is criterion for mediate Sparaxis remain
generic relationships of
the recognition of a genus. This
particularly so
is uncertain,
in the case of Sparaxis and Synnotia, where
there
a clear progression from actinomorphy
is com-
to
Review of Taxonomic Characters and
zygomorphy.
plete floral
Character
Polarization
Sparaxis and Synnotia
are so
similar vegeta-
CORM
tively that they cannot be TUNICS
distinguished except
in
The
flower.
floral bracts, often good
indicators of
corms are
In most species the relatively small
relationship
in Ixioideae, are
similarly
specialized
outgroup
enclosed which
in fine, pale, netted fibers,
They
in both. are scarious and more
or
less pale-
suggests
comparison
r with either Tritonia or Ixia
membranous
but
T7 are
i"rregularly streaked with
^
'"''''
^ rox-
the basal Synnotia, S.
condition. In section
'''
h n n
brown, and .
are often lacerate and
lone-cusped. .•• j
q- :Ur I ^ ^ bl,urghii and Sc. gal7eat. a hu ave .tuhe *tunic ^evxttPennddlinngg
bimilar bra„ct,s occur i. n t,,he tropi.cal, and, ,
southeast-
„^ j j u *u^
upwa„rd • r npliaanitus^
oe™m in a nec1k around t1he base of the
Af.;^o„ n-«
Atncan Dierama, c L- T
which . , .
for reason has been
it J^a m olW Hdeennssee
®"" t*V",^^ »tuni• cs accumul1ate • an unusually
c
-^lot^j
,c'onn.rs^i^\dAe^r,-e^dA related * Sparaxis• . T T . , _ ^
to In though, t
Ixia, m c -n^^n S
"^^^^' part.i- cul1 arl1 v • t,he l, atter. It n S. viUosa,
tn \^ r«la.^J H"
to be ncllro^sce^lKyr related Dierama ,1 , r..r.[ ^
to „
(Lewis, 19'6"2^)A,
t„nirs
' caryoph.yllacjea,raind,iSr..vari.egata t,he corm tunics
tWhr«rt<.a^o„c.,.ii u iij
the bracts are usually pale-membranous
1 and
short-
cu^rc
". " TT^ " es-
\^^ll
cusped. These
four genera have
angular ,o globose 1
.
'k'^
.
'^''^'"^ "
5,""
''''""'•
seeds »i.h a surface whieh ''' *T „''
in the outline of the
7"
^" "''''"''''T e^.
structure . probably 4n m,portant s
epidermal
cells is nearly ,o completely
obliterated,
'*"''
"''''""'°" ""''?"
"""'"«
"""^ '"'
a
fairly typical seed type IxioLae
in In Sparaxis '\
^"'"
""«'' "^"^^^
and Synnotia "" "
the seeds are completely
spherical
and
perfectly smooth.
LEAVES
Lewis
assigned Synnotia
together with Sparax-
Streptnntheray and Dierama
is, Ixia,
to subtribe
describe
Ixiinae of the Ixieae. Since then
composition
of
distinguished
Ixiinae has not changed except
that Streptanthera but
Iridaceae somewhat (soft
in their succulent
Number
Volume
79, Goldblatt 145
1
1992 Review and
Phylogenetic Analysis
Sparaxis
of
They
firm) nonfibrotic texture. are always a rather markably similar to those of Dierama, which
is
and presumed
pale green, the fine, pale lateral veins are to be convergent. Bracts of Ixia and
perhaps part of the reason for their being easily Tritonia are typically membranous and uniformly
seem
recognized. In sunlight the leaves sprinkled pale or with the veins a darker color, and in both
with gold, probably a reflection of the light from genera the bracts are most often short-cusped.
the epidermal papillae. Most species of the genus Three species of section Sparaxis stand out in
have
also obtuse leaves generally obliquely apic- having especially large but non-cusped bracts. In-
ulate, unlike both possible outgroups in which the stead, the margins are rather irregularly lobed. This
The assumed
condition here to be the plesiomorphic
is
have a basis in internal anatomy, but in any event state. In S.fragrans and S. maculosa the margins
should be regarded as derived. The leaf margin of the bracts including the cusps are irregularly
The enough
epidermis of the several species examined anatom- fringed (Fig. 1 A). condition is distinctive
As make from
ically is unusual. in the case of Tritonia and to possible to identify the species the
it
close Crocosmia and Chasmanthe{de Vos, bracts alone and must be regarded as apomorphic.
its allies
1982, 1984) and
Tritoniopsis (Goldblatt, 1990),
the marginal epidermal from laminar
the
cells differ
PERIANTH
iunuiar
The somewhat perianth actinomorphic
ened on The fleshy
the margins lack sub- is
anticlinal walls.
Sparaxis, but medianly zygomorphic
epidermal sclerenchyma, as do the margins in the in section is
&
Syn-
a derived condition, in section
Tritonia group but not Tritoniopsis (Goldblatt (Figs. 2, 3),
nnm notia. in which the flowers face to the side, the
leaf margin apomorphic upper tepal always largest, and the lower three
features are regarded as is
one another
and tepals are typically smaller, united to
strongly link Sparaxis to the Tritonia group,
AU
and grooved proximally.
especiaUy because morphology equivocal con- for a short distance,
is
cemin members of section Sparaxis have a fairly short
most thought
irelationshios in Ixioideae./xm,
do Synnotia.
to be allied to Sparaxis (Lewis, 1962; Goldblatt, perianth tube, as several of section
An roxburghii and
elongate tube characterizes S.
1969), has margin anatomy,
the plesiomorphic
leaf
variegata, and the
two subspecies of in
with the epidermis no on margin and lam- the 5.
different
and the tube bent sharply (geniculate) at the
ina, weU-developed subepidermal sclerenchy- latter is
ma top of the lower cylindric part of the tube (Fig.
associated with a marginal vein.
3B).
Sparaxis
section limited.
Floral variation in is
INFLORESCENCE AND
BRACTS
and
seven species have a regular perianth, five
All
stamens
There have actinomorphic flowers with erect
nothing remarkable about the inflores-
is
around
cence
of Sparaxis, a spike, but the patterns of sy
Two have asymmetrically arranged
sta-
branching members species
are variable. In most of section
below.
Sparaxis mens, described
branching the base,
restricted to al-
is
Synnotia the tepals are
though may In the flowers of section
plants produce up axes per
to five
eorm
and unequ
this seems to be the basic condition in
and
distance,
united for a
th e genus. In and Synnotia, lower three small, ..Iiurt
S. bulbifera section
more
and or
stamens are unUatcral
branching, when presumably grooved. Tlie
occurs, cauline,
it is
a lying close to the pot,terior tepal.
contiguous,
reversal to the basic kind of branching in Iri- less
orange or pink, foimd
of red to
<laceae Perianth colors
including the possible outgroups of the ge-
and pillansn,
elegans,
nus. Sparaxis tricolor, S. 5.
Flowers are more or spiraUy arranged on in
less
correspondmg as they
plesiomorphic,
a probably
flexuose axis section Sparaxis, but the spike are
in
predominant
secund
*s zygomorphic
in those species with a peri-
Sparaxis have
The remaining species of
may
anth and
(sect. Synnotia). The two characters be Jxia.
purple perianths.
white or
correlated, but are here regarded as separate syn- yellow to
odor
apomorphies.
three fragrant hpecies,
the
The most because the scents in
perhaps the
bracts of Sparaxis are
and galea
caryophyllacrn, S. fa,
striking They dry and S.frfigrnns, S.
are
feature of genus.
the
...
and thus doot evidence for rela-
seem
firm in texture, somewhat creased, and usually different,
on o.h.r rr.tena. the
hasrd
Moreover, all
long-cusped they are irreg- lionshlp.
(Fig. In addition,
1).
..rm .mm.d.atHy
do not tu lie
scented '^pecie^
ularly medium dark brown on the three
streaked
with to
related.
pale The are
translucent background. bracts re-
146 Annals the
of
Garden
Missouri Botanical
\
i
I
I
I
I
I
I
I
I
yf//
I
Figure
1.
axis maculosa (A) and xO.5;
S, hulhifera Habits singi'
(B).
I
and
separated inner and
outer bracts (Drawn by Manning
full size. C.
J.
stamens and
STAMENS AND tiie style are asyrametricall
STYLE
The
lower
(abaxial) stamen curved
is
Whereas
most
in species of section Sparaxis and
stamens
the arc arranged
symnietrically around
bet
the
TfMitral grandijiora and
style, in 5.
S. bulblfera
This
Number
Volume
79, Goldblatt 147
1
1992 Review and Phylogenetic Analysis
Sparaxis
of
Habits x0.5;8tngl
Figure
2.
and (Dr
inner outer bracts full size.
A
branchrs are derived. similar inlerpretarion
short
uni
seems reasonable with Ixia a^ uutgroup.
stamens
In and section
S. graadifiora and bulbifera
S.
POLLEN
Synnotia
the anthers face inward and arc introrse
at of section
anthesis, unlike the other species
The grains of several species
pollen
Sparaxis, which have extrorse anthers, the ba&al banded operculum
anthers
condition the
for Iridaceae. In 5. pillansii verrurate
iinutely tec-
elegans
«^e and
slightly coiled (S shaped), in 5. «:• In features
xh^'-o
all
i.
they (Gold-
are strongly twisted around the style lively to the presumed
blatt, 1969).
basal
Three
Sparaxis^ tricolor^
species of section S. ually small grains
5. elegans. and have short, terminally
S. pillansii, banded oper
single
expanded from the
style branches, different fili-
form, charac-
branches that
relatively long style SFFW
AND
FRUIT
Com-
terize the genus (GoldWatl, 1969).
rest of the
rnpules
The
ihin-wttll.d
r^rwon with Tritonia suggests that the long filiform
hftpH
!»arT^I—
less
and the
that
•tylc branches are the basal condition
148
Annals
of the
Garden
Missouri Botanical
^rf
I
1
t
I
Figure
Habit and
3. floral details of Sparaxis variegata metelerkampiae (B).
subsp. variegata and
(A) subsp.
Tawn
n
lubglobose
en seen from The
above. capsules with a greater number of seeds than the
membrano
^alls of the ripe capsules average (10-12 a little
per varies
Size
locule).
ded
impression of the seeds, between the species, but the large seeds of 5.
a consequence mm
of seed shape and
texture rather up apo-
parviflora, to 2.5 diam,, are clearly
than any
of
intrinsic feature
of the capsule morphic. The smaU subsp.
variegata
itself. seeds of
S.
The
seeds of both mm
sections Sparaxis and Synnotia
metelerkampiae, probably
.6- are
diam.,
1 .8
1
are spherical, hard, and The
shiny. raphe remains also apomorphic. above two taxa,
Curiously, the
conspicuous, and
there is a sHght swelling around one with many smaU with few
and other
seeds the
the micropyle. Color medium brown
deep
is to red- large seeds are most self-com-
also the strongly
Seed numbe
patible and autogamous genus.
the
species
in
an 8-10
species, with average
of seeds per
locule
seeds
KARYOLOGY
//7
per locule) and variegata
S. subsp, variegata (5- "^
Basic chromosome number Sparaxis ^
in is
6 per locule), while subspecies metelerkampiae
has 10 are
(Goldblatt, 1969, The chromosomes
1971).
Number
Volume
79,
1 Goldblatt
149
1992
Review
and
Phylogenetic Analysis
of Sparaxis
and
small exhibit a small range in size. Original roxhurghii renders informed comment on the tree
counts here two
for species of section Synnotia, impossible.
caryphyllacea and A
variegata
5. 5. subsp. mete- notable parallelism in the tree the elongate
is
=
lerkampiae, both conform
f 2/i 20, to the pattern perianth tube in 5. roxhurghii and 5. variegata^
already described A
for the genus. This karyotype species which have different types of corms.
accords closely with that of Ixia and Dierama, longer tree results when these two species are treat-
=
Tritonia and Crocosmia have 11^ but oth- ed as related by their long tubes, for there then
:r
is
erwise have a similar chromosome morphology. It a reversal in corm tunic texture and two parallel-
= tox=
only a small change from x
is 11 10, isms, one for an erect upper tepal and another for
and there ample
is precedent for single-step de- tunics thick and forming a neck. Other parallelisms
creasing
dysploid events
in flowering plants, in- in the tree, including a small flower in S, parvifiora
eluding
several within Iridaceae. and S. villosa^ spotted leaf bases in 5. galeala and
S. caryopkyllacea, and few and large seeds in S.
and galeata, are not unusual
parvifiora S. for
The
plants in general. presence of cauline branch-
Phylogeny
of Sparaxis
Synnotia group and
ing for the clade including the
The
characteristics outlined above and sum- S. hulbifera treated as a reversal, basal b h.
is
manzed
> in Table 1 form the basis for the cladistic ing being regarded here as apomorphic for Sparax-
analysis. Characters were polarized using The reversal of one of the basal characters for
initially
is.
both A
Ixia and Tritonia as the outgroup. more Sparaxis, oblique apiculate leaf apices, in S. fra-
parsimonious
resolution was obtained with Trito- grans not surprising in view of its narrow leaves,
is
The
nia. use
of Ixia as outgroup founded largely which leave opportunity for the expression of
little
is
on
karyotypic
and
similarity, the resulting clado- the character.
gram
thus assumes the convergent development of The analysis confirms the integrity of Synnotia
the two The
evidently unlinked and striking leaf ana- as a monophyletic assemblage. Synnotia clade
tomical
characters, columna supported by three synapomorphies, related
all
is
and
cells, lack of subepidermal epidermis at the to floral zygomorphy and thus perhaps intimately
^i^argins in Sparaxis and the Tritonia lineage. linked. The analysis also indicates that while the
in
I prefer to assume genus Sparaxis monophyletic and supported
that a single decreasing dysploid is is
event
occurred Sparaxis by synapomorphies, none evidently linked, the
in the resulting in five
line,
the
apparently identical karyotypes of Ixia and seven species of Sparaxis sensu Goldblatt 1 969),
(
Sparaxis. The cladogram was produced excluding Synnotia, do not themselves con-
(Fig. 4) i.e.,
but rather consist of four
enni stitute a single lineage,
puter more equivalent the Synnotia
analysis produced three trees of equal length, clades, or less to
the one
figured, a second that not supported by clade.
is
the data, and Treatment of these seven species as a section
a third that discussed below, that
is
have a Sparaxis taxonomically convenient, although
tree length of 31 and a consistency index is
(CI) of phylogenetically unacceptable because section
0.77, ignoring non-informative characters
The
for the Sparaxis so constituted paraphyletic.
ingroup. Ignorance of seed type (character as is
25) in S. roxhurghii and variegata makes recommended phylogenetic solution, to recognize
5.
it
^possible sec tions
to resolve completely the Synnotia clade.
Depending on two section Synnotia (or no sections at all) has little
the character these
state for
species Excepting the basal clade that
a shorter tree leneth of 30 and CI of 0.8 practical merit. ui-
imeag
many
e
obtained assuming
(as figured),
is
synapomorphy
small supported by only one each.
are
On
for
5. roxhurghii.
rhe
most araxis
significant point of difference bet
must
Sparaxis.
the and paraphyletic section It
three computer-generated the position residual
trees
is
hue
«f Sparaxis be noted that within the latter, the 5. tricolor
clade
parvifiora. position as sister
Its
remauung
to lour species
the
remaining Synnotia
species of section (Fig.
y primary clades.
where three
defined by two parallelisms the one
it is is
related that they are in practice
fact so closely
ifi
disimpmh, and as
their validity
clade
to
to 5. galeata-S. roxhurghii, and the cluster often difficult
been
has not alwa)
*8 supported by the presumed presence of few large species .^
^^^s
gnit!
(character 2S). Ignorance of the seeds of S.
150 Annals of the
Garden
Missouri Botanical
SPARAXIS
r
24 bases
leaf
23 flower small
spotte d
24 bases
leaf
25 seeds few
spotted
17 tube elongate
7 tube elongate
1
13 anthers twisted
3 tunic necked
26 throat streaked
8 upper tepal
1
2 tunic fibers coarse
recurved
23 flower
small
25 seeds tew and large 21 perianth partly purple
perianth bilabiate
1 1
5 stamens arcuate
1
19 secund
spike
22 branching cauline
14 stamens asymmetric
Tritonia
Crocosmia
9 bracts cuspidate
Chasmanthe
12 perianth yellow to white
Freesia
Anomatheca
Duthieastrum
X = 10
1
6 leaf apices oblique apiculate
7 leaf succulent
_+.
8 bracts scarious
22 branching basal
4 leaf margin epidermis columnar
5 leaf margin w/o sclerenchyma
and
Figure 4. Phylogeny of Sparaxis, with Tritonia and {Crocosmia, Chasmanthe, Freesia,
close
its allies
Tree
Anomatheca) as outgroup. The cladogram based on the character and data matrix presented in Table 1.
is list
30 and constructed
length is the consistency index (CI) is 0.8, with autapomorphies excluded for the manually
lines
color has merit, but because of small formal Sparaxis has an actinoxnor-
its size fact that the section
treatment seems unnecessary at present. phic perianth and Synnoda has a zygo-
section
morphic
one.
Review
Systematic
1802,
Sparaxis Mag. 548.
Ker, 15:
Curtis's Bot.
t.
The S
SDecies of
& &
1804 as
225.
Konig Sims, Ann. Bot.
1:
&
full Abb-
1805. Linnaea 32: 747, 1863,
Klatt,
tions are provided in a treatment of the genus
for 1882,
HaUe 389
Naturf. Ges. (Erganz. 56).
15:
Southern
if Afi
Ba-
not including Synnotia or Streptanthera.
synonymy
&
Cap.
Handb. Flora
ker, 196. 1892, in
Irid.
except where new
combinations
are presented. Ex-
or
Synnotia
115, 1896,
6: not including
tended
descriptions are provided two new
for spe-
Bot.
African
Streptanthera.
Goldblatt in S.
J.
maculosa
and
cies, S. caryophyllacea and
S.
for Synnom^
219-252.
35: 1969, not including
variegata, which
S. has a complex pattern of vari-
TYPE SPE-
Contr. Bolus Herb 13: 57. 1991-
Complete
ation. species descriptions and ample
ci- here
Ker
CIES: S. hulhifera (lectotype,
(L.)
tation of specimens are included
in revisions of
designated).
Synnotia
(Lewis, and
raxis
(Goldblatt, 1969). have not included
I sectional
'
descriptions here because they have ^^'^P^^^^hera Sweet, Brit. Fl. Card. ser. 1, 3: t. ^!!
little utility in
(&
Ia 390
...». Q.....;. 1 1827, Klatt, Abh. Naturf. Ges. HaUe 15:
. c^.ll l;to Sparaxis ,„...i^ „„i, r„_
like
(1892)
160
ganz.'56). 1882. Baker, Handbk.
Irid.
repetition of the generic description except Gen
for the
Number
Volume
79. Goldblatt
1 151
1992 Review and
Phylogenetic Analysis
Sparaxis
of
Table 1. Data matrix and character list for the cladogram (Fig. 4). The derived condition is denoted by 1;
ancestral by 0; state unknown by ?. Polarity of characters discussed in the text. Autapomorphies are not included.
is
Abbreviations for species will be evident from the text and Figure 4.
number
Character
Taxon 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26
1
1000000000000000000000
0001
Tritonia
Sparaxis
1111000000010001010000
1001
1000010010001010000
trie. 1001
11000010010001010000
eleg.
1001
1 1 1
10100000000010000
pill.
1001
1 1
101
11
.00111111101000000000100?0
10011111100101000000010000
tnacu.
11100101000000000000
grand.
1001
?0011111101101100010001010
bulb,
1 1
10111111101101100110100110
parvi.
0111111101101101110100070
gale.
11011111101101100010101001
roxb.
11011111001101100010100101
vilL
110111111011011010101000?1
caryo.
varie.
Characters:
the derived (apomorpl
1. basic chromosome number x 10 X 11
corm
2. tunics of coarse thickened below ^tunics fibers fine
fibers
mass—
3. corm tunics forming a neck and accumulating in a dense tunics not forming
—
4. leaf margin epidermis columnar epidermis normal
5. margins sclerenchyma— subepidermal sclerenchyma present
without subepidermal
6. leaf apices obtuse and oblique apiculate apices acute
—
±
7. leaves succulent and gold-reflecting leaves firm, not gold-reflecting
membranous and
8. rious, creased and brown streaked— bracts self-colored
—
9. bracts long-cuspidate and lacerate bracts not cuspidate or hardly scarious
—
10. bract edges edges smooth
irregularly serrate-fringed bract
—
11.
perianth zygomorphic and perianth actinomorphic
bilabiate
12. perianth shades of yellow to white or purple— perianth with red. orange or pink
—
13.
anthers weakly
twisted anthers straight
iynunetri
14. abaxial stamen opposed to adaxial tepal and style arcuate
—
15. stamens
unilateral and arcuate stamens not unilateral
16.
style branches short and broad above style branches filiform
—
17.
perianth
tube elongate perianth tube short
—
18. upper hooded
tepal erect-recurved upper tepal
^
19. spike secund—
spike spiral
,
marked or bicolored)
20. drably (uni-
perianth marked perianth usually
vividly (tricolored)—
marks
purple
21. purple— perianth without
perianth
with at least upper tepal partly
—
22.
branching
basal only branching cauline
23.
flowers small—
especially flowers fairly large
—
24.
leaf bases
spotted leaf bases self-colored
25.
seeds few and —seeds many and moderate-sized
large
^
26.
perianth throat streaked— throat not streaked
globose
Afr. Fl. PI. ed. 2: 218. 1951. TYPE SPECiES: S,
mod
elegans
Sweet (= Sparaxis elegans (Sweet) Gold-
blatt). nneetttteedd fniobeerrss oorr ooif ccooaarrssee hnaarrdu fuiubceis,
^ "'**^M- erately ffiinnee
^ynnotia 1826
Sweet, Card. 150. (as
Brit. FI. 2: t, ^^j clawlike below. Leaves several,
^j^j^j^^^^j
^ynnetial coxx Synnotiahy Sweet in Hort Brit,
.nmPtim^
, but sometimes aallssoo ccaauulliinnee., tthhee llooww-
mostly basal,
Linnaea
membranous
Ann.
Baker, Flora Cap. 134-135. 1896. Lewis,
6:
S. African Mus. 137-151. 1956. type species; remaining laminate and green, lanceo-
40: the
phylls),
S. variegata Sweet (= Sparaxis variegata (Sweet) acuminate.
obtuse to acute,
j^^^ or falcate,
linear
^^
Goldbl
losely
Anactorion 1838 1836).
Raf., Telluriana 34. (as
Fl.
fine
type: a, hicolor (Thunb Sparaxis villosa
±
glahroiui, simple
erect,
and thick,
relatively
firm
(Burm.
N.
E. Br.).
f.)
.A
152
Annals of the
Garden
Missouri Botanical
or branched, then either from near the base or largest and the lower three smallest, lanceolate
to
above the ground, sometimes with a few large ax- obovate or spathulate, acute to obtuse. Stamens
cormlets in inserted at the base of the expanded part
(throat)
Spike
(l-)few-
all axils. to several-flowered, lax, of the tube, symmetrically or asymmetrically
dis-
distichous in bud, loosely spiral or nearly secund posed; filaments filiform, straight and surrounding
in bloom; bracts scarious and creased, pale with the style, or curved outwards and the abaxial
twist-
±
brown
streaks, entire to lightly lacerate, or deep- ed to opposite the adaxial or aU
lie tepal, three
±
ly lacerate with long tapering cusps, the outer filaments parallel and opposite the adaxial
tepal;
than
(abaxial) larger the inner (adaxial). Flowers anthers straight, curved or spirally coiled,
sxib-
actinomorphic or zygomorphic, then Ovary
with
either basifixed, extrorse or introrse. ovoid; style
stamens
the alone asymmetrically arranged or the filiform, straight and erect or unilateral, the branch-
perianth also asymmetric, then the upper expanded
(adaxial) es either short, often with apices, or long
tepal larger and often hooded and the lower three and Capsules barrel-shaped oblong,
filiform. to firm-
tepals smallest, with contrasting marking and membranous,
ex- showing the outline of the seeds;
±
tended forward horizontally; perianth tube short seeds (2-)4-l 5 per
locule, globose, relatively large,
funnel-shaped (cylind smooth, Chromosome number x
usually shiny.
panded
above), sometimes
obliquely so, or elongate, 10.
much
A
exceeding the bracts and dimorphic, the low- genus of 3 South Africa and
1 species, all in
er part cylindric and abruptly bent and widened
restricted to the southwestern Cape, southern Na-
±
above; tepals with the whorls equal or the upper maqualand, and the western Karoo.
Key
to the
Species
\
la. Perianth actinomorphic; stamens
either symmetrically disposed around a central style or the abaxial stamen
lying opposite the adaxial and
tepal, style opposite the adaxial tepal (section Sparaxis).
2a. Bracts entire to slightly lacerate; stamens symmetrically when
disposed around erect style (this includes
anthers are
coiled).
3a. Anthers and
spirally coiled reaching only apex elegans
to of the style 3. S.
3b. Anthers
straight or and
slightly twisted cm-ved, extending
well past style branches.
4a. Anthers 2-5
straight, yellow to ochre; spikes -flowered; tepals orange, marked with yellow and
black
tricolor
5.
1^
4b. Anthers incurved and
slightly twisted, red to brown; spikes (2-)5-10-flowered; tepals rose
marked
pink, with yellow and red to purple 5. pillansii
2.
2b. Bracts deeply
lacerate with cusps
at least as long as the bract7stamensTy^^ or
rest of "the
asymmetrical
but not
coiled.
5a. Bract edges
irregularly serrate (fringed); stamens symmetrically arranged around
style.
Oa, Tepals 27-35 x 8- mm; mm
leaves oblong 9-
1 to lanceolate, 2 wide and obliquely apiculate
1
maculosa
5,
5_
6b. Tepals 18-25 x 5-8 mm; mm
— leaves linear to narrowly 2-5(-9) wide and acute
lanceolate,
fragrans
5.
_ _ 4,
±
5b. Bract edges
entire, not
serrate or fringed; stamens
asymmetrically
"disposed,
Stem
7a. usually branched
in mid-axis and bearing many produced
a cauline leaf; small cormlets
flowermg
after at nodes
all hulhifera
7, 5.
7b. Branches
present produced from
if base, and axes lacking a cauline cormiets not produced
leaf;
numbers
and
'" never above ground grandipra
„ 6 S
lb. Perianth zygomorphic,
with
the
adaxial (upper)
tepal largest, erect, and often h"ooded, and lower three tepals
smauest and ±
horizontal; stamens
and ±
unilateral arcuate, with the anthers and contiguous
parallel
^
(section Synnotia),
Narrow mm
8a,
part of perianth tube more than 25
long, well exserted from the bracts; tube bent abruptly
(geniculate) apex
at of
the slender
part 5 variegata
13
„
Narrow
8b.
part of perianth
tube than 25 nun
less long, exserted"or included b7^"tube gently to
in the
sharply curved
at the apex of the narrow
part.
Corm
9a.
tunics of mm
fine fibers; flowers small, the tepals than 10 parviflora
less long S.
8.
Vb. Lorra tumcs
either of fine or coarse
fibers thickened into claws below; flowers usually larger, the
mm
upper
tepals
at least 15 long, but shorter the corm
if tunics of coarse
fibers,
Lorm
lUa. tumcs
hard
of
reticulate fibers not extending upwards
in a neck; upper tepal directed
forward and somewhat
hooded
over
the stamens.
Ua.
Upper
tepal ca. 16 nmi
long, bases of the leaves uniformly dividing
colored; style
opposite the lower half of the anthers, mm viUosa
the
style branches ca. 2 long.^ 11. S.
mm
lib. Upper 22-24
tepal
lone: ha«»« nf ths. Io.,.«o i;„i,*i,. i.i.j -.t.^ 1^ ir.nt
.
