Bot. Macaronésica 24: 87-106 (2003)
87
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN
ANDROCYMBIUM (COLCHICACEAE)
NÚRIA MEMBRIVES1, JOAN PEDROLA -MONFORT1 & JULI CAUJAPÉ-CASTELLS2
1
Estació Internacional de Biologia Mediterrània-Jardí Botànic Marimurtra. Passeig Karl Faust, 10.
17300 - Blanes. Girona. Apdo. Correos 112. España (nuriamem@jazzfree.com; jpedrola@grn.es).
Jardín Botánico Canario «Viera y Clavijo». Apdo. 14 de Tarifa Alta. 35017 Las Palmas de Gran Canaria, islas Canarias. España (julicaujape@granca.step.es).
2
Recibido: Agosto 2000.
Palabras clave: Androcymbium, Colchicaceae, morfología seminal, filogenia, África.
Key words: Androcymbium, Colchicaceae, seed morphology, phylogeny, Africa.
SUMMARY
Macromorphological and micromorphological seed characteristics are described in 32 populations
belonging to 19 taxa of the genus Androcymbium distributed in Southwest Africa. The heterogeneity
shown in the seed characteristics in Southwest African species is compared with the uniformity described previously in the Northern African congeners. The principal seed characteristics are evaluated in
the taxonomic classification of the genus, and their evolution is considered under a recent cladistic
analysis from morphological data. According to this phylogeny, all seed characters studied show complex patterns of parallelisms and reversals in the evolution of the genus.
RESUMEN
Se describen las características macromorfológicas y micromorfológicas de las semillas de 32 poblaciones pertenecientes a 19 taxones del género Androcymbium distribuidas en Sudáfrica Occidental.
La heterogeneidad observada en las características seminales en las especies sudafricanas se compara con la uniformidad descrita previamente en los congéneres norteafricanos. Las principales características seminales se evalúan en la clasificación taxonómica del género y la evolución de estos caracteres se analiza en un reciente análisis cladista a partir de datos morfológicos. Según esta filogenia,
todos los caracteres seminales estudiados presentan complejos patrones de paralelismos y reversiones
en la evolución del género.
ISSN 0211-7150
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NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
INTRODUCTION
The genus Androcymbium Willd. (Colchicaceae) is represented by about 50
species (ARNOLD & WET, 1993; MÜLLER-DOBLIES & MÜLLER-DOBLIES, 1998;
PEDROLA-MONFORT et al., 1999a, 1999b, 2000) with a disjunct distribution in Africa
(Fig. 1A). The genus has been historically divided into three sections: Androcymbium, Erythrostictus, and Dregeocymbium (KRAUSE, 1920). Most of the species
belong to section Androcymbium and are distributed in South Africa, either in the
western (about 40 species) and in the eastern (seven species) region. Only two
species are ascribed to section Dregeocymbium (A. dregei and A. exiguum). These
distribute in Southwest Africa and Namibia. Seven species belong to section Erythrostictus and occur mainly in Northern Africa (four in the Mediterranean basin
and two in the Canary Islands), and only one of them (A. roseum), occurs in
Southern Africa. MÜLLER-DOBLIES & MÜLLER-DOBLIES (1998) published a new supraspecific classification of section Androcymbium where they propose its segregation into two subsections with eight series and four subseries. Recently,
MEMBRIVES (2000) proposes a new generic classification under phylogenetic tenets
based on the cladistic analysis of morphological and reproductive traits.
Given that seed morphology has been shown to be a conservative character in
many plant species (DAVIS & HEYWOOD, 1963), it is generally inferred that it bears
taxonomic value (STEBBINS, 1974; STUESSY, 1990). Within the Colchicaceae, variability in seed size, colour, caruncula presence and in the microstructure of the external layer has been reported in some genera (BUXBAUM, 1936; NORDERSTAM,
1982; PEDROLA-MONFORT, 1993). In Ornithoglossum, the microstructure of the external layer is of proven taxonomic usefulness (NORDERSTAM, 1982). Previous surveys in the genus Androcymbium allowed species differentiation based on seed
size alone (GREUTER, 1967; ROESSLER, 1974; SANTOS, 1977; REIFERBERGER, 1990;
PEDROLA-MONFORT, 1993). Within section Erythrostictus, the shape of the cells of
the internal layer showed inter-specific variability, with the Northern African A. wyssianum and the Southern African A. roseum displaying rectangular cells, and the
rest of species showing quadrangular cells (PEDROLA-MONFORT, 1993).
Our objectives are, first, to describe seed morphology and biometry in a broad
representation of Androcymbium species distributed in Southwest Africa (Fig. 1B)
in order to assess their value for taxonomic classification within the genus. And
second, to assess the evolution of these traits in the frame of a recent cladistic
analysis based on morphological data.
MATERIALS AND METHODS
We sampled 32 populations representing 19 taxa of genus Androcymbium distributed in Southwest Africa (Appendix 1; Fig. 1B), and the six species from
Northern Africa already studied by PEDROLA-MONFORT (1993). Seeds were obtained from plants grown in culture except from those belonging to A. cruciatum
and A. volutare which correspond to herbarium specimens (U. Müll-Doblies & D.
Müll.-Doblies 80099n -Botanischer Garten und Botanisches Museum Berlin-
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
89
Dahlem- for A. cruciatum, and P. Goldblatt 6238 -Royal Botanic Garden, Kew- for
A. volutare).
Figure 1.- A. Geographic distribution of genus Androcymbium. The numbers show the distribution of
Northern African species (1: A. hierrense; 2: A. psammophilum; 3: A. gramineum; 4: A. wyssianum; 5:
A. rechingerii; 6: A. palaestinum). B. Geographic distribution of the Southwest African populations
studied. The abbreviations of the Southwest African populations are described in Appendix 1.
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NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
Macromorphological traits.- Following previous studies within the Colchicaceae
(NORDERSTAM, 1982; PEDROLA-MONFORT, 1993), we studied seed shape, diameter,
colour, and caruncula presence/absence. Morphological observations and biometrical measurements were carried out in a binocular lens. Seed diameter was obtained by pooling 50 measurements of each of five individuals per population.
Micromorphological traits.- The seeds of the species of genus Androcymbium
show two different layers that wrap the endosperm and form the testa (PEDROLAMONFORT, 1993). Accordingly, the micromorphological descriptions of the seeds
considered both the external and the internal layers. To observe the internal layer,
the seeds were boiled during 10-15' and then the external cell layer was removed
using a nail. Then, seeds were subjected to ultrasonic waves to remove cellular
remains of the external layer. The studied traits were the shape of the cells in both
layers. Observations were carried out using the Scanning Electron Microscope
(SEM) at the University of Valencia (Spain).
RESULTS
The irregularities due to the pressure within the capsula notwithstanding, the
seeds of genus Androcymbium are always globose (Figs. 4-8) except in A. cuspidatum (Fig. 7C), where they show a slightly elliptical shape. Seed diameter was
mostly around 1.5 mm (Table 1, Fig. 2) and varied between 0.7 mm in A. dregei
and 3.5 mm in A. burchellii subsp. pulchrum. The seed colour varied among light
brown, brown, brown-reddish and black (Table 1).
Figure 2.- Seed diameter in Androcymbium. The boxes show the median, the quartiles and the
extreme values. In all cases, the names of species are coded with the first four letters.
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
Population
Seed color
Seed diameter
CAR
EXT
91
INT
Section Androcymbium
ALBA-PK
Brown-reddish
(1.20) 1.52 (1.80)
P
Reticulate (II)
Quadrangular
AUST-GH
Brown-reddish
(1.50) 1.92 (2.50)
A
Mosaic (IV)
Rectangular
AUST-WP
Brown-reddish
(1.30) 1.79 (2.40)
A
Mosaic (IV)
Rectangular
BELL-VI
Black
(1.00) 1.07 (1.20)
A
Mosaic (III)
Indifferenced
BURC-HX
Brown
(1.80) 2.07 (2.60)
A
Rough
Quadrangular
CAPE-HO
Brown
(1.35) 1.49 (1.60)
P
Reticulate(II)
Quadrangular
CIRC-SB
Black
(1.70) 1.84 (1.90)
P
Mosaic (III)
Rectangular
CUSP-CA
Brown
(1.50) 1.62 (1.90)
A
Reticulate (I)
Quadrangular
CUSP-MO
Brown
(1.20) 1.45 (1.70)
A
Reticulate (I)
Quadrangular
EGHI-CI
Brown
(1.30) 1.51 (1.70)
A
Mosaic (IV)
Rectangular
EGHI-PK
Brown
(1.20) 1.41 (1.60)
A
Mosaic (IV)
Rectangular
HANT-CA
Brown-reddish
(1.90) 2.14 (2.40)
P
Mosaic (II)
Quadrangular
HENS-EK
Brown-reddish
(1.30) 1.52 (1.65)
P
Mosaic (I)
Quadrangular
HUNT-EK3
Black
(1.30) 1.44 (1.60)
A
Mosaic (I)
Indifferenced
IRRO-EK
Black
(1.20) 1.43 (1.60)
P
Mosaic (III)
Variable
IRRO-EK2
Black
(1.50) 1.74 (1.90)
P
Mosaic (II)
Variable
IRRO-EK6
Black
(1.50) 1.69 (1.90)
P
Mosaic (II)
Variable
IRRO-KA
Black
(1.20) 1.49 (1.80)
P
Mosaic (I)
Variable
IRRO-KW
Black
(1.10) 1.33 (1.60)
P
Mosaic (I)
Variable
IRRO-VP
Brown-reddish
(1.30) 1.49 (1.70)
P
Mosaic (I)
Variable
IRRO-VY
Black
(1.20) 1.39 (1.60)
P
Mosaic (I)
Variable
POEL-CO
Black
(1.00) 1.25 (1.50)
A
Mosaic (I)
Rectangular
POEL-NB
Black
(1.20) 1.42 (1.60)
A
Mosaic (I)
Rectangular
POEL-ST
Black
(1.20) 1.26 (1.40)
A
Mosaic (I)
Rectangular
PULC-CA
Brown
(1.90) 2.30 (2.80)
A
Rough
Quadrangular
PULC-NI
Brown
(2.20) 2.76 (3.50)
A
Rough
Quadrangular
VILL-EK
Black
(1.40) 1.58 (1.80)
P
Mosaic (III)
Rectangular
VILL-ST
Black
(1.60) 1.83 (2.00)
P
Mosaic (III)
Rectangular
?
A
Mosaic (II)
Quadrangular
(1.00) 1.35 (1.70)
A
Mosaic (I)
Quadrangular
Rough
Quadrangular
Reticulate (I)
Quadrangular
VOLU-ST
?
WALT-ST
Black
Section Erythrostictus
CRUC-CA
?
?
A
Section Dregeocymbium
DREG-PK
Light brown
(0.70) 0.94 (1.20)
A
Table 1.- Macromorphological seed characteristics of genus Androcymbium. CAR: caruncula
(P=present; A=absent). Seeds are measured in mm. EXT: Microstructure of the external layer; INT: Cell
morphology of the internal layer. The abbreviations of the populations are described in Appendix 1.
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NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
In species A. albanense subsp. clanwilliamense, A. capense, A. circinatum, A.
hantamense, A. henssenianum, A. irroratum, and A. villosum, the raphe develops
in a caruncula (Table 1; Figs. 4A,E; 5C; 6A,C; 7E,G), a phenomenon that has also
been reported in other genera of the Liliales like Erythronium, Gagea, Uvularia,
Colchicum, Hermodactylus, and Patersonia (DAHLGREN, 1985). The width of this
caruncula-like raphe is never higher than 1 mm.
The external layer is rough in A. austrocapense, A. bellum, A. burchellii subsp.
burchellii, A. burchellii subsp. pulchrum, A. circinatum, A. cruciatum, A. dregei, A.
eghimocymbion, A. gramineum, and A. villosum (Figs. 5G; 6; 7A; 8) and smooth in
the rest of species studied. Cell shape and disposition as analyzed in the SEM
revealed three distinct patterns of microstructure in the external layer (Table 1):
a) Type mosaic. It features cells with walls within the cell surface. It is observed
in most species studied and four subtypes can be distinguished. Mosaic I is characterized by a smooth pavement with regular, polygonal cells with 5-6 faces of
different size that have sharp edges and a thin cell wall (Figures 4; 5B). It was
observed in A. henssenianum, A. huntleyi, A. irroratum (populations IRRO-KA,
IRRO-KW, IRRO-VP and IRRO-VY), A. poeltianum, and A. walteri. Mosaic II is
characterized by a smooth pavement with regular cells, blunt edges and a thick cell
wall (Fig. 5D,F). It was observed in A. hantamense, A. volutare, and A. irroratum
(populations IRRO- EK2 and IRRO-EK6). Mosaic III is characterized by an irregular
rough pavement with differently sized polygonal cells with 5-6 faces arranged in
different levels, sharp edge and a thin cell wall (Figs. 5H; 6B,D). It was observed in
A. bellum, A. circinatum, A. irroratum (population IRRO-EK), and A. villosum.
Mosaic IV is characterized by a smooth pavement with irregular cells with rounded
or elliptical edges and undifferentiated cell walls (Fig. 9F,H). It was observed in A.
austrocapense and A. eghimocymbion.
b) Type reticulate. It features an external layer micromorphology with walls
overtaking the cell surface. Two subtypes were observed among the studied species. Reticulate I is characterized by irregular polygonal cells (Fig. 4B,D) and was
observed in A. cuspidatum and A. dregei. Reticulate II corresponds to a suprareticulate structure with a thin reticle on top of the thick reticle unerneath (Fig.
7F,H) and was observed in A. albanense subsp. clanwilliamense, and A. capense.
c) Type rough. It features irregular pavement without clear cell edges (Fig. 8)
and was described in A. burchellii subsp. burchellii, A. burchellii subsp. pulchrum,
A. cruciatum, and in all the Northern African species of the genus.
The surface of the internal layer showed a rough surface in A. bellum and A.
huntleyi, where the cell edges cannot be precisely delimited (Fig. 9A, table 1). In
contrast, the rest of species showed a smooth or almost smooth wall (Fig. 9B,C,D),
where the edges of the cells in the internal layer are well delimited and group the
species in three general types: 1) quadrangular cells in A. albanense subsp.
clanwilliamense, A. burchellii subsp. burchellii, A. burchellii subsp. pulchrum, A.
capense, A. cruciatum, A. cuspidatum, A. dregei, A. gramineum, A. hantamense,
A. henssenianum, A. rechingerii, A. volutare and A. walteri (Fig. 9B); 2) rectangular
cells in A. eghimocymbion, A. austrocapense, A. circinatum, A. poeltianum, A.
villosum, and A. wyssianum (Figs. 9C); and 3) irregular cells (generally quadrangular tending to rectangular) in A. hierrense, A. irroratum, A. palaestinum, and A.
psammophilum (Fig. 9D).
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
93
DISCUSSION
Seed morphology in the species of Androcymbium shows a remarkable interspecific heterogeneity, while it behaves quite uniformly at the intra-specific level
except for the seven studied populations of A. irroratum. On the one hand, population IRRO-EK shows a rough external layer, whereas the rest show a smooth external layer. On the other hand, seeds of population IRRO-VP are brown-reddish,
and the rest of conspecific populations are black (Table 1).
Seed size in the South African taxa (Table 1) is similar to that of the North African congeners (Table 2) and, of which A. palaestinum and A. rechingerii show the
smallest seeds (averaging 1.6 mm), and A. hierrense shows the biggest seeds,
with an average of 2.3 mm (ARDANUY, 1997). Remarkably, seed size in genus Androcymbium is small as compared with other genera of the Colchicaceae like Colchicum, Merendera, Bulbocodium, Ornithoglossum, and Gloriosa (NORDERSTAM,
1982; GIBERT, unpubl.).
Taxa
A. gramineum
Seed diameter
1.75 ± 0.20
A
CAR
EXT
Rugose
INT
Quadrangular
A. hierrense
2.26 ± 0.29
A
Rugose
Irregular
A. palaestinum
1.60 ± 0.12
A
Rugose
Irregular
A. psammophilum
1.75 ± 0.23
A
Rugose
Irregular
A. rechingerii
1.60 ± 0.15
A
Rugose
Quadrangular
A. wyssianum
1.93 ± 0.26
A
Rugose
Rectangular
Table 2.- Macromorphological seed characteristics of Northern African species of genus
Androcymbium (from PEDROLA-MONFORT, 1993). CAR: caruncula (P=present; A=absent). EXT:
Microstructure of the external layer; INT: Cell morphology of the internal layer. The diameter of the
seeds are measured in mm.
The caruncula might act as a nourishing reward and, therefore, it could be related to seed dispersal mechanisms. All South African species studied show dehiscent capsule. Hence, seed dispersal can be carried out individually, indicating that
dispersal agents could be influencing the distribution of individuals in populational
space. In contrast, some of the species distributed in North Africa feature a dehiscent capsule and, consequently, the dispersal unit is the whole stock of seeds per
individual. A study in a stand of A. gramineum revealed that this is a paramount
factor in determining the spatial distribution of individuals in neighbourhoods of
genetically related individuals (CAUJAPÉ-CASTELLS & PEDROLA-MONFORT, 1997).
A close relationship between A. wyssianum (distributed in Northern Africa) and
A. roseum (distributed in Southern Africa) was inferred by PEDROLA-MONFORT
(1993) from the observation of rectangular cells in the internal layer of the seeds in
both species. This fact led to the consideration that A. wyssianum and A. roseum
might be the same species or were closely related. This also indicated that A.
roseum, distributed in South Africa, could be of paramount importance to under-
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NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
stand the dispersal path of the genus into North Africa. Recent studies allowed us
to observe additional seed material and indicate that the cells in the internal layer
of A. hierrense, A. palaestinum, and A. psammophilum (which were defined as
quadrangular in PEDROLA-MONFORT, 1993) show variable morphologies (Table 2).
A recent phylogeny based on cpDNA RFLPs (CAUJAPÉ-CASTELLS et al., 1999)
groups A. gramineum, A. palaestinum, and A. rechingerii, and the species A. hierrense, A. psammophilum, and A. wyssianum in diferent clades. This topology is
well supported and hints at the possibility that the present distribution of the North
African species might be accounted for by the existence of two different dispersal
lines starting in South Africa (CAUJAPÉ-CASTELLS et al., in mss.). The shape of the
cells in the internal layer might support this hypothesis under the consideration that
the species that exhibit rectangular cells (A. roseum and A. wyssianum, and with
variability A. hierrense, A. psammophilum, and A. palaestinum) could be more
primitive than A. gramineum and A. rechingerii, whose cells tend to be quadrangular. The morphological variability observed in the cells of the internal layer of South
African species makes it difficult to substantiate this hypothesis.
Taxonomic implications
None of the seed characters allows to support the classification of the genus in
three sections (Androcymbium, Dregeocymbium and Erythrostictus). All of the seed
characters examined (size, shape, colour, presence/absence of caruncula, type of
external and internal layer) show variability in the 17 taxa of section Androcymbium
included in this study. Therefore, seed characters cannot be used as an indicator
of sectional taxonomic category in the genus Androcymbium.
The only representative studied of section Dregeocymbium (A. dregei) shows
two seed traits that differentiate it from the rest of species: a small seed size (almost always < 1 mm) and the brown light colour of the external cover. This species
had been included in section Erythrostictus by previous authors (MÜLLER-DOBLIES
& MÜLLER-DOBLIES, 1990). The seed traits exclusive of A. dregei in the genus are
not shared by the rest of species within section Erythrostictus (PEDROLA-MONFORT,
1993; ARDANUY, 1997), which have bigger and darker seeds. From the studied
species, only A. bellum and some individuals of populations IRRO-KW and POELCO, present a seed diameter that overlaps only partially with the range observed in
A. dregei. However, their seeds are black.
Androcymbium cruciatum has been included historically within section Erythrostictus (Appendix 1). This species shares the absence of caruncula with the North
African species (belonging to this section), and differst from them in the external
layer type. MEMBRIVES (2000) argues for the inclusion of A. cruciatum in a new
monotypic section based on morphological traits.
The species A. bellum was segregate from section Erythrostictus and include it
in section Androcymbium by MÜLLER-DOBLIES & MÜLLER-DOBLIES (1998). This species shares with other species in section Erythrostictus the absence of caruncula
and the roughness of the testa’s external layer. However, the seeds of A. bellum
show a smaller size and a rougher internal layer than the others (PEDROLAMONFORT, 1993). According to seed traits, A. bellum is most similar to A. huntleyi
(that belongs to section Androcymbium) than the other species of section Erythros-
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
95
tictus. Both of them share seed colour and diameter, the absence of caruncula,
mosaic external layer and rough internal layer, and they only differ in the size and
shape of the cells in the external layer.
Only 15 of the 34 taxa classified within section Androcymbium (Appendix 2) by
MÜLLER DOBLIES & MÜLLER DOBLIES (1998) were included in our study. The two
subsections of these authors are not supported by any of the studied characters.
Subsection Gradatocymbium includes the species A. albanense, A. poeltianum, A.
henssenianum, and A. cuspidatum, that differ in most of the studied traits (seed
shape, colour, caruncula's presence, type of external layer and shape of the cells
of the internal layer). Neither does subsection Androcymbium share any of the
studied seed traits. For instance, A. irroratum and A. eghimocymbium (series
Eghimocymbia) differ in seed colour, presence/absence of caruncula, type of mosaic and the shape of the internal layer cells (Table 1).
A cladistic analysis of genus Androcymbium based on morphological and reproductive data (MEMBRIVES, 2000) concludes that none of the seed traits analysed
is monophyletic (Fig. 3) and shows that caruncula presence appeared three times
in the evolution of the genus. First in A. henssenianum, then in A. hantamense and
A. capense, and finally in A. irroratum, A. albanense subsp. clanwilliamense, A.
villosum, and A. circinatum. Furthermore, this character would have undergone a
reversal in A. volutare, A. walteri, A. bellum, and A. poeltianum. When the evolution
of the microstructural patterns of the external layer is analysed, the result is that
none of the four types behaves as a monophyletic trait either. Rough forms appear
at the base of the morphological phylogenetic tree as primitive external layer
microstructures, and then again in the terminal clade formed by A. burchellii subsp.
burchellii and A. burchellii subsp. pulchrum. Reticulate types appear three times in
that phylogeny: in the clade made up by A. dregei and A exiguum, in A.
cuspidatum and in A. crispum. Suprareticulate types appear independently in
species from two distinct clades (A. capense and A. albanense subsp.
clanwilliamense). Mosaic types appear three times also: in A. henssenianum and
A. huntleyi, in A. melanthioides and A. hantamense, and in the clade that groups A.
eghimocymbion, A. austrocapense, A. irroratum, A. villosum, A. circinatum, A. volutare, A. walteri, A. bellum, and A. poeltianum.
Given that none of the seed traits analysed in the genus Androcymbium are
good indicators of phylogenetic relatedness (MEMBRIVES, 2000), they have no
usefulness as taxonomic descriptors of supra-specific classification under a phylogenetic perspective. Therefore, we can conclude that the seed characters considered do not indicate either phylogenetic taxonomic assemblages, or morphological groupings like the historical classification in three sections (KRAUSE, 1920),
or the MÜLLER-DOBLIES & MÜLLER-DOBLIES (1998) proposal.
96
NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
Figure 3.- Phylogenetic tree of genus Androcymbium from morphologic and reproductive data
(MEMBRIVES, 2000). The species with seminal data unknown have been excluded. The evolution of
the external layer seed microstructure (squares at the end of the branches) and the
presence/absence of caruncula (where the presence of caruncula is represented by a picture of the
seed with a developed raphe) is represented in the tree. Types of microstructure: R=rough;
Re=reticulate; S=suprareticulate; M=mosaic; ?=unknown data.
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
97
ACKNOWLEDGEMENTS
We thank Isabel Mateu-Andrés for the facilities given to undertake this study
and for her suggestions in the classification of cell patterns. Amparo Ardanuy provided for the welfare of the material in cultivation. The Karl Faust Foundation gave
financial support for both the expedition to South Africa that allowed us to collect
the analysed samples and the subsequent investigations in conservation genetics
of Androcymbium carried out in the “Estació Internacional de Biologia MediterràniaJardí Botànic Marimurtra”.
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stand of Androcymbium gramineum (Cav.) McBride (Colchicaceae) Heredity 79: 341-349.
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Phylogeny of the Genus Androcymbium (Colchicaceae) Syst. Bot. 24(4): 581-597.
DAHLGREN, R., H.T. CLIFFORD & P.F. YEO, 1985.- The families of the Monocotyledons: Structure, Evolution, and Taxonomy. Springer-Verlag. Berlin.
DAVIS, P.H. & V.H. HEYWOOD, 1963.- Principles of Angiosperm Taxonomy. Ed. Olivier. Edinburg.
GREUTER, W., 1967.- Contributiones floristicae Austro-Aegeae. Candollea 22(2): 233-253.
KRAUSE, K., 1920.- Revision der Gattung Androcymbium Willd. Notizbl. Bot. Gart. Berlin 7: 512-526.
MEMBRIVES, N., 2000.- Biologia evolutiva del gènere Androcymbium (Colchicaceae) a Sudàfrica Occidental. Tesis Doctoral. Universitat de Barcelona.
MÜLLER-DOBLIES, U. & D. MÜLLER-DOBLIES, 1990.- Zur Kenntnis der Gattung Androcymbium (Colchicaceae) mi südlichen Afrika: 2. Androcymbium exiguum hat doch Artwert. Willdenowia 19:453-470.
- & D. MÜLLER-DOBLIES, 1998.- De Liliifloris Notulae. 6. De decuria prima specierum novarum generis
Androcymbium sect. Androcymbium (Colchicaceae) in Africa Australi s.l. Praeterea novitates de
huius sectionis nonnullarum specierum distributione (praesertim aucta speciminibus
STEllenbosensibus i.a. collectis a botanico vero E. G. H. OLIVERO) Fedd. Reper. 109 (7-8): 551572.
NORDERSTAM, B., 1982.- A monograph of the genus Ornithoglossum (Liliaceae). Opera Botánica 64: 151.
PEDROLA-MONFORT, J., 1993.- Biologia poblacional del complexe Androcymbium gramineum (Cav.)
McBride (Colchicaceae). Tesis Doctoral. Universitat de València.
- N. MEMBRIVES, J.M. MONTSERRAT & J. CAUJAPÉ-CASTELLS, 1999a.- A new species from the Western of
South Africa: Androcymbium huntleyi (Colchicaceae) Fontqueria 53: 1-2.
- N. MEMBRIVES & J.M. MONTSERRAT, 1999b.- Two new Androcymbia (Colchicaceae) from Western
South Africa. Fontqueria 54(2): 7-9.
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species of the genus Androcymbium Willd. (Colchicaceae) in Western South Africa. Bot.
Macaronésica, 24: 107-126.
REIFERBERGER, U., 1990.- Androcymbium hierrense Santos ssp. macrospermum Reifenberger ssp. nov.
Liliaceae., ein neuer Endemit der insel La Gomera. Vierea 18: 251-259.
ROESSLER, H., 1974.- Die Gattung Androcymbium Willd. (Liliaceae) in Südwestafrika. Mitt. Bot.
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SANTOS, A. 1977.- Androcymbium hierrensis spec. nova para la flora canaria. Ed. Funchal. Pp: 53-59.
STEBBINS, G. L., 1974. Flowering Plants. Evolution above the species level. Harvard University Press.
Cambridge. Massachusset.
STUESSY, T. F., 1990.- Plant Taxonomy. Columbia University Press. New York.
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Appendix 1.- Sampled populations and localities of genus Androcymbium studied in Southwest Africa.
Abb: abbreviations of the population and locality.
Population
Section Androcymbium
A. albanense subsp.
clanwilliamense
A. austrocapense
A. austrocapense
A. bellum
A. burchellii subsp.
burchellii
A. burchellii subsp.
pulchrum
A. burchellii subsp.
pulchrum
A. capense
A. circinatum
A. cuspidatum
Abb.
Locality
ALBA-PK
3219AA (WUPPERTAL) Clanwilliam-Wuppertal Road. Km 10
AUST-GH
AUST-WP
BELL-VI
BURC-HX
3418AC (SIMONSTOWN) Road to the Cape of Good Hope
3418AD (SIMONSTOWN) Wheal's Point. Cape Point Reserve
2817DC (VIOOLSDRIFT) Steinkopf to Vioolsdrift Road. Km 40
3319BC (WORCESTER) Worcester to Towsrivier Road. Near
Hexrivierpass
3119DA (CALVINIA) Calvinia to Ceres Road, 7 km from the
deviation to Kreitzberg
3118AA (CALVINIA) Wild Flowers Reserve of Nieuwoudtville
PULC-CA
PULC-NI
CAPE-HO
CIRC-SB
CUSP-CA
A. cuspidatum
CUSP-MO
A. eghimocymbion
EGHI-CI
A. eghimocymbion
A. hantamense
EGHI-PK
HANT-CA
A. henssenianum
A. huntleyi
HENS-EK
HUNT-EK3
A. irroratum
IRRO-EK
A. irroratum
IRRO-EK2
A. irroratum
IRRO-EK6
A. irroratum
A. irroratum
IRRO-KA
IRRO-KW
A. irroratum
A. irroratum
IRRO-VP
IRRO-VY
POEL-CO
A. poeltianum
POEL-NB
A. poeltianum
POEL-ST
A. poeltianum
VILL-EK
A. villosum
VILL-ST
A. villosum
VOLU-CA
A. volutare
WALT-ST
A. walteri
Section Erythrostictus
CRUC-ST
A. cruciatum
Section Dregeocymbium
DREG-PK
A. dregei
3318AB (CAPE TOWN) Malmesbury to Hopefield Road. Km 49
2917DB (SPRINGBOK) 3 km W of Springbok
3119DA (CALVINIA) Calvinia to Ceres Road, 7 km from the
deviation to Kreitzberg
3320CD (MONTAGU) Near Montagu-Badskloof. W side of the
gorge.
3218DB (CLANWILLIAM) N-7 Road in the Piketberg Pass to
Citrusdal
3219AA (WUPPERTAL) Clanwilliam to Wuppertal Road. Km 28
3119DA (CALVINIA) Calvinia to Ceres Road, 7 km from the
deviation to Kreitzberg
2817CC (VIOOLSDRIFT) Eksteenfontein to Modderfontein Road
2917AD (SPRINGBOK) Springbok to Port Nolloth Road, 20 km
from the first entry to Eksteenfontein
2917AD (SPRINGBOK) Springbok to Port Nolloth Road, 6 km
from the first entry to Eksteenfontein
2917AD (SPRINGBOK) Springbok to Port Nolloth Road, 15 km
from the first entry to Eksteenfontein
2817CC (VIOOLSDRIFT) Eksteenfontein to Modderfontein
Road, first turn
3018CB (KAMIESBERG) Bitterfontein to Kliprand Road.
3118BC (VANRHYNSDORP) Vredental to Koekenaap Road,
100 from the train station
3119AC (CALVINIA) Vanrhynspass
3118AD (VANRHYNSDORP) Vrendendal to Vanrhynsdorp
Road.
2917DB (SPRINGBOK) Springbok to Concordia Road.
2917DB (SPRINGBOK) Springbok to Nababeep Road.
2917DC (SPRINGBOK) Road from Steinkopf to Springbok, 5 km
2817CC (VIOOLSDRIFT) 1 km S of Eksteenfontein
2917BC (SPRINGBOK) 3 km S of Steinkopf, parallel to N7.
3119BC (CALVINIA) Perdekraal Farm
2917DC (SPRINGBOK) Road from Steinkopf to Springbok. Km 5
2917BC (SPRINGBOK) 5 km SSW of Steinkopf, parallel to N7.
3219AA (WUPPERTAL) Clanwilliam to Wuppertal Road. Km 28
NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
100
Appendix 2.- Taxonomic classification for section Androcymbium proposed by MÜLLER-DOBLIES &
MÜLLER-DOBLIES (1998).
Section Androcymbium
Subsection Gradatocymbium U.Müll.-Doblies & D.Müll.-Doblies
Serie Gradatocymbia U.Müll.-Doblies & D.Müll.-Doblies
Subserie Gradatocymbia
A. longipes, A. natalense, A. decipiens
Subserie Swazicymbia U.Müll.-Doblies & D.Müll.-Doblies
A. swazicum, A. burkei
Subserie Leistnerocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. leistneri, A. albanense subsp. albanense, A. poeltianum
Subserie Pachystyla U.Müll.-Doblies & D.Müll.-Doblies
A. henssenianum
Serie Trifoliata U.Müll.-Doblies & D.Müll.-Doblies
A. cuspidatum, A. kunkelianum, A. worsonense, A. hughocymbium
Subsection Androcymbium
Serie Eghimocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. eucomoides, A. undulatum, A. albomarginatum, A. vanjaarsveldii, A. scabromarginatum, A. irroratum, A. eghimocymbion
Serie Myiocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. circinatum, A. villosum, A. volutare
Serie Therocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. burchellii, A. latifolium
Serie Melittocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. capense, A. ciliolatum, A. hantamense, A. austrocapense, A. crispum
Serie Androcymbium
A. melanthioides, A. orienticapense, A. striatum
Serie Schlechterocymbia U.Müll.-Doblies & D.Müll.-Doblies
A. bellum
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
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Figure 4.- Microstructure of seed external layer in Androcymbium. Type mosaic I. A,B. A.
henssenianum. C,D. A. huntleyi. E,F. A. irroratum (IRRO-KW). G,H. A. poeltianum. A,C,E,G. Seed.
The white lines indicate 500 µm. B,D,F,H. Detail of the microstructure of the external layer. The white
lines indicate 100 µm.
102
NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
Figure 5.- Microstructure of seed external layer in Androcymbium. Type mosaic I. A,B. A. walteri.
Type mosaic II. C,D. A. hantamense. E,F. A. volutare. Type mosaic III. G,H. A. bellum. A,C,E,G.
Seed. The white lines indicate 500 µm. B,D,F,H. Detail of the microstructure of the external layer. The
white lines indicate 100 µm.
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
103
Figure 6.- Microstructure of seed external layer in Androcymbium. Type mosaic III. A,B. A. circinatum.
C,D. A. villosum. Type mosaic VI. E,F. A. austrocapense. G,H. A. eghimocymbion. A,C,E,G. Seed. The
white lines indicate 500 µm. B,D,F,H. Detail of the microstructure of the external layer. The white lines
indicate 100 µm.
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NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
Figure 7.- Microstructure of seed external layer in Androcymbium. Type reticulate I. A,B. A. dregei.
C,D. A. cuspidatum. Type reticulate suprareticulate. E,F. A. albanense subsp. clanwilliamense. G,H.
A. capense. A,C,E,G. Seed. The white lines indicate 500 µm. B,D,F,H. Detail of the microstructure of
the external layer. The white lines indicate 100 µm.
MORPHOLOGICAL SEED STUDIES OF SOUTHWEST AFRICAN ANDROCYMBIUM
105
Figure 8.- Microstructure of seed external layer in Androcymbium. Type rough. A,B. A. burchellii subsp.
burchellii. C,D. A. cruciatum. E,F. A. gramineum. G,H. A. burchellii subsp. pulchrum. A,C,E,G. Seed.
The white lines indicate 500 µm. B,D,F,H. Detail of the microstructure of the external layer. The white
lines indicate 100 µm.
106
NÚRIA MEMBRIVES, JOAN PEDROLA -MONFORT & JULI CAUJAPÉ-CASTELLS
Figure 9.- Shape of the internal layer cells. A. Rough (A. bellum). B. Quadrangular polygonal (A.
henssenianum). C. Rectangular polygonal (A. austrocapense). D. Irregular polygonal (A. irroratum).
The white lines indicate 60 µm.
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