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Taxonomic revision of Ornithogalum subg. Ornithogalum (Hyacinthaceae) in the Iberian Peninsula and the Balearic Islands

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As a part of a taxonomic revision of the Iberian and Balearic taxa of Ornithogalum, the results concerning O. subg. Ornithogalum are reported. Quantitative and qualitative characters were studied in detail, and they were evaluated for the taxonomy of the three accepted taxa: O. baeticum Boiss., O. bourgaeanum Jord. & Fourr. and O. divergens Boreau. A complete description is presented of all accepted species, and data on their nomenclature, biology, ecology, and distribution are also included. Relationships to other European taxa of the subgenus are also discussed. Moreover, a neotype is designated. Finally, a key is provided to facilitate identification.
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ISSN 0378-2697, Volume 289, Combined 3-4
ORIGINAL ARTICLE
Taxonomic revision of Ornithogalum subg. Ornithogalum
(Hyacinthaceae) in the Iberian Peninsula and the Balearic Islands
Mario Martı
´
nez-Azorı
´
n
Manuel B. Crespo
Ana Juan
Received: 2 July 2009 / Accepted: 13 August 2010 / Published online: 23 September 2010
Ó Springer-Verlag 2010
Abstract As a part of a taxonomic revision of the Iberian
and Balearic taxa of Ornithogalum, the results concerning
O. subg. Ornithogalum are reported. Quantitative and
qualitative characters were studied in detail, and they were
evaluated for the taxonomy of the three accepted taxa:
O. baeticum Boiss., O. bourgaeanum Jord. & Fourr. and
O. divergens Boreau. A complete description is presented
of all accepted species, and data on their nomenclature,
biology, ecology, and distribution are also included.
Relationships to other European taxa of the subgenus are
also discussed. Moreover, a neotype is designated. Finally,
a key is provided to facilitate identification.
Keywords Ornithogalum bourgaeanum
Ornithogalum baeticum Ornithogalum divergens
Taxonomy Distribution Ecology Mediterranean flora
Iberian Peninsula
Resumen En el marco de una revisio
´
n de las especies
ibe
´
ricas y baleares de Ornithogalum, se presentan los
resultados correspondientes a los ta
´
xones de O. subg. Orni-
thogalum. Se estudian con detalle y se evalu
´
a cualitativa-
mente y cuantitativamente el valor taxono
´
mico de los
caracteres morfolo
´
gicos de los tres ta
´
xones considerados:
O. bourgaeanum Jord. & Fourr., O. baeticum Boiss. y
O. divergens Boreau. Para cada especie se presenta una des-
cripcio
´
n completa, y se ofrecen datos sobre su nomencla-
tura, biologı
´
a, ecologı
´
a y distribucio
´
n. Se discuten asimismo
sus relaciones con otras especies europeas del subge
´
nero.
Adema
´
s, se designa un neotipo. Finalmente se aporta una
clave para facilitar la identificacio
´
n de los ta
´
xones
aceptados.
Introduction
Ornithogalum L. is widely distributed in Europe, Asia
(reaching Afghanistan to the East), Africa (except the
Tropic of Cancer) and Madagascar (Zahariadi 1965). The
number of species included in Ornithogalum
, and hence
its circumscription, has been a matter of controversy.
In a traditional sense, it includes about 120–130 taxa
(Landstro
¨
m 1989), or even 200–250 according to Obermeyer
(1978) and Mu
¨
ller-Doblies and Mu
¨
ller-Doblies (1996). On
the basis of floral and reproductive characters, many sections
(Engler 1888; Zahariadi 1965, 1977) and subgenera (cf.
Baker 1872; Zahariadi 1965, 1977, 1980; Obermeyer 1978)
have been accepted within Ornithogalum, many of them
based on previously described genera (cf. Gray 1821;
Rafinesque 1837; Parlatore 1854; Salisbury 1866).
In recent times, very different arrangements, almost
opposite to one another, have been proposed. On the one
hand, Speta (1998) circumscribed Ornithogalum to the
Eurasian and North African taxa of O. subg. Ornithogalum
(=O. subg. Heliocharmos Baker), which was hence con-
formed by 50–60 species, and he regarded the rest of the
groups as autonomous genera such as Albuca L., Cathissa
Salisb., Coilonox Raf., Dipcadi Medik., Eliokarmos Raf.,
Honorius S.F. Gray, Loncomelos Raf., Melomphis Raf.,
Neopatersonia Scho
¨
nland, Pseudogaltonia Kuntze, Stella-
rioides Medik. and Zahariadia Speta. On the other, Manning
et al. (2004) reduced all these genera to Ornithogalum,
which resulted in over 300 species and was thus coincident
with subfamily Ornithogaloideae
. Finally, Manning et al.
(2009) reconsidered the taxonomic identity of taxa within
M. Martı
´
nez-Azorı
´
n(&) M. B. Crespo A. Juan
CIBIO (Instituto de la Biodiversidad), Universidad de Alicante,
PO Box 99, 03080 Alicante, Spain
e-mail: mmartinez@ua.es
123
Plant Syst Evol (2010) 289:181–211
DOI 10.1007/s00606-010-0343-9
Author's personal copy
this subfamily and recognized only four genera: Ornitho-
galum, Albuca, Dipcadi and Pseudogaltonia.
However, phylogenetic studies we are undertaking in the
Ornithogaloideae, based on sequencing of plastid (trnL-F,
rbcL and matK) and nuclear (ITS) regions combined with
morphological data (Martı
´
nez-Azorı
´
n et al. 2010b) indicate
the monophyly of O. subg. Ornithogalum, as well as of
other related subgenera. Due to this fact, a narrower cir-
cumscription of Ornithogalum can also be assumed in
which up to 19 genera are accepted, being well charac-
terized morphologically, phylogenetically and biogeo-
graphically. In the present study O. subg. Ornithogalum is
circumscribed in the sense of O. subg. Heliocharmos
(Baker 1872), and corresponds fully to Ornithogalum sensu
stricto (cf. Speta 1998; Martı
´
nez-Azorı
´
n et al. 2010b).
Ornithogalum subg. Ornithogalum includes plants with:
inflorescences wide and short corymbose or pseudocorym-
bose; floral and fruiting pedicels erect-patent to reflexed;
tepals white on the adaxial face bearing a central green band
on the abaxial face; filaments linear or tapering contracted
abruptly at their apexes; ovary oblong, ovoid or obovoid,
truncate at the apex with six evident ribs in section; long and
filiform style and small trigonous stigma; and seeds globose
with reticulate testa (Moret et al. 1990). About 50 species
are usually included in this group (cf. Speta 1998; Martı
´
nez-
Azorı
´
n et al. 2010b), which are distributed mainly in the
Mediterranean basin, but reaching the Middle East.
Many taxa have been described in O. subg. Ornithogalum
on the basis of subtle morphological variations, usually with
little biological significance (Moret and Galland 1992).
Following Landstro
¨
m(1989), more than 150 taxa were
described in O. subg. Ornithogalum from the Mediterranean
and Irano-Turanian regions, but only 20 or 30 species could
be accepted, and he emphasized the need for a global revi-
sion in the whole subgenus. This complex taxonomy can be
due to intraspecific variation, the plasticity of individuals,
their habit strongly dependent on the environment and poor
conservation of the types (Moret and Galland 1992). Fur-
thermore, the original concepts of taxa were usually modi-
fied by later authors, who made very different links among
taxa on the basis of very different criteria. All these factors
led to a wide diversity of taxa conceptions, and generated an
extremely confusing taxonomy (Peruzzi et al. 2007). The
description by Greuter (1988)ofOrnithogalum as ‘being an
execrable genus’ can be perfectly applied to the current
understanding of most taxonomists about this genus.
Brief history of O. subg. Ornithogalum in the Iberian
Peninsula and the Balearic Islands
The taxonomy of the species included in O. subg. Orni-
thogalum in the Iberian Peninsula and the Balearic Islands
is very diverse and confusing. During the last 250 years, up
to 11 species (O. algeriense Jord. & Fourr., O. baeticum
Boiss., O. bourgaeanum Jord. & Fourr., O. collinum Guss.,
O. divergens Boreau, O. exscapum Ten., O. kochii Parl., O.
monticolum Jord. & Fourr., O. orthophyllum Ten., O. ten-
uifolium Guss. and O. umbellatum L.) have been cited for
the Iberian and Balearic floras (Boissier 1838; Lange 1860;
Willkomm 1862; Jordan and Fourreau 1866; del Amo y
Mora 1871; Ruiz-Rejo
´
n 1978; Colmeiro 1889; Willkomm
1893; Zahariadi 1980; van Raamsdonk 1984, 1985a, b,
1986; van Raamsdonk and Heringa 1987; Moret et al.
1987; Pastor 1987; Moret and Galland 1992; Franco and
Rocha Afonso 1994; Devesa 1995; Aizpuru et al. 2000
;
Bolo
`
s and Vigo 2001), most of them being described from
France, Italy or Algeria (cf. Linnaeus 1753; Tenore 1811,
1829, 1830; Gussone 1825, 1827; Boreau 1847; Parlatore
1857; Jordan and Fourreau 1866), but their relationships
with the Iberian and Balearic plants have never been
shown. As an exception, Boissier (1838) described
O. baeticum from the summits of S
a
Bermeja, near
Estepona (Ma
´
laga, southern Spain), a taxon that he related
to O. umbellatum and O. exscapum Ten. Both latter species
were later cited by Lange (1860). First, he mentioned
O. umbellatum from central and southern Spain, whereas
O. exscapum was restricted to Sacromonte in Granada, a
locality that was recorded by several authors (Willkomm
1862; del Amo y Mora 1871; Colmeiro 1889). Lange
(1860) also noted that some young exemplars collected
near Co
´
rdoba were akin to O. comosum L., because of their
very long, acuminate floral bracts.
Willkomm (1862) reported O. umbellatum, a plant
whose type locality is Blois (France) (cf. Stearn 1983), as
widespread in most of the eastern half of Spain. He
accepted two varieties differing in capsule shape and length
of the floral bracts: (1) var. genuinum Willk. from Castilla,
Arago
´
n and Catalun
˜
a; and (2) var. longebracteatum Willk.
(incl. O. baeticum) from Granada, Ma
´
laga and Ca
´
diz, in
which he segregated a form named nevadense Willk. (as
‘‘ Nevadensis’). Both taxa were revisited by Pau (1922),
who accepted O. umbellatum with two additional varieties:
(1) var. baeticum (Boiss.) Pau from S
a
Bermeja, which he
considered synonymous with O. umbellatum var. longe-
bracteatum Willk., and (2) var. nevadense (Willk.) Pau
from Cerro Lucero in Co
´
mpeta (Ma
´
laga). Moreover,
Willkomm (1862) also suggested the strong possibility of
both O. divergens and O. tenuifolium being found in
Catalun
˜
a and Valencia. In fact, the former was reported by
Pau (1888) from Segorbe (Castello
´
n), and later Willkomm
himself (1893) and Colmeiro (1889) added some localities
for both taxa from several sites in northeastern Spain.
The last taxon of this subgenus to be described from the
Iberian Peninsula was O. bourgaeanum (Jordan and Four-
reau 1866), with its type locality near Plasencia, Ca
´
ceres
182 M. Martı
´
nez-Azorı
´
n et al.
123
Author's personal copy
(western Spain). It has been almost overlooked until today,
being implicitly included in the traditional concept of
O. orthophyllum subsp. baeticum (Boiss.) Zahar. (cf.
Devesa 1995).
In recent times, Zahariadi (1980) accepted five taxa for
the Iberian Peninsula (O. collinum, O. exscapum, O. um-
bellatum, O. orthophyllum subsp. baeticum and O. diver-
gens), to which Moret and Galland (1992) later added
O. algeriense. However, according to most recent authors
(Tutin et al. 1980; Devesa 1995; Valde
´
s et al. 1987;
Aizpuru et al. 2000; Bolo
`
s and Vigo 2001), traditionally at
least two taxa of O. subg. Ornithogalum have been
accepted in the Iberian Peninsula and the Balearic Islands:
O. umbellatum and O. orthophyllum subsp. baeticum. They
were differentiated mostly by the disposition of the capsule
ribs and number of bulbils (equidistant and numerous in the
former, paired and absent in the latter).
As a result of the scenario discussed above, identifi-
cation of Iberian and Balearic individuals of Ornitho-
galum subg. Ornithogalum is extremely difficult. This is
due to different factors: (1) their wide morphological
plasticity produces the overlapping of most of the mor-
phological characters used in the available identification
keys (e.g. the relative length of the bracts with regard to
their pedicels); and (2) species mostly accepted for the
Iberian and Balearic territories (O. orthophyllum and
O. umbellatum) were described from Italy and France,
respectively, and their identity has never been confirmed
by detailed comparison with plants from the type locali-
ties. On this matter, Gadella (1972), Gadella and van
Raamsdonk (1981), van Raamsdonk (1986), Moret
(1992), and Moret and Galland (1992) found evidence
that O. umbellatum corresponds to the triploid plants with
medium sized secondary bulbils which bear leaves in the
first year, and have not reached the Iberian Peninsula.
Similarly, Garbari et al. (2003, 2008), after comparing
populations from the classic locality of O. orthophyllum
sensu stricto in central Italy, concluded that this taxon is a
diploid species endemic to the Abruzzi mountains. On the
other hand, the existence of Iberian populations of
O. monticolum and O. algeriense has been demonstrated
on a solid cytological and morphological basis (van
Raamsdonk 1986; Moret and Galland 1992; Bolo
`
s and
Vigo 2001; Bolo
`
s et al. 2005).
Data are presented here for a revision of O. subg.
Ornithogalum in the Iberian Peninsula and the Balearic
Islands, as a part of a global study on the genus that is
being carried out (cf. Martı
´
nez-Azorı
´
n et al. 2007a, 2009,
2010a, b). The morphological data shown here are conso-
nant with a genetic study of the group using the AFLP
technique (Martı
´
nez-Azorı
´
n et al. 2007b; Mar
´
nez-Azorı
´
n
et al., unpublished data), which strongly supports the tax-
onomy adopted here.
Materials and methods
In this study, a review of a total of 901 vouchers conserved
at the herbaria ABH, BIO, BC, BCN, COI, GDA-GDCA,
HUAL, K, LISI, MA, MACB, MGC, MUB, OVI, P,
SALA, SANT, SEV and VAL (acronyms following Thiers
2010) was undertaken.
As some of the herbarium materials were not complete,
fragments on vouchers were commonly found to be prob-
lematic, since some important structures were lacking (e.g.
bulbs, fruits, or seeds, among others), or measuring certain
structures (e.g. dimensions of the filaments of the stamens
or ovary) would involve serious damage to the material.
Therefore, morphological studies have usually been
undertaken on living material from natural populations, and
within a few hours of collection. A total of 339 fresh plants
from several populations were studied in order to achieve
the maximum morphological variation of each taxon. For
O. bourgaeanum, 235 specimens from 28 populations were
studied, for O. baeticum 46 specimens from 8 populations,
and for O. divergens 24 plants from 5 populations
(Table 1). Additionally, 13 individuals from three French
populations of O. monticolum and 21 individuals from
three Moroccan sites of O. algeriense were also included.
Morphological studies were based on comparison of
both qualitative and quantitative features of bulbs, leaves,
flowers, fruits and seeds, as described in detail by Martı
´
-
nez-Azorı
´
n et al. (2007a, 2010b), as well as on other fea-
tures related to both scanning electron microscopy
techniques and fresh plant scanning. Moreover, the fol-
lowing new characters were also considered:
Bulbs
Number and type of bulbils: The number of bulbils was
counted in each individual when they were present. Fur-
thermore, the type of bulbils was considered, differentiat-
ing between secondary bulbils, usually small and
pediculate, and sessile and larger bulbils generated by
division of the mother bulb into portions of similar size
which shared the basal plate.
Leaves
White band width: The maximum width of the central
white band of the leaf was measured, and values are
expressed in millimetres.
Fruits
Morphology of the capsule section: The capsules were cut
transversally and their morphology and rib structures were
recorded.
Taxonomic revision of Ornithogalum subg. 183
123
Author's personal copy
Table 1 Fresh studied specimens of taxa of O. subg. Ornithogalum used in the morphological and statistical analyses
Taxon Locality UTM coordinates Number of specimens Voucher
O. algeriense Medio Atlas, Azrou, Morocco 30STB9797 10 ABH 51612
O. algeriense Medio Atlas, Aguelmame Azigza, Morocco 30STB7051 5 ABH 51613
O. algeriense Alto Atlas, collado Tizi-n-tichka, Morocco 29RPQ5266 6 ABH 51615
O. baeticum Ca
´
diz, P.N. Grazalema, Spain 30STF9161 12 ABH 47136
O. baeticum Ca
´
diz, P.N. Los Alcornocales, Spain 30STF6345 7 ABH 50136
O. baeticum Granada, Llano de la Perdiz, Spain 30SVG5014 8 ABH 51029
O. baeticum Ma
´
laga, S
a
de las Nieves, Spain 30SUF2273 9 ABH 50148
O. baeticum Ma
´
laga, Estepona, S
a
Bermeja, Spain 30SUF0239 3 ABH 50150
O. baeticum Navarra, Viana, Cicujano, Spain 30TWN5702 3 ABH 51044
O. baeticum Sevilla, Alcala
´
de Guadaira, Spain 30STG5235 1 ABH 50984
O. baeticum Zaragoza, Calcena, Spain 30TXM0411 3 ABH 51045
O. bourgaeanum Albacete, S
a
de Alcaraz, Spain 30SWH5168 4 ABH 50125
O. bourgaeanum Albacete, Calar del Mundo, Spain 30SWH4551 15 ABH 47134
O. bourgaeanum Albacete, El Jardı
´
n, Spain 30SWH5589 6 ABH 51041
O. bourgaeanum Alicante, S
a
Aitana, Spain 30SYH3982 15 ABH 47142
O. bourgaeanum Alicante, Planes, Spain 30SYH3298 17 ABH 47129
O. bourgaeanum Alicante, Coll de Rates, Spain 30SYH5590 5 ABH 50138
O. bourgaeanum A
´
vila, El Barco de A
´
vila, Spain 30TTK8570 10 ABH 51054
O. bourgaeanum Badajoz, P.N. Cornalvo, Spain 29SQD4422 9 ABH 51030
O. bourgaeanum Burgos, Sargentes de la Lora, Spain 30TVN2836 7 ABH 51052
O. bourgaeanum Ca
´
ceres, Alı
´
a, Spain 30SUJ1270 8 ABH 51038
O. bourgaeanum Ca
´
ceres, Las Villuercas, Spain 30STJ9868 25 ABH 51032
O. bourgaeanum Ca
´
ceres, Plasencia, Spain 29TQE5239 6 ABH 51301
O. bourgaeanum Ca
´
ceres, Oliva de Plasencia, Spain 29TQE4750 4 ABH 51302
O. bourgaeanum Ciudad Real, Navas de Estena, Spain 30SVJ66 10 ABH 50072
O. bourgaeanum Cuenca, Can
˜
ada del Hoyo, Spain 30SWK9423 10 ABH 51047
O. bourgaeanum Huelva, Fuenteheridos, Spain 29SQB0498 12 ABH 50129
O. bourgaeanum Jae
´
n, Calar del Cobo, Spain 30SWH3931 6 ABH 51042
O. bourgaeanum Jae
´
n, Calar del Espino, Spain 30SWH3938 6 ABH 51043
O. bourgaeanum Jae
´
n, Pto. de Despen
˜
aperros, Spain 30SVH5649 6 ABH 51040
O. bourgaeanum Jae
´
n, Aldeaquemada, Spain 30SVH6748 5 ABH 51039
O. bourgaeanum Leo
´
n, Collada de Valdeteja, Spain 30TTN9857 5 ABH 51050
O. bourgaeanum Leo
´
n, Valporquera, Spain 30TTN9054 7 ABH 51051
O. bourgaeanum Le
´
rida, A
´
ger, Montsec d’Ares, Spain 31TCG1454 5
O. bourgaeanum Madrid, Pto. de Somosierra, Spain 30TVL5153 7 ABH 51055
O. bourgaeanum Orense, Rubia
´
, Vilardesilva, Spain 29TPH7803 3 ABH 50075
O. bourgaeanum Salamanca, El Cerro, Spain 30TTK4965 10 ABH 51303
O. bourgaeanum Segovia, Carabias, Spain 30TVL4389 7 ABH 51056
O. bourgaeanum Zamora, Cuzcurrita, Spain 29TQF2792 5 ABH 47143
O. divergens Huesca, Jaca, Spain 30TYN0016 5 ABH 51059
O. divergens Jard. Bot. Brest (origin unknown), France 6 ABH 51607
O. divergens Loire Valley, Angers, France 30TXT9360 3 ABH 52834
O. divergens Loire Valley, Nazelles-Negron, France 31TCN4653 5 ABH 52829
O. divergens Tuscany, Siena, Porta Tufi, Italy 32TPN89 5 ABH 51608
O. monticolum Hautes Alpes, col du Lautaret, France 32TKQ9589 3 ABH 52835
O. monticolum Hautes Alpes, Le Mone
ˆ
tier-les-Bains, France 32TLQ0282 5 ABH 52836
O. monticolum Hautes Alpes, Saint-Ve
´
ran, France 32TLQ3151 5 ABH 52837
184 M. Martı
´
nez-Azorı
´
n et al.
123
Author's personal copy
Among all of the characters, only 39 quantitative char-
acters (Table 2) were used for the statistical tests. An initial
principal component analysis (PCA) was conducted
including all measured quantitative characters to explore
morphological variation within the five considered taxa of
O. subg. Ornithogalum: O. algeriense, O. baeticum, O.
bourgaeanum, O. divergens and O. monticolum. Later, a
second PCA was done according to the highest values of
the previous variance obtained from the first and second
principal components, which therefore would assess the
main statistical support among the taxonomic identities. A
discriminant analysis was also performed to check the fit-
ness of the latter PCA results. To identify those features
that had the highest potential as diagnostic characters, we
submitted the dataset to a stepwise discriminant analysis.
The value of the inclusion criterion was set at F = 3.84
and the exclusion criterion at F = 2.71 (default values in
SPSS). A classificatory discriminant analysis was per-
formed with the characters that were included in the step-
wise discriminant analysis. The morphological distinction
among taxa was determined by studying the frequency
distribution of the discriminant scores, and the classifica-
tion was evaluated by cross-validation. All analyses were
conducted with the software SSPS v. 14.
Plant names and authorities are in accordance with the
International Plant Names Index (IPNI 2010). Dots on the
distribution maps correspond to both herbarium material
and studied living populations. For each taxon, only a
selection of localities are listed to avoid repetition and to
shorten information.
Results
Statistical analyses
In the initial PCA of all 39 studied quantitative morpho-
logical characters, nine principal components were
obtained, which explained together 75.6% of the variance,
the two first components being responsible of 42.7% of the
variance (31.8% and 10.9%, respectively). All five taxa
considered in the subgenus appeared rather mixed (Fig. 1).
However, a clear gradient was observed along the first axis
after delimiting three groups: O. bourgaeanum (plus
O. monticolum), O. baeticum (plus O. algeriense) and
O. divergens. All three biological entities showed a wide
overlap of many morphological characters, some of which
were rather discriminative concerning secondary bulbils,
inflorescence and flower features (Table 2). The second
principal component was not effective in distinguishing
taxa. In fact, the values obtained and percentages of vari-
ance explained were very low in all cases.
When only characters with the highest values were
selected (Table 2), a clear differentiation of three groups
was again shown in the subsequent PCA (not shown) and
discriminant analysis (Fig. 2). Accordingly, only three
species are hereafter accepted as O. bourgaeanum (incl.
O. monticolum), O. baeticum (incl. O. algeriense) and
Table 2 Studied morphological characters and values obtained in the
first PCA of all 39 quantitative morphological characters
Character Component
1
Component
2
Length of bulb 0.275 0.393
Diameter of bulb 0.481 0.531
Number of secondary bulbils 0.617 0.012
Number of leaves 0.352 0.360
Maximum length of leaves 0.354 0.588
Maximum width of leaves 0.519 0.316
White band of leaves width 0.451 0.296
Length of the stem 0.154 0.529
Number of flowers 0.184 0.660
Length of the inflorescence 0.311 0.743
Width of the inflorescence 0.637 0.401
Length of the bract 0.506 0.472
Width of the bract 0.719 -0.123
Length of the lowermost floral pedicel 0.578 0.539
Length of the middle floral pedicel 0.648 0.500
Length of the uppermost floral pedicel 0.228 0.011
Insertion angle of the floral pedicels 0.016 -0.017
Insertion angle of the fruiting pedicels 0.334 -0.229
Length of the outer tepal 0.820 -0.071
Length of the inner tepal 0.839 -0.056
Width of the outer tepal 0.809 -0.239
Width of the inner tepal 0.820 -0.198
Width of the green band of the outer tepal 0.758 -0.156
Width of the green band of the inner tepal 0.711 -0.135
Length of the outer filament 0.826 -0.202
Length of the inner filament 0.800 -0.245
Width of the outer filament 0.823 -0.149
Width of the inner filament 0.711 -0.186
Length of the anther 0.723 -0.257
Width of the anther 0.362 -0.427
Length of the ovary 0.808 -0.210
Width of the ovary 0.718 -0.122
Length of the style 0.460 -0.373
Length of the capsule 0.444 -0.040
Width of the capsule 0.339 -0.035
Number of seeds per capsule 0.166 -0.108
Length of seed 0.218 -0.217
Width of seed -0.090 0.161
Weight of seed -0.134 0.171
Taxonomic revision of Ornithogalum subg. 185
123
Author's personal copy
O. divergens. The results of both tests were fully congru-
ent, although the discriminant analysis produced a clearer
segregation (Fig. 2). Cross-validated classification results
of this discriminant analysis showed that more than 95% of
the specimens that were assigned a priori to the three
groups were classified correctly. Four functions were
obtained: the first canonical axis explained 54% of the
variance and was based mainly on the number of bulbils,
and the second one explained the remaining 38.7% of the
variance and was related basically to the style length
(Table 3).
Morphological analysis
The morphological results are presented in relation to the
three taxonomic entities cited above.
Fig. 1 Plot of the first two axes
of the PCA of all 39
morphological characters
Fig. 2 Plot of the two functions
of the discriminant analysis
186 M. Martı
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Bulb
Bulb sizes were similar in the three considered taxa. The
bulb of O. bourgaeanum (incl. O. monticolum) showed the
largest and smallest values, whereas O. baeticum (incl.
O. algeriense) and O. divergens usually showed middle
sizes (Table 4). Regarding the shape of the bulb, O. bae-
ticum and O. bourgaeanum produced ovate bulbs, whereas
the bulb in O. divergens was usually slightly depressed.
The number, size and shape of the bulbils were also sig-
nificant features. O. divergens showed several small,
spherical and pediculate secondary bulbils (Table 4),
whereas the other ones did not show these features, though
they produced the largest and sessile primary bulbils gen-
erated by the division of the mother bulb, sharing its basal
plate. Finally, bulb scales were almost completely conc-
rescent in O. divergens, whereas they were free and
imbricate in O. bourgaeanum and O. baeticum.
Leaves
The number, size and morphology of the leaves were quite
similar among the three studied taxa. The number of leaves
usually varied from four to eight, although in O. bour-
gaeanum (incl. O. monticolum) the number reached 12
(Table 4). The longest and widest leaves were again seen in
O. bourgaeanum, although the usual values were quite
similar with much overlap in the three taxa (Table 4).
Moreover, all three taxa showed a central white band on
the adaxial side of the leaves which was due to an inter-
ruption in the palisade tissue in the central portion of the
leaf (cf. Peruzzi et al. 2007). In the studied taxa, the white
band was very variable in width, although some individuals
of O. baeticum (incl. O. algeriense) showed the widest
white bands (Table 4). Finally, leaves of O. baeticum were
glaucous green, at least when sprouting, whereas the other
taxa had bright green leaves.
Inflorescence
Species of O. subg. Ornithogalum produced inflorescences of
similar length and width, usually corymbose or pseudoc-
orymbose-racemose, with the lowermost pedicels much
longer than the upper ones. The size and shape of the inflo-
rescences, as well as the floral pedicel length were extremely
variable (Table 4). However, the relative disposition of the
lowermost pedicels yielded differences among the studied
taxa. O. bourgaeanum (incl. O. monticolum)andO. baeticum
(incl. O. algeriense) had erect to erect-patent pedicels,whereas
O. divergens showed very long and patent or slightly deflexed
lowermost pedicels, showing clear differences in relation to
the other species. Regarding the bracts, their absolute length
and width were very similar in the three considered taxa, with
much overlap. Floral bracts of O. bourgaeanum and O. bae-
ticum were usually either a bit shorter than the pedicels or a bit
longer, and exceptionally were much shorter and even up to
half of the pedicel length. On the contrary, at least the lower-
most bracts of O. divergens were constantly much shorter than
their pedicels, around half of their length.
Perigone
The length and width of the tepals and their shape in the
three considered taxa were extremely variable. Tepals
varied from acute to obtuse, even in the same population.
Moreover, their size could vary strongly even in a single
individual, when considering mature flowers of the lower
part of the inflorescence or the younger ones from the
upper part. The largest values of tepals was seen in O.
divergens (Table 4), although it produced a lower number
of flowers per inflorescence (Table 4). On the contrary, O.
bourgaeanum (incl. O. monticolum) showed the smallest
tepals (Table 4), although again it generates the highest
number of flowers per inflorescence (Table 4). O. baeticum
(incl. O. algeriense) usually shows intermediate values.
Table 3 Canonical structure
expressing the correlations of
those selected morphological
characters with the highest
standardized significant
contribution to the discriminant
function of the stepwise
discriminant analysis
Characters F-to-remove value Functions
Axis 1 Axis 2
Number of secondary bulbils 87.218 0.835 -0.084
Length of the lowermost floral pedicel 10.094 -0.026 -0.308
Width of the inner tepal 10.824 -0.162 0.245
Width of the green band of the outer tepal 12.141 0.105 -0.411
Length of the inner filament 8.414 0.274 0.125
Width of the outer filament 22.724 0.248 0.069
Width of the inner filament 6.611 -0.279 -0.124
Length of the anther 4.384 0.072 0.272
Length of the ovary 6.475 0.369 -0.020
Width of the ovary 5.175 -0.065 -0.090
Length of the style 88.798 0.082 0.901
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Finally, the width of the dorsal green band of the tepals
was also variable with much overlap among the taxa. The
outer tepals usually showed wider green bands than the
inner tepals. In O. divergens, the band usually reached
7 mm, whereas in the other taxa it did not usually exceed
5 mm (Table 4).
Androecium
The stamen laments of both whorls were lanceolate to
tapering and abruptly contracted at the apex. The inner fila-
ments were slightly broader and longer than the outer fila-
ments, which embrace the septal nectaries of the ovary,
Table 4 Observed values and ranges of all studied morphological characters in Ornithogalum spp.
Character O. algeriense O. baeticum O. bourgaeanum O. divergens O. monticolum
Total bulb length (cm) (1.7) 2.1–3.5 (4) (1.5) 2–3 (3.6) (1.3) 1.4–3.7 (4.1) (1.2) 1.5–3 (3.1) 2.5–3.5 (4.5)
Bulb diameter (cm) (0.9) 1.1–2.8 (0.8) 1.4–2.5 (2.8) (0.8) 1–3 (4) (1.1) 1.5–2.8 (3.5) (1.2) 1.4–2.3 (4)
Number of secondary bulbils 0 0 0 (7) 10–30 (41) 0
Leaf number (3) 4–6 (8) (3) 4–8 (10) (3) 4–10 (12) 4–8 (10) 4–7
Leaf length (cm) (5) 9–25 (30) (8) 10–30 (38) (6) 8–35 (48) (8) 11–35 (10) 12–31
Leaf width (cm) (0.2) 0.4–1.2 (1.5) (0.3) 0.4–1 (1.2) (0.2) 0.3–1 (1.5) (0.2) 0.3–0.8 0.2–0.7
White band width (mm) 0.5–2.5 1–3 (0.2) 0.5–1.5 (2) (0.2) 0.5–1 (1.5) 0.5–1
Stem length (cm) (2.6) 3–6.7 (7.4) (2) 3–11 (19) (1.7) 3–18 (27) (5) 6–13 (17) (5) 8.5–18 (21)
Number of flowers (2) 3–17 (20) (3) 4–18 (21) (3) 4–25 (36) (3) 4–17 (20) (3) 5–10 (13)
Inflorescence length (cm) 2–7 (9) 3–4 (11) (1.5) 2–10 (14) (2) 3–10 (12) (2.1) 3–7.6 (8.0)
Inflorescence width (cm) 2–6 (8) (2) 3–8 (1.7) 2.2–9 (10) (6) 8–16 (18) (2.5) 4–10.5
Bract length (mm) 21–50 (60) (20) 30–70 (15) 17–55 (70) (21) 30–50 (60) (18) 27–42
Bract width (mm) (5) 6–10 (12) (4) 5–10 (12) (3) 4–10 (13) (6) 7–12 (13) 5–8
Lowermost floral pedicel length (mm) (17) 19–32 (40) (21) 25–45 (55) (10) 17–65 (80) (40) 50–100 (120) (20) 33–70 (80)
Middle floral pedicel length (mm) (10) 13–25 (27) (17) 20–40 (48) (10) 14–50 (60) (30) 35–80 (85) 30–50 (54)
Uppermost floral pedicel length (mm) (2) 3–12 (18) (3) 6–25 (35) (1) 2–20 (30) (3) 6–30 (35) (7) 12–30 (35)
Insertion angle pedicels in fruit (308)408–808(908) (308)408–808(908) (308)408–808(908) (758)808–1008 (358)408–508
Outer tepal length (mm) (15) 16–23 (26) (16) 17–25 (27) (9) 12–24 (27) (21) 23–30 13–18
Inner tepal length (mm) (14) 15–22 (24) (15) 16–24 (27) (8) 11–22 (26) (20) 22–26 (28) 12–17
Outer tepal width (mm) (4.5) 5–7.5 (8) 5–9 (10) (3) 4–8 (10) (6) 7–8 (9) 3.5–6
Inner tepal width (mm) (4) 4.5–7 (8) (4) 4.5–8 (9.5) (3) 3.5–6.5 (8) (5) 6–7.5 (8) 3.5–6
Outer tepal green band width (mm) (2) 2.5–4.5 (5) (2) 2.5–4.5 (5) 2–5 (7) (4) 4.5–6.5 (7) (1.8) 2–3
Inner tepal green band width (mm) (1) 1.5–2.5 (3) (0.7) 1–3 (1) 1.2–3.5 (4) 3–4 (4.5) (1) 1.2–2
Outer filament length (mm) 7–9 (10) 7–10 (12) (4.5) 5–9 (10) (6) 10–12 6–8
Inner filament length (mm) 7–11 (7.5) 8–11 (12) (5) 6–10 (12) (6.5) 11–12 (13.5) 6–8
Outer filament width (mm) (1) 1.2–1.7 1.5–2 (2.2) 1–2 (1.8) 2–2.5 (2.8) 1.3–2
Inner filament width (mm) 1.2–2 (2.1) (1.8) 2–2.5 (3) 1.2–2.5 (3.5) (2) 2.2–2.5 (3) 1.3–2
Anther length (mm) 2–3 (3.5) (2) 2.5–3 (4) (1.5) 2–3 (4) 2–3.5 1.5–2.5
Anther width (mm) 0.7–1 0.7–1 0.5–1.5 (2) 0.8–1.5 0.8–1
Ovary length (mm) 3.5–5 (6) 4–6 (7) (2) 2.5–5 (7) 6–7 (8) 3.5–4
Ovary width (mm) (2) 2.2–3.5 (4) (2) 3–4 (5) (1.5) 2–3.5 (4) 3–4 (5) 2–2.5
Style length (mm) 4–5 (5.5) (3.8) 4–6 (6.5) (2) 2.5–3.5 (3.7) 3–3.8 (4) 2–3
Capsule length (mm) 9–10 (10) 12–20 (22) (6) 8–15 (18) (10) 11–18 (20) 9–14
Capsule width (mm) 6–7 (6) 8–13 (5) 6–15 (19) (7) 8–13 (15) 6–9
Number of seeds per capsule 15–28 15–30 (35) (8) 10–30 (35) (10) 11–15 (17) 11–27
Seed length (mm) 2–2.5 (2.8) (2) 2.3–2.6 (2.8) (1.3) 1.6–2.2 (2.4) (1.8) 1.9–2.4 (2.6) 1.6–2.2 (2.4)
Seed width (mm) 1.4–1.7 (1.9) (1.4) 1.5–1.8 (1.9) (1.2) 1.3–1.9 (2.1) (1.3) 1.5–1.7 (1.8) 1.3–2
Seed weight (mg) 2.2–4.5 (2) 2.5–4 (4.9) (1.5) 2–5 (5.7) (2) 2.5–4.2 (4.6) 2–5.1
The data are presented as usual ranges together with the minimum and maximum values in parentheses
188 M. Martı
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gathering the nectar and offering it to the insects. The three
studied taxa showed very similar filaments. However, the
filaments were widest in O. divergensand were a bit smaller in
the other two taxa with much overlap among them (Table 4).
Gynoecium
The ovary was very similar in all studied taxa, being obo-
vate to oblong, truncate at the apex and showing six distinct
ribs in section. However, O. divergens showed the longest
ovary, O. bourgaeanum (incl. O. monticolum) the shortest
ovary, and O. baeticum (incl. O. algeriense) an ovary
intermediate in length (Table 4). O. bourgaeanum showed
the narrowest ovary, and O. baeticum and O. divergens
showed the widest ovary (Table 4). However, there was
great overlap among the studied taxa for all these characters
(Table 4). On the contrary, the absolute length of the style
was a crucial and easy character to differentiate these
taxa, as supported by the discriminant analysis (Table 3).
O. divergens and O. bourgaeanum showed a similar style
length (up to 3.8 (4) mm), whereas in O. baeticum the style
length was usually 4 mm or more (Table 4).
Fruit and seeds
The size and morphology of the capsule allowed the species
to be differentiated. The capsules of all the studied species
were oblong, ovate or obovate, and truncate at the apex
(Figs. 3, 4 and 5; Table 4). Nonetheless, in O. baeticum (incl.
O. algeriense) the capsules were mostly oblong with a length
up to twice the width, whereas in O. bourgaeanum (incl.
O. monticolum) (Fig. 4) and O. divergens (Fig. 5) the cap-
sules were usually ovate or obovate with a length a little more
than or the same as the width. On the other hand, the mor-
phology seen in transverse sections of the capsules showed
interesting differences. In O. baeticum all furrows were
similar, flattened or obtuse, giving a trigonous or polygonal
appearance in section (Fig. 3). However, in O. bourgaeanum
the furrows on the suture of the carpels, coinciding with the
septal nectaries, were deep and wide or obtuse, whereas the
furrows on the midveins of the carpels were much narrower
and acute, giving a paired aspect to the ribs (Fig. 4). Finally,
in O. divergens the capsules had six evident and more or less
equidistant ribs with deep furrows (Fig. 5).
Seeds of the studied taxa were globose and slightly
apiculate (Fig. 6), with the seed testa reticulate. First, O.
baeticum (incl. O. algeriense) showed very prominent ret-
icules, in which the cellular walls could easily be identified
(Fig. 6). Moreover, they were clearly apiculate, or they
could show a distinctive crest from the apex to the opposite
pole. On the other hand, the seeds of O. bourgaeanum (incl.
O. monticolum) and O. divergens (Fig. 6) were globose and
scarcely apiculate and their testa showed less prominent
reticules with the cellular walls not so evident. Regarding
the size of the seeds, O. baeticum showed the largest seeds,
whereas O. bourgaeanum and O. divergens had smaller
seeds with a wide overlap between them (Table 4). Seed
weight was an overlapping character, since O. bourgaea-
num showed the highest and the lowest seed weights, and
both the other species showed medium values (Table 4).
Thus, O. baeticum and O. bourgaeanum showed the highest
number of seeds per capsule, whereas O. divergens showed
the lowest (Table 4). However, there was great overlap in
all these characters. Regarding seed colour, O. baeticum
had bright blackish seeds, whereas O. bourgaeanum and
O. divergens had caramel brownish or dull blackish seeds.
Discussion
After the morphological studies and the statistical analy-
ses, only three biological entities—O. bourgaeanum,
Fig. 3 Morphology of the
capsules of Ornithogalum
baeticum (incl. O. algeriense)
(upper row lateral views, middle
row apical views, bottom row
transverse sections): a S
a
Bermeja, Ma
´
laga, Spain (type
locality); bc Llano de la
Perdiz, Granada, Spain; df S
a
de las Nieves, Ma
´
laga, Spain;
g P.N. Grazalema, Ca
´
diz, Spain;
h Tizi-n-tichka, High Atlas,
Morocco. Vertical scale bar
1cm
Taxonomic revision of Ornithogalum subg. 189
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O. baeticum and O. divergens—were accepted in O. subg.
Ornithogalum in the Iberian Peninsula and the Balearic
Islands. They showed different morphological traits and
particular ecological behaviours. Both O. monticolum and
O. algeriense were indistinguishable from O. bourgaeanum
and O. baeticum, respectively, and hence they are here
included in their synonymy. Combining all morphological
characters yielded poor complete segregation of these taxa,
as shown by the total PCA analysis (Fig. 1), due to their
wide morphological plasticity, and overlap in most of their
vegetative and reproductive characters. This could explain
the confusing taxonomy adopted in this subgenus over
recent centuries. However, among all the studied charac-
ters, some certain vegetative and reproductive features
were revealed as diagnostic for the identification of the
three taxa, as deduced from the values obtained in the
discriminant analysis (Table 3; Fig. 2), and they are dis-
cussed below.
With regard to bulbs, the number and type of bulbils
allowed easy differentiation between O. divergens and both
O. bourgaeanum and O. baeticum. The presence of bulbils
has commonly been used by authors dealing with the
Iberian and Balearic taxa (cf. Coutinho 1913, 1939;
Zahariadi 1980; Pastor 1987; Devesa 1995; Aizpuru et al.
2000; Bolo
`
s and Vigo 2001) as a diagnostic character in
O. subg. Ornithogalum. They characterized O. divergens
(mostly referred as O. umbellatum’) as bearing bulbs with
numerous bulbils, and O. bourgaeanum and O. baeticum
(usually merged under O. orthophyllum subsp. baeticum’)
as bearing bulbs lacking or with very few bulbils. How-
ever, there are important differences not only in the number
of bulbils but in their type, as discussed by Landstro
¨
m
(1989) and Moret (1991). On the one hand, O. divergens
produces many (up to 41) secondary bulbils, called ‘bul-
bils’ by Landstro
¨
m(1989) and ‘bulbilles’ by Moret (1991),
which mostly lack leaves and rest for one or more years
before sprouting and becoming mature plants. They are
solid with a firm external coat, very small (5–8 mm in
diameter), spherical and pediculate, and do not share the
basal plate with the mother bulb, being generated under its
outer tunic. This type of bulbil is found only in polyploid
taxa (cf. Landstro
¨
m 1989). On the other hand, both
O. bourgaeanum and O. baeticum can generate primary
bulbils by splitting of the mother bulb into two or more
identical parts, which were therefore called ‘sister bulbs’
by Landstro
¨
m(1989), and which can sprout and bloom just
from the next year (cf. Moret 1991). These sister bulbs
Fig. 4 Morphology of the capsules of Ornithogalum bourgaeanum
(incl. O. monticolum)(upper row lateral views, middle row apical
views, bottom row transverse sections): a Plasencia, Ca
´
ceres, Spain
(type locality of O. bourgaeanum); b Can
˜
amero, Ca
´
ceres, Spain;
c Navezuelas, Ca
´
ceres, Spain; d P.N. Cornalvo, Badajoz, Spain;
e Barco de A
´
vila, A
´
vila, Spain; f La Alberca, Salamanca, Spain; g P.N.
Caban
˜
eros, Ciudad Real, Spain; h Fuencaliente, Ciudad Real, Spain;
ik Calar del Mundo, Albacete, Spain; l, m Siles, Jae
´
n, Spain; n S
a
de
Alcaraz, Albacete, Spain; o, p Le Mone
ˆ
tier-les-Bains, Hautes Alpes,
France (type locality of O. monticolum); qs Saint-Ve
´
ran, Hautes
Alpes, France; t Cuzcurrita, Zamora, Spain. Vertical scale bar 1cm
Fig. 5 Morphology of the ovaries and capsules of Ornithogalum
divergens (upper row lateral views, middle row apical views, bottom
row transverse sections): a, b Botanic Garden Brest, France; c Maze,
Loire Valley, France; d Jaca, Huesca, Spain; e Luna, Zaragoza, Spain;
f Saumur, Loire Valley, France; g Siena, Tuscany, Italy. Vertical
scale bar 1cm
190 M. Martı
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appear in low numbers, are sessile, large, and share the
basal plate and the outer tunics with the mother bulb. This
second type is more commonly found in the diploid taxa of
the subgenus (cf. Landstro
¨
m 1989).
A third type of bulbil is found in the subgenus, which
includes few, leafy and medium-sized secondary bulbils,
external to the outer tunic of the mother bulb. They were
named ‘caı
¨
eux’ by Moret (1991) or ‘side bulbs’ by Lands-
tro
¨
m(1989), and can be observed in the type (iconotype)
O. umbellatum s.s. (cf. Stearn 1983). As we have not found
any Iberian or Balearic plant matching these types of sec-
ondary bulbils, according to van Raamsdonk (1986)we
excluded the presence of O. umbellatum in both territories.
Summarizing, the depressed bulb shape and concrescence of
the bulb scales in O. divergens (cf. Speta 2000), together with
the number and mainly the type of bulbils, are shown here
to be crucial characters for segregating that taxon from
O. bourgaeanum and O. baeticum (Fig. 2; Tables 3 and 4).
Moreover, in both latter species bulb scales were free and
imbricate, allowing easy distinction from O. kochii, in which
they are completely concrescent (cf. Garbari et al. 2008).
Morphology and the size of leaves have also been used
for taxonomy of O. subg. Ornithogalum (cf. Zahariadi
1980; Pastor 1987; Devesa 1995; Franco and Rocha
Afonso 1994). Similarly, Bolo
`
s and Vigo (2001) have
argued that the existence of a marked central white band on
the adaxial side of the leaf allow the differentiation of
O. divergens (as ‘O. umbellatum subsp. umbellatum’) from
both O. bourgaeanum (as O. umbellatum subsp. monti-
colum’) and O. baeticum (as O. umbellatum subsp. bae-
ticum’), in which it was weakly visible. However, our
studies on the Iberian and Balearic plants show extreme
variability in the length and width of the leaves, as well as
in the features of the central band that is present in all
individuals, with a wide overlap among the three accepted
taxa (Table 4
). Conversely, the colour of the young leaves
can be used to differentiate O. baeticum (with glaucous
leaves) from O. bourgaeanum and O. divergens (both with
bright green leaves), as pointed out by Moret and Galland
(1992). Therefore, on the basis of leaf characters, O. exs-
capum, a plant described from Naples (Italy) and distrib-
uted throughout the central and eastern Mediterranean (cf.
Fig. 6 Seeds of the species of
Ornithogalum subg.
Ornithogalum: a, b O. baeticum
(Ca
´
diz, Spain); c, d O.
bourgaeanum (Alicante, Spain);
e, f O. divergens (Angers,
France). Scale bars: a, c,
e 1 mm; b, d, f 100 lm
Taxonomic revision of Ornithogalum subg. 191
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Landstro
¨
m 1989; Garbari et al. 2008), should be excluded
from the Iberian and Balearic floras since its leaves do not
show such a white band (cf. Tenore 1811). All references
to its presence in Sacromonte, Granada (Lange 1860;
Willkomm 1862; del Amo y Mora 1871; Colmeiro 1889)
surely correspond to young, immature forms of O. baeti-
cum, and hence do not show the characteristic deflexed
fruiting pedicels of O. exscapum. In a similar way, all
Iberian cites of O. collinum should be rejected, since this
species bears leaves ciliate on the margins (cf. Gussone
1825; Garbari and Giordani 1984), whereas all studied
Iberian and Balearic plants show completely glabrous
leaves.
Some authors have used the inflorescence structure
(Coutinho 1913, 1939; Aizpuru et al. 2000; Bolo
`
s and Vigo
2001) and/or the length of the floral bracts relative to their
pedicels (cf. Coutinho 1913, 1939; Zahariadi 1980; Bolo
`
s
and Vigo 2001) as diagnostic characters. In agreement with
those authors, our data show that O. divergens produces
wide inflorescences with very long and patent to deflexed
pedicels, with the length of the lowermost bract reaching
about half the length of its pedicel. On the contrary,
O. bourgaeanum and O. baeticum showed narrower inflo-
rescences with shorter and erect-patent to patent pedicels,
and the length of most of the bracts usually three-quarters
or more of the length of their pedicels (Table 4).
Morphology and size of the tepals have traditionally
been used to describe or to discriminate different taxa (cf.
Jordan and Fourreau 1866; Pastor 1987; Bolo
`
s and Vigo
2001). In fact, PCA shown in Fig. 1 revealed that the
length and width of the tepals represented an important
percentage of the variance (Table 2), mostly when the taxa
were compared by pairs. Thus, O. bourgaeanum produces
smaller tepals than O. divergens, although O. baeticum was
in an intermediate position and there was a wide overlap
between the taxa. Therefore, our studies revealed that the
flower characters vary widely, even in a single population
or individual, it being common that tepals are usually much
larger in the lower flowers of a given inflorescence. Hence,
they are only useful for differentiation between some par-
ticular taxa, and not in a global sense.
As previously remarked by Moret and Galland (1992),
the style length is as a crucial character to segregate
O. baeticum, with consistently longer styles, from
O. bourgaeanum and O. divergens, in which they are
shorter (Tables 3 and 4). This character itself allows an
easy and confident identification of herbarium sheets
lacking bulbs, leaves or capsules.
Capsule morphology is an interesting character to be
considered for taxonomy of the Iberian and Balearic taxa
of O. subg. Ornithogalum. Traditionally, segregation of
O. divergens (referred as O. umbellatum’) from O. bour-
gaeanum and O. baeticum (both united as ‘O. orthophyllum
subsp. baeticum’) has been based on the disposition of the
capsule ribs, which are equidistant in the former and
grouped by pairs in the two latter (cf. Coutinho 1913, 1939;
Zahariadi 1980; Pastor 1987; Aizpuru et al. 2000; Bolo
`
s
and Vigo 2001). However, the situation appears to be a bit
more complicated. On the one hand, capsules of O. di-
vergens bear six prominent and more or less equidistant
ribs (Fig. 5), a quite homogeneous character in this taxon.
On the other, capsules of O. baeticum are more or less
polygonal in section, with all the furrows between the ribs
(both on sutures and the midveins of the carpels) similarly
obtuse or flattened (Fig. 3) and not clearly paired. Finally,
capsules of O. bourgaeanum are extremely variable in size
and shape, although they can be recognized by the nar-
rower and acute furrows along the midveins of the carpels,
usually giving ribs a markedly paired appearance (Fig. 4).
Other features such as the shape and size of the
androecium, ovary and seeds are not absolutely conclusive
for the taxonomy of the three considered species. None-
theless, they are informative enough when combined with
other diagnostic characters.
Three types of seeds were described by Moret et al.
(1990) on the basis of the ornamentation pattern of testa
cells. Although these types were thought to characterize
different subgenera of Ornithogalum, a direct relationship
has not been demonstrated as shown by Coskuncelebi et al.
(2000) and Moret et al. (1990), who found more than one
seed type in various Eurasian sections of the genus.
However, all species of O
. subg. Ornithogalum show typi-
cally globose and slightly apiculate seeds with reticulate
testa (type 1, sensu Moret et al. 1990). Therefore, this testa
character can be regarded as an apomorphy of this group
(cf. Martı
´
nez-Azorı
´
n et al. 2010b).
The complex caryology of the Iberian and Balearic
members of O. subg. Ornithogalum is related to the high
frequency of ploidy, aneuploidy, translocations and dele-
tions (Neves 1952). All these processes have important
implications in the morphology and reproductive behaviour
of plants (van Raamsdonk 1986; Moret and Favereau 1991;
Moret and Galland 1992), since usually they constitute
reproductively isolated populations with autonomous
descendent lineages (van Raamsdonk 1985a, b) and can
generate morphological aggregates with complex taxon-
omy. Accordingly, some authors accept several species in
the polyploid complex of O. umbellatum sensu lato (Gad-
ella 1972; Gadella and van Raamsdonk 1981; Raamsdonk
1986), though others only accept just one variable species
with different ploidy levels (Moret and Galland 1992).
As deduced from previous morphological and cary-
ological combined studies carried out on Iberian plants,
O. bourgaeanum is mainly a diploid species (2n = 18)
or exceptionally triploid (2n = 27) (cf. Neves 1952, 1973;
Gime
´
nez-Martı
´
n 1958;Lo
¨
ve and Kjellquist 1973;
192 M. Martı
´
nez-Azorı
´
n et al.
123
Author's personal copy
Ruiz-Rejo
´
n 1978; van Raamsdonk 1986; Moret et al. 1987;
Moret and Galland 1992), whereas O. divergens is usually
a hexaploid species (2n = 54) (cf. Neves 1952, 1956;
Gadella and van Raamsdonk 1981), both belonging to
the cited O. umbellatum complex’ (x = 9). Conversely,
O. baeticum is an amphidiploid (2n = 50, 52, 54; x = 25)
(cf. Ruiz-Rejo
´
n 1974, 1978; van Raamsdonk 1986; Moret
and Galland 1992) derived from an uncertain ancient
allopolyploid crossing, in which both a polyploid taxon of
the O. umbellatum complex and a relative of the current
lineage of O. platyphyllum Boiss. would act as presumed
parents (van Raamsdonk 1986). Therefore, Iberian cites of
O. gussonei Ten. (=O. tenuifolium Guss., non Redoute
´
)by
Willkomm (1893), Ruiz-Rejo
´
n(1978) and Franco and
Rocha Afonso (1994) should be excluded, since it is an
eastern Mediterranean plant with chromosome number
2n = 14 (x = 7) (cf. Garbari and Tornadore 1971;
Landstro
¨
m 1989; Garbari et al. 2003), quite different from
the Iberian diploids 2n = 18 (x = 9). Moreover, it pro-
duces concolour and filiform leaves which have not been
observed in any Iberian material.
Preliminary results of the phylogenetic studies we are
carrying out based on sequences of plastid and nuclear
DNA regions (Martı
´
nez-Azorı
´
n et al. 2010b) place the taxa
of O. subg. Ornithogalum (incl. O. subg. Myogalum)ina
monophyletic clade, clearly apart from other Mediterra-
nean subgenera such as Cathissa or Beryllis. Furthermore,
our AFLP studies have also revealed a wide differentiation
among the three species here accepted (Martı
´
nez-Azorı
´
n
et al., unpublished data). Such molecular markers indicate
a closer position of O. bourgaeanum and O. divergens with
regard to O. baeticum, as suggested by van Raamsdonk
(1986) and Moret and Galland (1991) on the basis of
caryological and morphological data. Moreover, other
related European taxa such as O. orthophyllum, O. kochii,
O. exscapum, O. collinum and O. gussonei, show a suffi-
cient genetic differentiation from the Iberian and Balearic
populations, warranting recognition at the species rank
(Martı
´
nez-Azorı
´
n et al., unpublished data). Conversely,
North African populations of O. algeriense are mixed with
those of O. baeticum, a result that supports synonymization
under the earlier name O. baeticum. In a similar way,
French populations of O. monticolum from the Maritime
Alps (its type locality) and Central Plateau, nest near the
base of all samples of O. bourgaeanum, with a very small
genetic distance. Aquaro and Peruzzi (2006) treated the
two taxa as belonging to two different lineages clearly
differing in the satellite type of one chromosome pair: the
former (treated as O. kochii subsp. monticolum (Jord. &
Fourr.) Peruzzi) showing terminal satellites, and the latter
(treated as O. orthophyllum subsp. baeticum) bearing
intercalary satellites. Their conclusion would also be sup-
ported by certain differences revealed by chromosome
banding (van Raamsdonk 1986). However, caryological
data reported previously by Moret et al. (
1987) show that
both terminal and intercalary satellites occur in different
Iberian populations (e.g. terminal in Leo
´
n, La Pola de
Gordo
´
n, and intercalary in Madrid) of the diploid taxon
here accepted as O. bourgaeanum, and therefore are not a
diagnostic character for specific differentiation. Similarly,
van Raamsdonk (1985a, b) found that crosses between
French and Iberian diploid plants are fully compatible,
yielding similar or even higher seed sets than crosses
between only French diploid plants. All these molecular,
caryological and reproductive data together with the wide
morphological plasticity observed in wild populations of
the two taxa suggest treating O. bourgaeanum and O.
monticolum as synonyms. However, further molecular
studies are still needed to clarify the relationships
among all European and North African diploids of the
O. umbellatum complex’.
Taxonomic treatment
Ornithogalum subg. Ornithogalum. Ornithogalum subg.
Heliocharmos Baker in J. Linn. Soc. (Bot.) 13: 258. 1872.
Ornithogalum sect. Heliocharmos (Baker) Engl., Nat.
Pflanzenfam. [Engler & Prantl] 4: 67. 1888. LECTOTYPE:
O. umbellatum L. (cf. Stearn 1983).
Celsia Heist. ex. Fabr., Enum. (ed. 2). [Fabr.]. 22. 1763,
pro parte, nom. illeg. [non Celsia L., Sp. Pl.: 621. 1753;
Scrophulariaceae]. LECTOTYPE: O. umbellatum L.
Key to the Iberian-Balearic taxa of Ornithogalum subg.
Ornithogalum
1. Bulb slightly depressed, surrounded by (7)10–30(41)
spherical and secondary bulbils, about 5 mm in length
and pediculate (1–8 mm). Mature inflorescence
(6)8–16(18) cm width (excluding flowers), lowermost
pedicels very long, (40)50–100(120) mm, patent or
slightly deflexed. Lowermost bracts around half the
length of their pedicels O. divergens
Bulb ovate, usually simple, or one to seven larger
(1–3 cm long) and sessile primary bulbils, usually
included in the outer tunic and sharing the basal plate of
the mother bulb. Mature inflorescence (1.7)2.2–9(10)
cm width (excluding flowers), lowermost pedicels
shorter, (10)17–65(80) mm, erect or erect-patent.
Lowermost bracts usually much longer — 2
2. Style (2)2.5–3.5(3.7) mm. Capsule pyriform,
(6)8–15(18) 9 (5)6–15(19) mm, with furrows on
suture carpels deep, wide and obtuse, although those
on midvein of carpels much narrower and acute, giving
Taxonomic revision of Ornithogalum subg. 193
123
Author's personal copy
a paired appearance in section. Young leaves bright
green. Seeds slightly apiculated with few evident
crests, where the cellular walls cannot be differentiated
O. bourgaeanum
Style (3.8)4–6(6.5) mm. Capsule oblong, (9)12–20
(22) 9 (6)8–13 mm, both suture and midvein furrows
similar, wide, flattened or obtuse, giving a polygonal
appearance in section. Young leaves glaucous. Seeds
strongly apiculated with prominent crests, where the
cellular walls can be observed O. baeticum
Ornithogalum baeticum
Ornithogalum baeticum Boiss., Elenchus 84. 1838. Orni-
thogalum orthophyllum subsp. baeticum (Boiss.) Zahar. in
Bot. J. Linn. Soc. 76(4): 356. 1978. Ornithogalum alger-
iense subsp. baeticum (Boiss.) Moret in Pl. Syst. Evol. 181:
200. 1992. Ornithogalum umbellatum subsp. baeticum
(Boiss.) O. Bolo
`
s & Vigo, Fl. Paı
¨
sos Catalans 4: 87. 2001.
Ornithogalum umbellatum var. longebracteatum Willk. in
Willk. & Lange, Prodr. Fl. Hispan. 1: 216. 1862. TYPE:
SPAIN. In summa
ˆ
Sierra d’Estepona [Ma
´
laga], May 1837,
Alt. 4,000
0
(lectotype: G-BOIS! The lowermost individual
on the right part of the sheet number one, as designated by
Burdet et al. 1982).
Ornithogalum algeriense Jord. & Fourr., Brev. Pl. Nov.
1: 52. 1866. TYPE: ALGERIA. In arvis Algeriae, circa
Alger, 1862, unknown collector (lectotype: LY!, desig-
nated by van Raamsdonk 1982).
Ornithogalum algeriense subsp. atlanticum Moret in
Canad. J. Bot. 66: 2185. 1988. TYPE: MOROCCO. Bou-
znika, re
´
gion de Rabat, 334.8 9 357.6, 13 Mar 1984,
Moret 84-24-01 (holotype: PCU!).
Ind. loc.
‘Hab. in summo monte Sierra d’Estepona alt. circ.
4,000
0
’’.
Illustrations
Jordan & Fourreau, Ic. Fl. Eur. 1: Tab. 74. 1867; Fig. 7.
Description
Geophyte. Bulb (1.5)2–3(4) 9 (0.8)1.4–2.5(2.8) cm, ovoid-
oblong with whitish outer tunics, generating contractile
roots, without secondary bulbils, sometimes with primary
bulbils in low number, which are generated by splitting of the
main bulb into similar parts, sharing the basal plate and the
outer tunics. Leaves (3)4–8(10), in basal rosette,
(5)10–30(38) 9 (0.2)0.4–1(1.5) cm, linear-tapering, erect,
slightly keeled, green, glaucous when sprouting, glabrous,
with a longitudinal white band 1–3 mm wide, with withering
apex, synanthous. Floral stem (excluding the inflorescence)
(2)3–11(19) 9 0.3–0.4 cm, erect, smooth, green, glaucous
and glabrous. Inflorescence corymbose-racemose, (2)
3–11 9 (2)3–8 cm (excluding the flowers but not their
pedicels), with (2)4–18(21) flowers; bracts (20)30–70 9
(4)5–10(12) mm, a bit shorter or longer than the pedicels,
ovate-lanceolate or triangular, with acuminate-acute apex,
membranous with various greenish nerves; floral pedicels
erect-curved, the lower ones (17)25–45(55) mm, the middle
ones (10)13–40(48) mm and the upper ones (2)3–25(35)
mm; fruiting pedicels (15)20–55(60) mm, curved and
ascending. Flowers 30–40 mm diameter, few fragrant; tepals
white in the adaxial side and white with a central green band
in the abaxial side, acute or obtuse; outers lanceolate-
obovate, (15)16–25(27) 9 4.5–9(10) mm, with green band
(2)2.5–4.5(5) mm wide; inners lanceolate, (14)15–24
(27) 9 (4)4.5–8(9.5) mm, with green band (0.7)1–3 mm.
Stamens six, one-half to two-fifths the length of the tepals;
filaments white, lanceolate-tapering, with the greatest width
in the basal half and narrowing progressively towards the
apex; the outers 7–10(12) 9 (1)1.2–2(2.2) mm; the inners
(7.5)8–11(12) 9 (1.2)2–2.5(3) mm, a bit longer and wider
than the outers; anthers dorsifixed, pale yellow or whitish,
4–5 9 1.2–1.5 mm before dehiscence and (2)2.3–3(3.5) 9
0.7–1(1.2) mm after dehiscence. Ovary 3.5–6(7) 9
(2)3–4(5) mm, pale green, obovate or cylindrical, truncate in
the apex, with six evident ribs and septal nectaries;
style whitish, filiform, (3.8)4–6(6.5) mm; stigma slightly
trigonous and glandulous. Capsule (9)12–20(22) 9 (6)
8–13 mm, oblong to obovate, with trivalvar dehiscence,
truncate in the apex, with six ribs and similar suture and
midvein furrows, which are obtuse or flattened, giving a
polygonal appearance in section. Seeds 15–30(35) per fruit
(
X ¼ 24), (2)2.3–2.6(2.8) 9 (1.4)1.5–1.8(1.9) mm, black or
sugar-brown colour, shining, globose, strongly apiculate,
(2)2.5–4.5(4.9) mg in weight; testa of type 1 (reticulate),
with reticules formed by very prominent crests, where the
cellular walls can be distinguished.
Chromosome number
2n = 54 (Ruiz-Rejo
´
n 1974, 1978 as O. umbellatum).
2n = 50 (van Raamsdonk 1986 as O. algeriense). 2n = 52
(Moret and Galland 1992 as O. algeriense).
Biology
Flowers from March to June, even July in higher areas.
Fructification occurs from May to July or August. This
species does not produce secondary bulbils, although the
194 M. Martı
´
nez-Azorı
´
n et al.
123
Author's personal copy
mother bulb can split in several parts that share the basal
plate and are similar in size. This usually follows trans-
plantation or can occur under conditions of stress such as
severe drought.
Ecology
This species grows mainly in open mountainous stony and
relatively dry grasslands, although it can reach forests or
coastal areas, including marshes. In the Iberian Peninsula,
it ranges from sea level to 1,600 m altitude, although in
northern Africa it extends up to 2,600 m altitude in the
High Atlas. It seems to prefer warm areas.
Distribution
North Africa (Morocco and Algeria) and the Iberian Pe-
ninsula. The Iberian populations are mainly distributed in
Fig. 7 Ornithogalum baeticum
(Grazalema, Ca
´
diz, Spain):
a flower, frontal view; b flower,
dorsal view; c whole plant;
d bract; e inner stamen (left) and
outer (right); f gynoecium;
g capsule, lateral view;
h capsule, apical view;
i capsule, transverse section;
j portion of one leave; k bulb.
Scale bars 1cm
Taxonomic revision of Ornithogalum subg. 195
123
Author's personal copy
the south (Andalucı
´
a, except Almerı
´
a and east of Granada
and Jae
´
n), though some disjoint populations are found in
the centre (environs of Madrid) and north (Sistema Ibe
´
rico)
(Fig. 8).
Observations
O. baeticum exhibits a wide morphological variability in
leaves, flowers and inflorescences. Some individuals of the
type locality, including the holotype, are sometimes small,
much resembling O. bourgaeanum. This fact led several
authors (cf. van Raamsdonk 1986; Aquaro and Peruzii 2006)
to misunderstand the type of O. baeticum and to assimilate it
with O. bourgaeanum, leaving O. algeriense for most of
the southern Iberian populations. However, the style of
O. baeticum (including plants of the type locality) usually
reaches or exceeds 4 mm in length; the capsule morphology,
oblong, with flattened or obtuse furrows on both sutures and
midveins of carpels; and the glaucous colour of the young
leaves, allow easy differentiation between the two species.
Moreover, the type specimens of O. algeriense conserved at
LY bear styles about 4.5 mm in length (Thie
´
baut pers.
comm.), perfectly matching O. baeticum and hence justify-
ing synonymization of the two taxa. Preliminary results of
AFLP studies (Martı
´
nez-Azorı
´
n et al., unpublished data) also
support this solution. The disjoint populations of O. baeticum
from the north of the Iberian Peninsula were consistently
smaller than those from the south, probably due to their more
stressful high-altitude habitat. The actual populations of this
species are numerous and are composed of many individuals;
therefore, it seems that it is not endangered in short or
medium term. However, the disjoint populations in the north
are scarce and they are composed of few individuals;
therefore, the evolution of these populations should be fol-
lowed to evaluate their risk of decline or extinction in this
area. IUCN (2001) category: LC.
Representative specimens examined
Spain Badajoz: Bienvenida, Sierra de Bienvenida, 21 Apr
1994, 700–750 m, 29SQC4840, E. Rico, F. Amich et al.,VI
Itinera Mediterranea, no. 348 (MA 717968); Olivenza,
Sierra de Alor, 19 Apr 1973, 600 m, A. Segura Zubizarreta
s.n. (MA 355364); Usagre, 10 May 1971, B. Casaseca s.n.
(OVI 05302); Manchita, ‘La Moneda’’, 14 Apr 1981,
T. Ruiz Te
´
llez s.n. (SALA 20779).
Ca
´
diz: Villaluenga del Rosario, Llanos del Republicano
(P.N. Grazalema), 1 May 2003, 840 m, 30STF9063, M.
Martı
´
nez-Azorı
´
n s.n. (ABH 47137); Barbate de Franco, 25
Apr 1979, 10 m, TF3809, Akina, Barra,
Lo
´
pez & Morales
s.n. (MA 503052); Jimena de la Frontera,
´
o Hozgarganta
cerca del pueblo, 26 Mar 2005, 30STF7934, M. Martı
´
nez-
Azorı
´
n s.n. (ABH 50149); Ubrique, Casa de la Patilla, 30
Apr 1983, TF8158, A. Aparicio & S. Silvestre s.n. (BC
806294); Camino al Can
˜
o del Agua, presa del Guadarran-
que, 1 May 1974, A. Asensi & B.
´
ez s.n. (MGC 1080);
El Cobre, Algeciras, Apr 1980, E. Salvo Tierra s.n. (MGC
6803); Sotogrande, 9 Apr 1971, S. Holmdahl s.n. (MGC
50633); Tarifa, Puerto del Cabrito, 27 Mar 1969, V.H.
Heywood, D.M. Moore et al. 162 (SEV 13678); Chiclana,
pinares de La Barrosa, 24 Mar 1968, N. Gutie
´
rrez,
S. Silvestre & B. Valde
´
s 1233/68 (SEV 20123); Entre
Jerez de la Frontera y El Puerto de Sta. Marı
´
a, 9 Mar 1978,
J. Pastor, S. Talavera & B. Valde
´
s s.n. (SEV 120211).
Fig. 8 Map of studied
populations of Ornithogalum
baeticum in the Iberian
Peninsula. Dots correspond to
both studied natural populations
and revised herbarium sheets
196 M. Martı
´
nez-Azorı
´
n et al.
123
Author's personal copy
Co
´
rdoba: Estacio
´
n del Zu
´
jar, 23 Apr 1982, J.A. Devesa,
T. Luque & B. Valde
´
s s.n. (SEV 120228); Be
´
lmez, Sierra
Palacios, 24 Mar 1978, J. Pastor s.n. (SEV 79638); Espiel,
Arroyo del Valle y faldas del Cerro del Molino, 20 Apr
1979, M.J.
´
az, P. Ferna
´
ndez & J. Mun
˜
oz s.n. (SEV
120465); Fuenteovejuna, 28 Apr 1979, E.F. Galiano, A.
Ramos & R. Elvira s.n. (SEV 121727); Posadas,
´
o
Guadazueros, 28 Mar 1980, P. Ferna
´
ndez & J. Varela s.n.
(SEV 120214); Entre Palma del
´
o y Hornachuelos, 21
Mar 1978, S. Talavera & B. Valde
´
s s.n. (SEV 217426);
Priego de Co
´
rdoba, desfiladero de Las Angosturas, 30 Mar
1979, M.L.
´
az, E. Dominguez & J. Mun
˜
oz s.n. (SEV
121818); Cabra, 3 May 1918, C. Vicioso s.n. (BC 61959,
MA 21767); Entre Fuencubierta y Fuente Palmera, 16 Mar
1979, J. Mun
˜
oz & J. Varela s.n. (SEV 120140); Carretera
de Zuheros a Luque, 23 Feb 1978, M.L.
´
az, E.
Domı
´
nguez et al. s.n. (SEV 121686).
Granada: Granada, Llano de la Perdiz, 16 Apr 2006,
1,016 m, 30SVG506145, M. Martı
´
nez-Azorı
´
n s.n. (ABH
51029); Sacromonte, cerca de la capital, 8 May 1949,
Mun
˜
oz Medina s.n. (GDA 37593, MA 345622); Gu
¨
ejar-
Sierra, Laguna de la Trinchera, 12 May 1989, 30SVG6013,
M
a
T. Vizoso & M. Zea s.n. (GDA 44139, GDA 44131);
Iznalloz, subida al Cerro del Frage, vertiente noroeste, 28
May 1989, 1,160 m, 30SVG4739, L. Gutie
´
rrez & F.J.
Garcı
´
a s.n. (GDA 44132); Loja, base del Hacho, orienta-
cio
´
n sur, 13 Mar 2001, 650 m, 30SVG0015, L. Baena & C.
Morales s.n. (GDA 44118); Cogollos de Vega, camino al
cortijo de Carialfaquı
´
n, 14 Apr 1988, 1,100 m,
30SVG5025, L. Cano s.n. (GDA 21246); Sierra Elvira,
23 Mar 1989, 30SVG3922, M
a
T. Vizoso Paz & A. Vizoso
Gonza
´
lez s.n. (GDA 44137); Ca
´
diar, valle del
´
oCa
´
diar,
bajo el pueblo, 5 Apr 2001, 700 m, 30SVF8387, C.
Morales, C. Quesada, L. Baena & M
a
T. Vizoso s.n. (GDA
44120); Montefrı
´
o, Illora, base de la S
a
de Parapanda, 18
May 2000, 1,500 m, 30SVG1830, C. Morales, C. Quesada,
L. Baena & E. Linares s.n. (GDA 42849); S
a
de Alfacar,
Apr 1963, F. Valle s.n. (GDAC 10026); Sierra Nevada,
Hotel del Duque, 1 May 1968, 1,170 m, C. Morales s.n.
(GDAC 780); Sierra Tejeda, Llanos de la Alcaicerı
´
a, 9 Apr
1980, J.M. Nieto s.n. (MGC 7507).
Guadalajara: Uceda, puente del
´
o Jarama, 23 Apr
1987, 30TVL5921, R. Morales 76RM, P. Blanco & S.
Cirujano s.n. (MA 448342).
Huelva: Palma del Condado, 16 Mar 1968, J. Borja, J.
Izco & J. Mansanet s.n. (MA 258866, VAL 128486); Pinar
de Bollullos del Condado, 19 Mar 1966, E.F. Galiano & J.
Novo s.n. (SEV 120146); Carretera de Almonte al Parador,
21 Mar 1982, J. Herrera s.n. (SEV 120215); Entre
Valdeflores e Higuera de la Sierra, Dehesa la Corchuela,
26 Apr 1975, B. Cabezudo s.n. (SEV 115538); Punta
Umbrı
´
a, 20 Mar 1982, M.J. Diez s.n. (SEV 120206);
Entrada a Santa Olalla, 18 May 1978, P. Candau, J.
´
ez
et al. s.n. (SEV 120225); Ayamonte, isla Canela, 6 Apr
1979, J.
´
ez, S. Talavera & B. Valde
´
s s.n. (SEV 121728);
El Rocio, Coto del Rey, Almonte, Puente del Ajolı
´
, 20 Mar
1981, J.A. Devesa, I. Fernandez, T. Luque, J. Pastor & S.
Talavera s.n. (SEV 61975).
Jae
´
n: Jae
´
n, La Imora, 21 Apr 1986, 600 m, 30SVG2782,
A. Crespi s.n. (BCN 46315, MA 505399, MGC 33940, VAL
77746); Prox. Estacio
´
n de Alcaudete, 10 Apr 1984, 500 m,
M. Ortiz & F. Valle s.n. (GDAC 17850).
La Rioja: Zubia, 4 May (MA 21804); Logron
˜
o, Recajo,
rives de l’Ebre, 10 May 1923, H. Elı
´
as s.n. (BCN 46329,
BCN 46318).
Madrid: Soto del Real, 1 May 1981,
D. Sa
´
nchez Mata
s.n. (OVI 10587); Alcala
´
(GDA 37610); Herbazales de El
Pardo, 4 May 1940, S. Rivas Goday s.n. (MA 202207);
Rivas de Jarama, 28 Apr 1918, C. Vicioso s.n. (MA 21808);
Lagunas de la Moncloa, 12 Apr 1919, H. Villar s.n. (MA
158388); Soto del Real, 1 May 1981, D. Sa
´
nchez Mata s.n.
(MA 258223); San Agustı
´
n de Guadalix, 4 Apr 1981,
750 m, 30TVL4806, J.C. Moreno Sa
´
iz s.n. (MA 451749);
Guadalix de la Sierra, 860 m, 30TVL4012, F. Go
´
mez
Manzaneque s.n. (MA 453059); El Vello
´
n, 12 May 1989,
870 m, 30TVL5213, F. Go
´
mez Manzaneque s.n. (MA
453060); Reduen
˜
a, 1 May 1983, 910 m, 30TVL5016, F.
Go
´
mez Manzaneque s.n. (MA 453058).
Ma
´
laga: Estepona, S
a
Bermeja, subida al pico Reales, 12
May 2005, 1,200 m, 30SUF0239, M. Martı
´
nez-Azorı
´
n,
M.B. Crespo & C. Pena s.n. (ABH 50150); Sierra Bermeja,
cerro La Real, 20 May 1919, Gros s.n. (BC 61963); Cortes
de la Frontera, La Sauceda (P.N. Los Alcornocales), 2 May
2003, 550 m, 30STF6846, M. Martı
´
nez-Azorı
´
n s.n. (ABH
47135); Antequera, El Torcal de Antequera, 6 Jun 1989,
30SUF6395, M
a
T. Vizoso Paz & J.C. de la Rosa Alamos
s.n. (GDA 44171); El Burgo, S
a
de las Nieves N, mirador
del Guarda Forestal, 8 Apr 2005, 30SUF2273, A. Martı
´
nez
& E. Azorı
´
n s.n. (ABH 50148); Ronda, La Nava de San
Luis, 4 May 1974, A. Asensi & B.
´
ez s.n. (MGC 1077);
Benaoja
´
n, Sierra del Palo, vereda entre la Fuente del
Saucillo y el Mayo del Artesa
´
n, 25 May 2002, 1,140 m,
30STF9462, M. Becerra, F. Sa
´
nchez Tundider s.n. (MGC
51981); Montejaque, El Quejigal, 5 May 1991, Martı
´
n&
Soria s.n. (MGC 31992); Casares, playa de Casares, 7 Apr
1976, UF02, A. Asensi & B.
´
ez s.n. (MGC 3402); Sierra
de Almola, Cartagima, 23 May 1932, L. Ceballos & C.
Vicioso s.n. (MA 21772); Ma
´
laga, Sierra del Hacho, 10 Apr
1974, 600–800 m, S. Talavera & B. Valde
´
s s.n. (SEV
119867); Alpandeire, Puerto de Encinas Borrachas, 16 Apr
1987, 1,000 m, 30SUF0261, D. Montilla s.n. (MGC 40529).
Navarra: Mendavia, 10 Apr 1988, 350 m,
30TWN5502, B. Fz. De Beton
˜
o & J.A. Alejandre s.n.
(MA 484245); Viana, Cicujano, Escuela Agraria, 11 May
2006, 375 m, 30TWN577026, P.M. Uribe-Echebarrı
´
a s.n.
(ABH 51044).
Taxonomic revision of Ornithogalum subg. 197
123
Author's personal copy
Segovia: La Granja?, 1 Jun 1893, A.E. Lomax s.n. (MA
21803).
Sevilla: Alcala
´
de Guadaira, Ermita de San Roque, 6
Apr 2005, 100 m, 30STG5235, M.A. Alonso, A. Juan, J.
Monerris & J.J. Montoya s.n. (ABH 50984); Almade
´
ndela
Plata, Finca El Berrocal, Arroyo Quevedo, 5 Apr 1982, A.
Aparicio, F. Garcı
´
a & S. Silvestre s.n. (SEV 121817);
Puebla de los Infantes, Arroyo Tamujoso, 11 Apr 1982,
100–400 m, M. Barrera, T. Caballero et al. s.n. (SEV
120132); El Garrobo, 1 Apr 1979, I. Ferna
´
ndez s.n. (SEV
79629); Sevilla, Universidad Laboral, alrededores, 12 Mar
1980, T. Luque & B. Valde
´
s s.n. (SEV 217461); Coria del
´
o, 23 Mar 1968, S. Silvestre & B. Valde
´
s s.n. (SEV
20125); Coripe, Pen
˜
o
´
n de Zaframago
´
n, 18 Mar 1984,
Martı
´
n, Mun
˜
oz, Ordon
˜
ez & Sequeiros s.n. (SEV 105949);
Entre Moro
´
n y Pruna, 18 Mar 1977, E. Ruiz de Clavijo s.n.
(SEV 31981); Villamanrique de la Condesa, Raya Real y
camino, 5 Mar 1978, J. Ferna
´
ndez & P. Lo
´
zar s.n. (SEV
120463); Alga
´
mitas, Pen
˜
o
´
n de Alga
´
mitas, 1 Jun 1975, E.
Ramos s.n. (SEV 31998); Guillena, Arroyo Carrascosa, 22
Mar 1981, J. Otal, M.J. Torrescusa et al. s.n. (SEV
120223); Sierra de Espartero, Moro
´
n, 24 Apr 1933, C.
Vicioso s.n. (MA 21769); Castilblanco de los Arroyos,
urbanizacio
´
n Las Minas, 17 Apr 1981, C. Andre
´
s, M.A.
Cobo et al. s.n. (SEV 121726).
Soria: Olvega, alto del puerto de la Carrasca, 9 May
1993, 1,180 m, 30TWM82, G. Mateo s.n. (VAL 80040);
Ventosilla de San Juan, cerca de Cerro Tin
˜
oso, 23 May
1995, 1,200 m, 30TWM52, Montamarta s.n. (VAL 94895);
Fuentelsaz, Cerro de San Juan, en la cima, 2 Jun 1990,
1,350 m, 30TWM43, Montamarta s.n. (VAL 86662);
Deza, Sierra Min
˜
ana, 23 May 1988, 1,100 m,
30TWL8095, M.L. Gil Zu
´
n
˜
iga & J.A. Alejandre 1448/88
(MA 484247); Narros, vertiente norte de la Sierra del
Almuerzo, 5 Jun 1980, 1,280 m, 30TWM6033, Granzow
123 & Zaballos (MA 258222); San Felices, May 1964,
800 m, A. Segura Zubizarreta s.n. (MA 355424); Almarail,
1 Jun 1968, 1,000 m, A. Segura Zubizarreta s.n. (MA
355376); Ciria, en las cercanı
´
as del castillo, 29 May 1988,
1,040–1,100 m, 30TWM8608, M.L. Gil Zu
´
n
˜
iga & J.A.
Alejandre 1153/88 (MA 484243); Madas de Lubia, 25 May
1934, L. Ceballos & C. Vicioso s.n. (MA 21799).
Toledo: Cigarral, junto al Barranco de la Degollada, 24
Mar 1982, 582 m, 30SVK1312, P. Egido 476 (MA 433116).
Zaragoza: Purujosa, S
a
del Moncayo, 11 Jun 1988,
1,362 m, 30TXM042193 (VAL 82451); An
˜
o
´
n, Macizo del
Moncayo, Sierra de la Muela, 17 Jun 1989, 1,500 m,
30TXM0219, M.L. Gil Zu
´
n
˜
iga & J.A. Alejandre s.n. (MA
484224); Sierra de Atea, 30 May 1909, 1,100 m, B. & C.
Vicioso s.n. (MA 21797); Calcena, entre Corrales del Roble
y Corrales de Valdenoria, 11 May 2006, 935 m,
30TXM047114, P.M. Uribe-Echebarrı
´
a s.n. (ABH 51045).
Portugal Algarve: Loule
´
, estrada para o Barranco do
Velho, acima do Olho de Agua, Cabec¸a Gorda, 20 Mar
1979, Malato-Beliz & J.A. Guerra 15174 (MA 285478);
Loule
´
, Amendoeira, Fonte Filipe, 20 Mar 1979, Malato-
Beliz & J.A. Guerra 15206 (MA 285477); Loule
´
, estrada
para o Barranco do Velho, entre o cruzamento e a Amen-
doeira, 21 Mar 1979, Malato-Beliz & J.A. Guerra 15254
(MA 285476); Loule
´
, estrada para o Barranco do Velho,
margem esquerda da Rib
a
das Merce
ˆ
s, 22 Mar 1979,
Malato-Beliz & J.A. Guerra </