taxonomic revision of ornithogalum subg. beryllis (hyacinthaceae)

23
Belg. J. Bot. 142 (2) : 140-162 (2009) © 2009 Royal Botanical Society of Belgium INTRODUCTION Ornithogalum L. is widely distributed in Europe, Asia (reaching Afghanistan to the East), Africa (except the Tropic of Cancer band) and Madagascar (ZAHARIADI 1965). In a traditional sense, it comprises about 120-130 taxa (LAND- STRÖM 1989), or even 200-250 according to OBER- MEYER (1978) and MÜLLER-DOBLIES & MÜLLER- DOBLIES (1996). Based on some oral and reproductive characters, many sections (ENGLER 1888, ZAHARIADI 1965, 1977) and subgenera (BAKER 1872, ZAHARIADI 1965, 1977, 1980, OBERMEYER 1978) have been recognised within Ornithogalum, many of them on the basis of pre- viously described genera (GRAY 1821, RAFIN- ESQUE 1837, PARLATORE 1854, SALISBURY 1866). Other recent studies have suggested very dif- ferent arrangements, almost opposite to one another. On the one hand, SPETA (1998) reduced Ornithogalum to the Eurasian and North African taxa of O. subg. Ornithogalum (= O. subg. Helio- charmos Baker), which was hence constituted by 50-60 species, and he placed the rest of groups as autonomous genera such as Albuca L., Cathissa Salisb., Coilonox Raf., Dipcadi Medik., Eliokar- mos Raf., Honorius S.F. Gray, Loncomelos Raf., Melomphis Raf., Neopatersonia Schönland, Pseu- TAXONOMIC REVISION OF ORNITHOGALUM SUBG. BERYLLIS (HYACINTHACEAE) IN THE IBERIAN PENINSULA AND THE BALEARIC ISLANDS Mario MARTÍNEZ-AZORÍN * , Manuel B. CRESPO and Ana JUAN CIBIO (Instituto de la Biodiversidad), Universidad de Alicante, Apartado 99, E-03080 Alicante, Spain ( * Author for correspondence; e-mail: [email protected]) Received 7 November 2008; accepted 18 June 2009. ABSTRACT. — As part of a taxonomic revision of the Iberian and Balearic taxa of Orni- thogalum, quantitative and qualitative characters are studied in detail in Ornithogalum subg. Beryllis (Salisb.) Baker, and they are evaluated for taxonomy of the two taxa: O. narbonense L. and O. pyrenaicum L. A complete description is presented for both species, and data on their biology, ecology, and distribution are also included. Moreover, a key is provided to facilitate identication. RESUMEN. — En el marco de una revisión de las especies ibéricas y baleares de Ornitho- galum, se estudia con detalle O. subg. Beryllis (Salisb.) Baker, y se evalúa cualitativa y cuanti- tativamente el valor taxonómico de los caracteres morfológicos de los dos táxones: O. narbo- nense L. y O. pyrenaicum L. Para cada especie se presenta una descripción completa y datos sobre su biología, ecología y distribución. Además, se aporta una clave para facilitar la identi- cación de los táxones aceptados. KEY WORDS. — Loncomelos, Beryllis, Ornithogalum narbonense, Ornithogalum pyrenai- cum, Andorra, distribution, Portugal, Spain, taxonomy.

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Page 1: taxonomic revision of ornithogalum subg. beryllis (hyacinthaceae)

Belg. J. Bot. 142 (2) : 140-162 (2009)© 2009 Royal Botanical Society of Belgium

INTRODUCTION

Ornithogalum L. is widely distributed in Europe, Asia (reaching Afghanistan to the East), Africa (except the Tropic of Cancer band) and Madagascar (ZAHARIADI 1965). In a traditional sense, it comprises about 120-130 taxa (LAND-STRÖM 1989), or even 200-250 according to OBER-MEYER (1978) and MÜLLER-DOBLIES & MÜLLER-DOBLIES (1996). Based on some fl oral and reproductive characters, many sections (ENGLER 1888, ZAHARIADI 1965, 1977) and subgenera (BAKER 1872, ZAHARIADI 1965, 1977, 1980, OBERMEYER 1978) have been recognised within

Ornithogalum, many of them on the basis of pre-viously described genera (GRAY 1821, RAFIN-ESQUE 1837, PARLATORE 1854, SALISBURY 1866).

Other recent studies have suggested very dif-ferent arrangements, almost opposite to one another. On the one hand, SPETA (1998) reduced Ornithogalum to the Eurasian and North African taxa of O. subg. Ornithogalum (= O. subg. Helio-charmos Baker), which was hence constituted by 50-60 species, and he placed the rest of groups as autonomous genera such as Albuca L., Cathissa Salisb., Coilonox Raf., Dipcadi Medik., Eliokar-mos Raf., Honorius S.F. Gray, Loncomelos Raf., Melomphis Raf., Neopatersonia Schönland, Pseu-

TAXONOMIC REVISION OF ORNITHOGALUM SUBG. BERYLLIS (HYACINTHACEAE) IN THE IBERIAN PENINSULA

AND THE BALEARIC ISLANDS

Mario MARTÍNEZ-AZORÍN*, Manuel B. CRESPO and Ana JUAN

CIBIO (Instituto de la Biodiversidad), Universidad de Alicante, Apartado 99, E-03080 Alicante, Spain

(* Author for correspondence; e-mail: [email protected])

Received 7 November 2008; accepted 18 June 2009.

ABSTRACT. — As part of a taxonomic revision of the Iberian and Balearic taxa of Orni-thogalum, quantitative and qualitative characters are studied in detail in Ornithogalum subg. Beryllis (Salisb.) Baker, and they are evaluated for taxonomy of the two taxa: O. narbonense L. and O. pyrenaicum L. A complete description is presented for both species, and data on their biology, ecology, and distribution are also included. Moreover, a key is provided to facilitate identifi cation.

RESUMEN. — En el marco de una revisión de las especies ibéricas y baleares de Ornitho-galum, se estudia con detalle O. subg. Beryllis (Salisb.) Baker, y se evalúa cualitativa y cuanti-tativamente el valor taxonómico de los caracteres morfológicos de los dos táxones: O. narbo-nense L. y O. pyrenaicum L. Para cada especie se presenta una descripción completa y datos sobre su biología, ecología y distribución. Además, se aporta una clave para facilitar la identi-fi cación de los táxones aceptados.

KEY WORDS. — Loncomelos, Beryllis, Ornithogalum narbonense, Ornithogalum pyrenai-cum, Andorra, distribution, Portugal, Spain, taxonomy.

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 141

dogaltonia Kuntze, Stellarioides Medik. and Zahariadia Speta. On the other hand, MANNING et al. (2004) merged all those genera into Ornithog-alum, which resulted in over 300 species and was, therefore, equivalent to subfamily Ornithogaloi-deae.

In this study, we accept provisionally the subgeneric rank proposed by BAKER (1872), since it explains better the naturalness of the group based on morphological characters and it was fol-lowed mostly in the European fl oras. However, our preliminary phylogenetic studies in the Orni-thogaloideae, based on sequencing of plastid (trnL-F and rbcL) and nuclear (ITS) regions, together with morphological studies (MARTÍNEZ-AZORÍN et al., unpubl. data), point out the mono-phyly of O. subg. Beryllis, as well as of other related subgenera. Therefore, acceptation of Lon-comelos, in the sense of SPETA (1998), could also be acceptable.

Ornithogalum subg. Beryllis (Salisb.) Baker (= Loncomelos Raf.) includes plants with very long and narrow racemose infl orescences; pedi-cels patent or erect-patent in fl ower and erect and appressed to the stem in fruit; tepals white or yel-lowish on the adaxial face, bearing a central green band on the abaxial face; fi laments linear, lanceo-late or tapering, abruptly expanded in their basal half; ovary oblong or subsphaerical, with long and fi liform style and small stigma; seeds polygo-nal or irregularly compressed, with verruculose or smooth and puzzle-like testa. About 20 species are usually included in that subgenus (WITTMANN 1985, SPETA 1998), which is represented in the Iberian Peninsula and the Balearic Islands by O. narbonense L. and O. pyrenaicum L. Data are presented here for a revision of the subgenus in those territories, as a part of a global revision of the genus that is being carried out (MARTÍNEZ-AZORÍN et al. 2007).

BRIEF HISTORY OF O. SUBG. BERYLLIS

Among the 13 genera in which SALISBURY (1866) split the Linnaean Ornithogalum, he described Beryllis to include O. pyrenaicum and O. narbonense (under O. stachyodes Sol.), and also proposed the new Eustachys Salisb. to

segregate O. latifolium L. and O. pyramidale L. Although the latter is a homonym of Eustachys Desv. (1810) and, therefore, is to be treated as illegitimate, it is usually treated as synonym of Beryllis. However, FABRICIUS (1759) had previ-ously segregated Celsia to include O. latifolium L., O. luteum L. (currently included in Gagea Salisb.), and O. umbellatum L. (a member of O. subg. Ornithogalum). Although Fabricius’s genus, as fi rst described, is quite heterogeneous and confusing, it is a later homonym of Celsia L. (LINNAEUS 1753) and therefore is illegitimate. A similar proposal was later effected by RAFIN-ESQUE (1837), who described Loncomelos to group O. pyrenaicum L., O. narbonense L., O. latifolium L. and O. pyramidale L. (together with other South African species currently assigned to O. subg. Osmyne sensu OBERMEYER 1978), and Loncoxis to segregate O. sulphureum Schult. f. (a species usually synonymised to O. pyrenaicum L.). Therefore, all those genera are to be treated as synonyms of Loncomelos Raf., the name having priority at the genus rank (SPETA 1998).

Nonetheless, BAKER (1872) used Beryllis Salisb. at the subgeneric rank, making the combi-nation O. subg. Beryllis (Salisb.) Baker, whereas ENGLER (1888) regarded it as a mere section of Ornithogalum. Both authors used Beryllis in an expanded sense that included the Mediterranean taxa (Beryllis s.s.) plus several South African taxa currently related to O. subg. Aspasia (Salisb.) Baker and O. subg. Urophyllon (Salisb.) Baker, p.p. (OBERMEYER 1978).

When the name O. subg. Beryllis (Salisb.) Baker was published it was superfl uous nomen-claturally, since the type of Urophyllon Salisb. was included (Art. 52.1, 52.2e; MCNEILL et al. 2006), and consequently it was a synonym of O. subg. Urophyllon (Salisb.) Baker, though it was not illegitimate (Art. 52.3). When both subgenera are treated as different, both names recover their validity and they can be used, each one with its corresponding circumscription. In this way, the new name O. subg. Spetagalum proposed by MÜLLER-DOBLIES & MÜLLER-DOBLIES (1996) to replace the supposedly illegitimate O. subg. Beryllis, is certainly superfl uous, since these

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142 BELGIAN JOURNAL OF BOTANY 142

authors explicitly remarked that this new name should be used exclusively for the Mediterranean plants, maintaining the South African ones in O. subg. Urophyllon (Urophyllon s.s.).

More recently, ZAHARIADI (1965) followed Baker’s treatment and even combined Eustachys Salisb. as a subgenus of Ornithogalum. Later, he recognized the sections Involuta, Galactea and Albedo within O. subg. Beryllis, as well as sec-tion Eustachys in O. subg. Eustachys (ZAHARIADI 1977).

On the basis of molecular, morphological and chemotaxonomical data, PFOSSER & SPETA (1999) and SPETA (1998, 2001) revived Loncome-los Raf., though in a narrower sense which exclu-sively included Euro-Asian and North African taxa. Recently, this solution has been followed by an increasing number of authors (SPETA 2006, RAVENNA 2007, GARBARI et al. 2007, PERUZZI et al. 2007). Conversely, MANNING et al. (2009) accepted it as a synonym of O. sect. Ornitho-galum based on cladistic analyses of chloroplast DNA sequences mostly from sub-Saharan taxa of Ornithogaloideae.

MATERIALS AND METHODS

A total of 409 herbarium vouchers of O. narbon-ense and 164 of O. pyrenaicum were revised from the herbaria ABH, BIO, BC, BCN, COI, FCO, GDA-GDCA, HGI, HUAL, K, LISI, MA, MACB, MGC, P, SALA, SANT, SEV and VAL (acronyms according to HOLMGREN & HOLMGREN 1998).

As fragments on vouchers usually were problem-atic because some important structures were lacking -e.g., bulbs, fruits, or seeds —, or because measuring some others -e.g., dimensions of the fi laments of the sta-mens or ovary- would imply serious damage to mate-rial, morphological studies were undertaken mostly on living material from natural populations, and within a few hours after collection. However, when complete dried herbarium specimens were available, some mor-phological features were complemented with data from vouchers. Consequently, morphological studies here presented are based on a total of 47 complete individu-als selected from 10 populations of O. narbonense, and 19 from 6 populations of O. pyrenaicum (Table 1). For each taxon, only a selection of localities is listed to avoid repetitions and to shorten information.

The studied morphological characters were as follows.

Table 1. Studied specimens of taxa of O. subg. Beryllis used in the morphological studies, and the statistic analyses (*).

Taxon Locality Number of specimens VoucherO. narbonense* ESP., Alicante, Benimantell 10 ABH 47144O. narbonense ESP., Alicante, Sª de Mariola 12 –

O. narbonense ESP., Alicante, Gata — Javea 6 ABH 50143O. narbonense ESP., Castellón, Tirig 2 MA 540009O. narbonense ESP., Huelva, Almonaster 2 ABH 50130O. narbonense ESP., Teruel, Torre de Arcas 2 VAL 78567O. narbonense ESP., Valencia, Algar de Valencia 2 VAL 7839O. narbonense ESP., Zaragoza, Peñafl or 2 VAL 99587O. narbonense ESP., Cádiz, El Bosque 4 ABH 47139O. narbonense ESP., Leon, Carbajal de la Legua 5 ABH 51028O. pyrenaicum* ESP., Huelva, Puebla de Guzmán 9 ABH 50135O. pyrenaicum ESP., Palencia, Velilla del Rio Carrión 3 VAL 104621O. pyrenaicum ESP., Lérida, Espot 1 VAL 142592O. pyrenaicum ESP., Soria, Villar del Ala 4 VAL 99185O. pyrenaicum ESP., Pontevedra, Vila de Cruces 1 SANT 41677O. pyrenaicum ESP., Málaga, Villanueva del Rosario 1 BC 61929

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 143

BULBS

Length and diameter of the bulb. Length was consid-ered as the vertical distance from bulb base to its apex, including neck. Diameter was expressed as the maxi-mum width of the bulb.

Bulb scales structure. After the transversal cut of the bulb, the level of fusion or concrescence of scales, as well as their arrangement was noted.

LEAVES

Leaf length and width. Maximum length and width of leaves were measured in each specimen.

Total number of leaves. The total number of basal sheaths was determined in each specimen; fragmented or dried leaves were included in the counts.

STEMS AND INFLORESCENCES

Stem length. It corresponded to the stem length from the apex of the bulb up to the fi rst fl ower (base of the infl orescence).

Infl orescence length and width. The distance from the point where the lowermost fl oral pedicel inserts to the apex of the completely developed infl orescence was considered as the infl orescence length, whilst the maxi-mum horizontal distance between the apexes of the longest pedicels (without considering tepals) was regarded as its width.

Plant height. It corresponded to the total length of the stem plus the total length of the infl orescence.

Number of fl owers. The total number of fl owers was counted, including those at the tip of the immature infl orescence or prior to anthesis.

Flower and fruit pedicel length. Three kinds of fl owers were distinguished in each infl orescence: lowermost, middle and uppermost. The middle fl ower was that whose pedicel was on the nearest point to the middle of the infl orescence. Fruit pedicels were measured after capsule ripening.

Flower and fruit pedicel insertion angle. Average angle of insertion of the pedicels on the infl orescence axis was measured separately for fl ower and fruit pedicels.

Bract maximum length and width. The maximum length and width of the lowermost bract in the infl orescence were measured.

FLOWERS

Flower diameter. Length between the apexes of two opposite tepals was measured on the lowermost fl ower of the infl orescence.

Tepal length and width. Both characters were measured on the lowermost fl ower of the infl orescence, differen-tiating between outer and inner tepals.

Tepal green band width. A longitudinal green band is present on the abaxial face of the tepals; the width of this band was measured separately for outer and inner tepals.

Length and width of stamen fi laments, anthers and ovary at anthesis. Maximum length and width of sta-men fi laments, anthers and ovary were measured.

Style length and stigma morphology. The shape and glandulousness of the stigma were described.

FRUITS

Capsule length and width. Maximum length and width of mature capsules were measured.

Number of seeds per capsule. Seeds were counted in 32 to 35 randomly selected capsules from different indi-viduals and/or populations for each species. In some cases, individuals collected in the wild were transferred into pots until capsules ripened. Then, they were cut off and isolated until they opened spontaneously.

SEEDS

Seed length and width. Maximum length and width of seeds were measured. Measurements were made on 40 seeds per population, in different populations when possible. A binocular Leica® MZ6, with incorporated micrometer was used.

Seed weight. Each measured seed was also weighted with an analytical balance HM-202 (AND®).

Measurements of width of tepals and green band of tepals, length and width of fi laments, anthers, ova-ries, capsule and seeds, as well as length of style, were made on a ruler with a precision of 0.1 mm. Moreover, weight of seed was measured with a precision of 0.1 mg. Other measurements such as length and diameter of bulb, length of neck of bulb, width of leaves and infl orescence, length and width of bracts, length of fl o-ral and fruiting pedicels, and length of tepals, were made with a precision of 1 mm. Only length of leaves, stem and infl orescence were measured with a precision of 1 cm.

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144 BELGIAN JOURNAL OF BOTANY 142

SCANNING ELECTRON MICROSCOPE

Images of seeds and pollen grains were taken with a scanning electron microscope (SEM) JEOL 840. As the material was dried, no special treatment was required prior to observation. Samples were directly glued on metallic stubs, and then coated with about 30 nm gold. Seed testa classifi cation follows MORET et al. (1990).

SCANNING OF FRESH PLANTS

After the morphological study of specimens col-lected in the wild, scannings were carried out of every living plant and their different vegetative and fl oral structures. For this purpose, an ‘Epson Perfection 1250’ scanner was used. General plates of both species were composed with those images.

STATISTIC ANALYSES

All studied Iberian populations of both O. nar-bonense and O. pyrenaicum were very homogeneous in their principal morphological features. Therefore, just one population representative of each species (Table 1) was selected for statistic analyses, in which measurements were taken from living individuals. For O. narbonense, 10 specimens from Alicante province were studied while for O. pyrenaicum, 9 plants from Huelva province were studied (Table 1). With regard to the morphological variables, a princi-pal component analysis (PCA) was conducted with SPSS ver. 10.

RESULTS

BULBS

Both O. narbonense and O. pyrenaicum pro-duce bulbs quite similar in shape, being both ovate with a more or less developed and narrow neck at their tip. Length and width show slight overlapping in both taxa (Table 2), though the former species reaches commonly larger values (Table 2). However, bulbs are longer in O. narbo-nense, because of the longer neck (16)20-34(40) mm, which is fragile and sometimes curved. Both species usually do not produce secondary bulbils, though exceptionally they can be found in very low number (1-3). Moreover, the bulb scales of both taxa are free and imbricate.

LEAVES

Both O. narbonense and O. pyrenaicum pro-duce very long and narrowly lanceolate or taper-ing, slightly glaucous leaves. However, in O. nar-bonense they are usually a bit shorter and wider than in O. pyrenaicum (Table 2). The number of leaves is similar in both species (Table 2). Moreo-ver, leaves of O. pyrenaicum usually almost com-pletely wither during early anthesis, while those of O. narbonense remain at least partially green until fruiting.

INFLORESCENCE

Taxa of O. subg. Beryllis produce a typical very long racemose infl orescence. Length and width of the infl orescences in both taxa are simi-lar, though O. pyrenaicum usually presents nar-rower infl orescences, with fewer fl owers and longer stems than O. narbonense (Table 2). In both species, the length of fl ower pedicels is approximately equal along most of the infl ores-cence, but shorter at the apex, which results in a conical ending. However, the lowermost and mid-dle pedicels can be longer in O. narbonense than in O. pyrenaicum (Table 2). The length of fruit pedicels are somewhat different in both taxa, although there is a wide overlap (Table 2). O. nar-bonense usually produces much longer pedicels, up to 40-50 mm long, but sometimes they are no longer than 15 mm. In O. pyrenaicum, fruit pedi-cels are commonly shorter, up to 29 mm. Finally, bracts are constantly longer and wider in O. nar-bonense (Table 2).

PERIGON

Inner and outer tepals showed a similar size in both species, though inners can be slightly shorter and wider than the outers. Tepals of O. narbonense are lanceolate or ovate-lanceolate, white-coloured on the adaxial face and white with a longitudinal green band on the abaxial face, rarely slightly curled at margins. On the contrary, in O. pyrenaicum they are narrowly lanceolate to almost linear, and typically yellowish on the adax-ial face and yellowish with a longitudinal green band on the abaxial face, usually with convolute

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 145

Table 2. Observed values and ranges of the studied morphological characters.

Morphological character O. narbonense O. pyrenaicumTotal bulb length (mm) (36)50-70(100) (32)34-50(58)Neck length (mm) (16)20-34(40) 10-20Bulb diameter (mm) (25)30-50(55) (22)26-36(38)Leaf number (3)5-8(12) (3)4-7(8)Leaf length (cm) (22)31-62(67) (38)40-75(80)Leaf width (cm) (0.3)0.5-1.6(1.8) (0.6)0.7-0.9(1.3)Stem length (cm) (11)25-57(61) (46)48-78(85)Number of fl owers (12)19-64(79) (20)25-55(64)Infl orescence length (cm) (12)15-48(50) (16)20-45(51)Infl orescence width (cm) (1)1.7-3.7(4.8) (0.7)1.5-2.6(3)Bract length (mm) (10)12-24(26) (7)8-15(19)Bract width (mm) (2)3-5(6) (2)2.5-3.7(4)Lowermost fl oral pedicel length (mm)

(12)15-38(48) (9)13-25(29)

Middle fl oral pedicel length (mm) (9)12-26(28) (8)11-17(19)Uppermost fl oral pedicel length (mm)

(2)3-16(20) (2)3-7(9)

Fruiting pedicels (mm) 15-40(50) 18-25(29)Outer tepal length (mm) (12)13-15(16) 7-11(12)Inner tepal length (mm) (11)12-14(15) 7-10(11)Outer tepal width (mm) (2.5)3-4(4.5) (1.3)2-2.3Inner tepal width (mm) (2.5)3-4.5(5) (1.5)2-2.5Tepal green band width (mm) (1)1.2-1.8(2) 1-1.2Outer fi lament length (mm) (5)6-7.5 5-6Inner fi lament length (mm) (6)6.5-7.5(8) 5.5-6.2Outer fi lament width (mm) 1.1-1.5(1.7) 1.2-1.4Inner fi lament width (mm) 1.2-2 (1.2)1.4-1.8Anther length (mm) (1.7)2-3(4) 1.5-2(3)Anther width (mm) (0.7)0.8-1.5(1.7) 0.7-0.8(1.2)Ovary length (mm) (2.5)3-4(5) (1.5)2-2.5(2.8)Ovary width (mm) (2)2.5-3(3.2) (1.2)1.5-2.5(2.7)Style length (mm) (2)3-4(5) (2)2.5-3.5(4)Capsule length (mm) (9)10-15(16) (7)8-9(9.5)Capsule width (mm) (4.5)5-6.5(7) (5)5.5-6.5(7)Number of seeds per capsule (6)8-15(17) (6)10-18(20)Average number of seeds per capsule

10 ± 2.83 12.25 ± 4.70

Seed length (mm) (2.3)2.6-3.8(4) (2.3)2.5-3.1(3.3)Seed width (mm) (1.6)1.8-2.4(2.6) (1.4)1.6-2(2.3)Seed weight (mg) (3.5)4-5.5(6) 5-7.5

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146 BELGIAN JOURNAL OF BOTANY 142

margins after anthesis. Tepals are commonly longer and wider in O. narbonense than in O. pyrenaicum (Table 2). Moreover, the dorsal green band of tepals is usually narrower in O. pyrenai-cum than in O. narbonense.

ANDROECIUM

Stamen fi laments of both whorls are similar in shape in both taxa, being abruptly expanded in their lower half and much narrower and fi li-form at the apex, though inners are slightly longer and wider than outers (Table 2). They are usually longer and a bit wider in O. narbonense than in O. pyrenaicum (Table 2). Anthers are equal in size in both whorls, though O. narbon-ense show slightly longer and wider anthers -(1.7)2-3(4) × (0.7)0.8-1.5(1.7) mm- than O. pyrenaicum -1.5-2(3) × 0.7-0.8(1.2) mm-. Pollen grains of O. narbonense and O. pyrenaicum are elliptical, anacolpate, with reticulate tectum, ret-icules being wider and more abundant in the equatorial zone (Fig. 1).

GYNOECIUM

In Ornithogalum narbonense the ovary is ovoid-lanceolate to cylindrical, whereas O. pyrenaicum shows an ovoid, spherical or shortly cylindrical ovary. Moreover, the ovary is wider and longer in O. narbonense than in O. pyrenai-cum (Table 2). The style is long and fi liform in both taxa, showing a wide overlap, though it can be longer in O. narbonense (Table 2). Finally, the stigma is usually small, punctiform and trigonous in both species.

FRUIT

Capsules typically stand appressed to the stem in Ornithogalum subg. Beryllis. The capsule of O. narbonense is elliptical to slightly ovoid, while in O. pyrenaicum it is ovate-cylindrical. Capsule width is similar in both species, though O. narbonense usually produces longer capsules than O. pyrenaicum (Table 2). Moreover, capsules of O. pyrenaicum usually contain more seeds (12 ± 4.7) than those of O. narbonense (10 ± 2.8).

Fig. 1. Pollen grains in Ornithogalum subg. Beryllis. a: O. narbonense (Alicante, Spain); b: O. pyrenaicum (Huelva, Spain). Scale = 10 mm.

SEEDS

Seeds of O. narbonense and O. pyrenaicum are blackish and dull-coloured, more or less polyg-onal or irregularly compressed, somewhat fl at-tened and angulose with more or less winged edges, and with irregularly rugose surface (Fig. 2).

Seeds are usually longer and wider in O. narbon-ense than in O. pyrenaicum, though seed weight is higher in O. pyrenaicum (Table 2). Seed testa is papillate or granulate (type 2) in O. narbonense, whereas in O. pyrenaicum it is rugose to minutely reticulate, sometimes with scattered small grains on edges (Fig. 2).

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 147

PRINCIPAL COMPONENT ANALYSES

In the PCA of all studied morphological char-acters in the two selected populations (Fig. 3), O. narbonense and O. pyrenaicum appear clearly sep-arated. Ten principal components have been obtained, which together explain 91.7% of the vari-ance, with the fi rst two components explaining 47% of the variance (32.2% and 14.8%, respectively). The fi rst one allows to separate perfectly O. narbo-nense from O. pyrenaicum and mainly refl ects some characters, such as stem length, tepal length and width, inner fi lament length or capsule length (Table 3). The second principal component was not effec-tive to distinguish the two species. The percentage of explained variance and correlations with mor-phological variables were quite low. When all stud-ied Iberian populations were analysed together, PCA yielded similar results (data not shown).

DISCUSSION

Many of the studied morphological charac-ters are not useful for an unequivocal identifi ca-tion of O. narbonense and O. pyrenaicum. Both taxa show a considerable morphological plastic-ity, which generates a wide overlap in many of their vegetative and reproductive characters.

With regard to bulbs, both species show rather similar features. However, those in O. narbonense bear a fragile and longer neck, which prevents easy extraction, and, therefore, could favour this species in disturbed or cultivated areas. This character is not present in O. pyrenaicum, a species that grows in meadows and grassland in mountain areas. Other features such as shape and size of leaves, infl orescences, androecium, and gynoecium, are not absolutely decisive for distinguishing both

Fig. 2. Seeds of the species of Ornithogalum subg. Beryllis. a-b: O. narbonense (Alicante, Spain); c-d: O. pyrenaicum (Huelva, Spain). Scales: a, c = 1 mm; b, d = 100 mm.

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148 BELGIAN JOURNAL OF BOTANY 142

species. Nonetheless, they are informative enough when combined with other diagnostic characters.

Among the studied characters, several veg-etative and reproductive features were revealed as fully diagnostic for identifi cation of O. narbo-nense and O. pyrenaicum by the PCA (Table 3; Fig. 3). Many authors (e.g., ZAHARIADI 1980, PIGNATTI 1982, TORNADORE 1985, 1986, PASTOR 1987, DE BOLÒS & VIGO 2001, GARBARI et al. 2007) had already emphasized leaf withering at anthesis, and colour, size and shape of tepals as crucial diagnostic characters. However, the stud-ied Iberian plants of O. pyrenaicum show smaller tepals (Table 2), which are similar to those from North African plants (8-10 mm; MAIRE 1958). They are slightly shorter and narrower than those from other European countries such as Britain (11-13 mm; STACE 1997, HILL & PRICE 2000), France and Switzerland (10-12 mm; COSTE 1906) or Italy (11-12 mm; TORNADORE 1985, GARBARI et al. 2007). Therefore, the overlapping with O. narbonense is higher in those populations. Con-versely, the Iberian populations of O. narbonense

show tepals up to 15-16 mm long, which is longer than the Italian plants (TORNADORE 1985, GAR-BARI et al. 2007). It is interesting to note here that the lectotype of O. pyrenaicum (LINN 428.5) shows tepals 8-10 mm long, hence falling into the variation rank of the Iberian and North Afri-can plants. According to that, the Linnean type more likely comes from the Pyrenees than from the Alps.

TORNADORE (1985, 1986) and GARBARI et al. (2007) reported clear differences in fruit pedicel length, being longer in O. narbonense (2-4 cm) than in O. pyrenaicum (up to 2 cm). However, our data indicate that the Iberian plants of the latter species can reach 29 mm, clearly exceeding meas-urements of the Italian ones.

Our results also point out that length of bulb neck, tepal green band width, stamen fi laments length, anther size, ovary size, and capsule length, are also variables contributing with high percent-ages of variance to the fi rst principal component (Table 3; Fig. 3). Therefore, they are also rather useful for distinguishing both species.

Fig. 3. PCA of all studied morphological characters.

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 149

Anatomical features of leaves have been used to distinguish among groups of taxa in O. subg. Ornithogalum, using O. narbonense [≡ Loncomelos narbonensis (L.) Raf.] as the out-group (PERUZZI et al. 2007). However, they were not suffi cient for taxa differentiation. Con-versely, GARBARI et al. (2007) found that leaves of Italian populations of O. narbonense pro-

duced about 41-55 vascular bundles, whilst those of O. pyrenaicum only 24-44. This differ-ence, which is directly related to leaf width, was used by the same authors in their identifi cation key. However, the taxonomical utility of leaf anatomy has not been tested here for O. subg. Beryllis, and should be the focus of further research.

Table 3. Studied morphological characters and principal components obtained in the analysis.

Character Component 1 Component 2Bulb length 0.570 -0.343Bulb diameter 0.531 -0.553Leaf number -0.506 0.388Leaf length -0.671 0.244Leaf width 0.661 0.359Stem length -0.790 0.212Number of fl owers 0.151 0.449Infl orescence length -0.101 0.547Infl orescence width 0.495 0.411Bract length 0.506 -0.015Bract width 0.111 0.731Lowermost fl oral pedicel length 0.163 0.310Middle fl oral pedicel length 0.381 0.249Uppermost fl oral pedicel length 0.193 -0.464Outer tepal length 0.853 0.085Inner tepal length 0.820 0.223Outer tepal width 0.948 0.117Inner tepal width 0.903 0.221Outer tepal green band width 0.664 0.487Inner tepal green band width 0.593 0.341Outer fi lament length 0.484 0.434Inner fi lament length 0.811 0.268Outer fi lament width 0.059 0.761Inner fi lament width 0.507 0.459Anther length 0.725 -0.441Anther width 0.701 -0.276Ovary length 0.520 0.651Ovary width -0.093 0.662Style length 0.296 0.228Capsule length 0.712 -0.313Capsule width -0.043 -0.015Seed length 0.595 -0.265Seed width 0.493 -0.068Seed weight -0.309 -0.230

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150 BELGIAN JOURNAL OF BOTANY 142

GARBARI et al. (2007) reported some minor differences pollen grain size between O. narbon-ense and O. pyrenaicum, with the former showing slightly larger pollen grains (64-79 × 31-33 mm) than the latter (58-66 × 25-29 mm). Our results (data not shown) fall within the variation range presented by those authors. Moreover, pollen microsculpture offers some remarkable differ-ences between both taxa. As reported by DÍEZ & PASTOR (1985), the exine surface of Ornithoga-lum taxa varies from irregularly reticulate to per-forate near both the apertures and the polar areas, with clearly wider and more abundant reticules in the equatorial zone. However, after our results such reticules are wider in O. pyrenaicum result-ing in an irregular pattern of perforations, whilst they are much narrower in O. narbonense (Fig. 1), showing a rather more regular pattern.

According to GARBARI et al. (2007), seed size is similar in both taxa, around 3.6-5.0 × 2.3-2.5 mm in O. narbonense, and 4.0-5.0 × 2.5 mm in O. pyrenaicum. Remarkably, the Iberian plants produce smaller seeds in both cases: they are 2.3-4 × 1.6-2.6 mm in O. narbonense and 2.3-3.3 × 1.4-2.3 mm in O. pyrenaicum (Table 2). How-ever, similar smaller seed size has been reported by BEDNORZ & CZARNA (2008) for Polish plants of the latter, suggesting that seed size is a variable character. Three types of seeds were described by MORET et al. (1990), on the basis of the ornamen-tation pattern of testa cells. Although those types were thought to characterize different subgenera of Ornithogalum (ZAHARIADI 1977), a direct rela-tionship has not been demonstrated. In fact, COSKUNCELEBI et al. (2000) and MORET et al. (1990) found more than one seed type in various Eurasian sections of the genus. Within O. subg. Beryllis, Type 2 (granulate) and Type 3 (ruminate or puzzle-like) have been identifi ed by MORET et al. (1990), who ascribed both O. narbonense and O. pyrenaicum to Type 2. As shown in Fig. 2, the studied Iberian plants of O. narbonense produce seeds with a granulate testa (clearly referable to Type 2), whilst in O. pyrenaicum they are irregu-larly rugose to almost minutely reticulate on the periclinal walls, sometimes with smaller granules scattered on the seed edges (but not clearly refer-able to any of the three described types). This

peculiar pattern has been also observed in plants of O. pyrenaicum from England (pers. observ.), and has also been recently reported by BEDNORZ & CZARNA (2008) from plants cultivated in Poland. As it does not correspond to the pattern described by MORET et al. (1990) for plants from the Middle Atlas (Morocco), this is hence a ques-tion to be addressed in the future to clarify the eventual geographical variation patterns of testa ornamentation in O. pyrenaicum.

Combination of all morphological characters yielded complete separation of both taxa, as shown in the PCA analysis (Fig. 3), which indi-cates that the input of every single character is important for a full identifi cation.

TAXONOMIC TREATMENT

KEY FOR THE IBERIAN-BALEARIC TAXA OF ORNITHOGALUM SUBG. BERYLLIS

1. Leaves almost completely withered at anthesis. Tepals yellowish on the adaxial face, 7-11(12) × (1.3)2-2.3(2.5) mm. Testa of the seed rugose to minutely reticulate, sometimes with scattered small grains on edges ............... O. pyrenaicum

– Leaves at least partially green at anthesis. Tepals white on the adaxial face, (11)12-15(16) × (2.5)3-4.5(5) mm. Testa of the seed granulate ................................ O. narbonense

DESCRIPTION OF TAXA

Ornithogalum subg. Beryllis (Salisb.) Baker in J. Linn. Soc. (Bot.) 13 (68): 260. 1872 ≡ Beryllis Salisb., Gen. Pl. [Salisbury]: 33. 1866 ≡ O. subg. Spetagalum U. Müll.-Doblies & D. Müll.-Do-blies in Feddes Repert. 107: 520. 1996 [nom. superfl .] – Lectotype: O. pyrenaicum L. (STEARN 1983)

= Celsia Heist. ex Fabr., Enum. (ed. 2) [Fabr.]: 22. 1763, pro parte [O. latifolium L.], nom. illeg. [non Celsia L., Sp. Pl.: 621. 1753; Scro-phulariaceae]

≡ Loncomelos Raf., Fl. Tellur. 2: 24. 1837 – Lec-totype: L. pyrenaicus (L.) L.D. Hrouda ex J. Holub (cf. SPETA 2001).

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 151

= Loncoxis Raf., Fl. Tellur. 3: 58. 1837 – Holo-type: L. sulfurea (Waldst. & Kit.) Raf. ≡ Anth-ericum sulfureum Waldst. & Kit. [Descr. Icon. Pl. Hung. 1: 98. 1802] ≡ Ornithogalum sul-fureum (Waldst. & Kit.) Schult. & Schult. f. [Syst. Veg. ed. 15 bis [Roemer & Schultes] 7(1): 518. 1829] = O. pyrenaicum L. (cf. WITTMANN 1985).

= Eustachys Salisb., Gen. Pl. [Salisbury]: 33. 1866, nom. illeg. [non Eustachys Desv. in Nouv. Bull. Soc. Philom. 2: 188. 1810; Poaceae] ≡ O. subg. Eustachys Zahar. in Rev. Roumaine Biol., Sér. Bot., 10(4): 281. 1965 – Lectotype (here indicated): O. latifolium L., Sp. Pl.: 307. 1753. Observations: ZAHARIADI (1965) selected O. arcuatum Stev. as the lec-totype. However, as it was not mentioned by Salisbury in the protologue of Eustachys, that lectotypifi cation is to be superseded.

= O. sect. Involuta Zahar. in Rev. Roumaine Biol., Sér. Bot., 10(4): 277. 1965 – Holotype: O. fl avescens Lam.

= O. sect. Galactea Zahar. in Rev. Roumaine Biol., Sér. Bot., 10(4): 277. 1965 – Holotype: O. ponticum Zahar.

= O. sect. Albedo Zahar. in Rev. Roumaine Biol., Sér. Bot., 10(4): 277. 1965 – Holotype: O. fi scherianum Krascheninn.

1. Ornithogalum narbonense L., Cent. Pl. 2: 15. 1756

≡ Loncomelos narbonensis (L.) Raf., Autik. Bot.: 56. 1840 ≡ O. pyramidale subsp. narbo-nense (L.) Asch. & Graebn., Syn. Mitteleur. Fl. 3: 255. 1905

Ind. loc. “Habitat in Galliae australis, Italiae agris”.

Lectotype. LINN 428.7 [designated by STEARN 1983].

Illustrations. PASTOR, Fl. Vasc. Andalucía Occid. 3: 436. 1987; WITTMANN in Stapfi a 13: 38, 39. 1985; Fig. 4.

Description. Geophyte. Bulb (36)50-70(100) × (25)30-50(55) mm, ovoid or sphaerical, generat-ing contractile roots, usually without secondary

bulbils, tapering into a long and usually curved neck (16)20-34(40) mm; outer tunics pale brown or whitish. Leaves (3)5-8(12), in a basal rosette, (22)31-62(67) × (0.3)0.5-1.6(1,8) cm, linear-ta-pering, deeply keeled, green or somewhat glau-cous, glabrous, erect, withering at the apex, synanthous. Floral stem (excluding the infl ores-cence) (11)25-57(61) × 0.3-0.5 cm, erect, smooth, glabrous and pruinose. Infl orescence racemose, very long and pyramidal at the apex, (12)15-48(50) × (1)1.7-3.7(4.8) cm, (excluding fl owers but not their pedicels), with (12)19-64(79) fl ow-ers; bracts (10)12-24(26) × (2)3-5(6) mm, shorter or longer than pedicels, usually half of their length, ovate-lanceolate or triangular, much wider at the base, with apex setose-acuminate, membranose, with 3 greenish nerves; fl oral pedi-cels erect-patent, the lower ones (12)15-38(48) mm, the middle ones (9)12-26(28) mm and the upper ones (2)3-16(20) mm; fruiting pedicels 15-40(50) mm, erect and appressed to the stem. Flowers 20-30 mm in diameter, slightly fragrant; tepals white in the adaxial side and white with a central green band in the abaxial side of (1)1.2-1.8(2) mm width, lanceolate-obovate, obtuse or acute, with smooth or undulate edges; outers (12)13-15(16) × (2.5)3-4(4.5) mm; inners (11)12-14(15) × (2.5)3-4.5(5) mm, being a little shorter and wider than the outers. Stamens 6, 1/2 to 2/3 of the tepal length; fi laments white, abruptly wid-ened in the basal half, outers (5)6-7.5 × 1.1-1.5(1.7) mm, inners (6)6.5-7.5(8) × 1.2-2 mm, being somewhat longer and wider than the out-ers; anthers dorsifi xed, pale yellow or whitish, 3-4 × 1-1.7 mm before dehiscence and 1.5-2 × 0.7-1.5 mm after dehiscence. Ovary (2.5)3-4(5) × (2)2.5-3(3.2) mm, pale green, ovoid-lanceolate to cylindrical, truncate at the apex, trigonous with 3 obtuse ribs and with septal nectaries; style whitish, fi liform, (2)3-4(5) mm; stigma slightly trigonous and glandulose. Capsule (9)10-15(16) × (4.5)5-6.5(7) mm, elliptical to slightly ovoid, trigonous, pale brown, with trivalvar dehiscence. Seeds (6)8-15(17) per fruit (n = 35, mean = 10, SD = 2.83), with (3.5)4-5.5(6) mg weight, (2.3)2.6-3.8(4) × (1.6)1.8-2.4(2.6) mm, angulose and irregularly compressed, and testa granulate (type 2).

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152 BELGIAN JOURNAL OF BOTANY 142

Fig. 4. Ornithogalum narbonense (El Bosque, Cádiz, Spain): a: fl ower, frontal view; b: fl ower, dorsal view; c: infl o-rescence; d: fruiting raceme; e: bract; f: inner (left) and outer (right) stamens; g: gynoecium; h: capsule, lateral view; i: capsule, transversal section; j: capsule after dehiscence, lateral view; k: capsule after dehiscence, apical view; l: bulb; m: leaves. Scale bars = 1 cm.

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ORNITHOGALUM SUBG. BERYLLIS IN THE IBERIAN PENINSULA 153

Chromosome number. 2n = 54 (NEVES 1952, 1973, LÖVE & KJELLQVIST 1973, PALOMEQUE & RUÍZ REJÓN 1976, RUÍZ REJÓN 1978, WITTMANN 1985, TORNADORE 1986, PASTOR & DIOSDADO 1994). 2n = 54 + (O-11B) (NEVES 1952, DARLING-TON & WYLIE 1955). According to TORNADORE (1985), O. narbonense is an autohexaploid taxon with basic chromosome number x = 9.

Biology. It fl owers from April to May or even June. Fruiting extends from May to July or August. Reproduction is mainly by seeds, which are produced in a high number per infl orescence. However, it can also produce a low number of secondary basal bulbils. Bulb shows a narrow neck usually curved at the apex, a characteristic that makes extraction diffi cult by easy breaking of the neck. Moreover, the bulb usually is located very deep in the ground. All these facts can be understood as an adaptation to anthropic activities in crops, where it is found as a weed.

Ecology. This species grows usually on roadsides, cereal crops, grasses and anthropic areas, on soils with basic or acidic nature.

Distribution. Coastal or slightly continental areas of southern Europe, northern Africa, southwestern Asia, and Macaronesia (cf. WITTMANN 1985). In the Iberian Peninsula and the Balearic Islands it is a very common species which inhabits most of the territory, excepting humid areas in the very north and northeast of the Iberian Peninsula (Fig. 5).

Observations. This species could be confused with O. pyrenaicum, mainly when only incom-plete herbarium materials are studied. However, the size and colour of tepals, and testa microstruc-ture clearly separate both taxa. IUCN (2001) cat-egory: LC.

Selected material. SPAIN. Álava. Santacruz de Kamp-ezo, 30TWN5226, 600 m, 13-VI-1985, J.A. Alejandre

Fig. 5. Map of studied populations of O. narbonense in the Iberian Peninsula and the Balearic Islands. Dots corre-spond to both studied natural populations and revised herbarium sheets.

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154 BELGIAN JOURNAL OF BOTANY 142

(MA 338379). Albacete. Sierra de Alcaraz, 11 km al SE de Alcaraz, 38º35’N 2º25’W, 1300 m, 24-VI-1979 (VAL 135663); Alrededores de Sta. Elena de Ruidera, 24-V-1933, Gz. Albo (MA 89011); Alborea, prox. cruce N-332 con ctra. a Casa de Ves, 30SXJ4252, 720 m, 21-V-2006, A. Juan (ABH 51027). Alicante. Villena, 30SXH8674, 500 m, 18-V-1994, M.A. Alonso, De la Torre & Vicedo (ABH 9912); Jávea, carretera de Gata a Jávea, 31SBC5096, 75 m, 08-V-2004, M. Martínez Azorín (ABH 50143); Orihuela, Embalse de la Pedrera, 30SXH8810, 100 m, 12-IV-2000, A. Juan, M. Fabregat & A. Ruiz de León (ABH 43130); Polop, proximidades de la Cova Polida, 27-V-1956, A. Rigual (MA 372393, VAL 138686). Almería. Los Gallardos, 01-V-1971, R. Regredo (VAL 135664); Sierra Cabrera, WG9606, 600 m, 31-V-1995, A. de la Torre (MUB 44594). Barcelona. Begas (Begues), IV-1871, M. Compañó (BC61916). Burgos. Frías, 30TVN7535, 520 m, 02-VI-1990, I. García Mijangos (BIO 16955, FCO 21356). Cáceres. Garrovillas, 09-V-1986, M. Ladero & Santos (SALA 75144); Moraleja, 29TPE8741, 04-VI-1983, A. Valdés Franzi (SALA 68249); Serrejón, 30STK6011, 26-V-1984, T. Ruiz Téllez (SALA 71792). Cádiz. El Bosque, puerta del camping, 30STF7772, 350 m, 04-V-2003, M. Mart. Azorín (ABH 47139); Sierra de las Cabras, 400-500 m, 14-V-1979, M.J. Díez & S. Silvestre (SEV 120447); Jerez de la Frontera, Laguna de Medina, 24-IV-1979, B. Cabezudo, F. García & J. Rivera (SEV 119873); Algodonales, Sierra de Líjar, 500-600 m, 31-V-1979, A. Aparicio, B. Cabezudo & J. Rivera (SEV 58786). Cantabria. Reinosa, Mata-porquera, 13-VI-1952, E. Guinea (MA 164762); Poza-zal, 22-VI-1985, Herrá (MA 682409). Castellón. Zorita del Maestrazgo, Mas de Rigores, río Bergantes, 30TYL3916, 31-V-2004, A. Juan & M.A. Alonso (ABH 50137); Tirig, 31TBE5178, 450 m, 23-V-1992, C. Fabregat & S. López Udías (MA 540009, VAL 89846). Ciudad Real. Villanueva de los Infantes, 30-V-1980, T. Luque, J.L. Ubera & B. Valdés 1149/80 (SEV 120448). Córdoba. Cabra, 03-V-1918, C. Vicioso (BC 61914, MA 21933); Sierra Morena, próximo Pista Sur, UH881195, 400 m, 01-V-1992, M. Melendo (GDAC 41924); Priego de Córdoba, faldas de la Tiñosa, 19-V-1978, J.M. Muñoz (SEV 120130). Cuenca. Fuentes, Laguna del ojo de la Corva, 30SWK8224, 1040 m, 04-VI-1994, Moreno Valdeolivas (VAL 84494); Gara-balla, Los Chicoteros, 1150 m, 01-VI-1978, G. Mateo (VAL 142598); Alrededores de Solán de Cabras, 11-VI-1942, A. Caballero (MA 21947); Cumbres de Cañiza-res, 10-VII-1932, A. Caballero (MA 21950). Gerona. Lladó, 200 m, VI-1895, S. Vayreda (MA 21957); Besalú, La Garrotxa, DG77, 150 m, 21-V-1987, X. Viñas

(HGI 15696); Cantallops, a Querafumat, Alt Empordà, DG9393, 180 m, 04-VI-1998, J. Font (HGI 14388). Formentera. Ses Salines, Can Miquel de Baix, 31SCC648877, 5 m, 16-V-2009, F. Martínez Flores & C. Pena (ABH 53929). Granada. Dílar, Muralla, 30SUF4921012 [incorrect, probably 30SVG4901], 24-V-1989, A. Arregui, C. Pérez & A. del Río (GDAC 43707); Illora, Sierra Parapanda, 30SUG1725 [incor-rect, probably 30SVG1725], 850 m, 21-V-1989, Mª M. Pérez & D. Molina (GDAC 43690); Granada. Frente al cementerio, 07-V-1988, P. Jiménez Tejada (GDAC 33032); Salida de Salar, hacia Alhama, 08-VI-1989, M. Trigo & J.M. Nieto (MGC 25398). Guadalajara. Entre Bujalaro y Mandayona, 04-VI-1970, F. Bellot, R. Carballal & M.E. Ron (SALA 6168); Santa María del Espino, camino hacia el barranco de la Hoz, 30TWL5737, 1160 m, 08-VI-2001, E. Carrillo, L.M. Ferrero & O. Montouto (MA 704368); Padilla de Hita, 04-VI-1970, Bellot, Carballal & Ron (MA 195871). Huelva. Almonaster, Era de la Cuesta Real, 29SPB9593, 23-V-2004, M. Martínez Azorín, M.B. Crespo & C. Pena (ABH 50130); Aroche, salida hacia El Mustio, 29SPC7901, 400 m, 22-IV-1994, E. Rico, F. Amich et al. (MA 717247); Paymogo, 20-V-1942, C. Vicioso (MA 21934). Huesca. Sariñena, cercanías de la estación del ferrocarril, 30TYM3635, 320 m, 02-VI-1979, P. Monserrat (MA 258882, SALA 23633); Barbastro, 4-5 km dirección Monzón, 31TBG65, 400 m, 26-V-1989, J. Pedrol (MA 487386). Ibiza. Sant Josep, Sa Talaia, Roques altes, 31SCD4907, 375 m, 05-V-1996, A. Barber & J.C. Cristóbal (ABH 33308); Ibiza, Sant Miquel, Puig Gros, 31SCD7226, 280 m, 04-V-1996, A. Barber & J.C. Cristóbal (ABH 33285); Ibiza, carretera de San Juan, km 10, 07-V-1980, Rivas-Martínez, M. Costa & J. Loidi (VAL 10767). Jaén. Villacarrillo, entre Mogón y Agrupación de Mogón, 30SYH9513, 520 m, 30-IV-2005, A. Juan & P. Vila (ABH 50074); Hornos, cerca del Collado de Hontan-ares, Sª de Segura, 30SWH2629, 950 m, 22-VI-1984, Pajarón & Rodríguez Pascual (MA 508215); Jaén, paraje Las Lagunillas, 30SVG3183, 440 m, 21-IV-1986, C. Fernández (GDA 25550, JAEN 860344, MA 505398, MGC 33438, FCO 18929); Sierra Mágina, Mojón Blanco, 1350 m, 13-VI-1926, Cuatrecasas (BC 61917). La Rioja. Logroño, 22-V, Zubia (MA 21929); Logroño, Armensia (MA 21954). León. Peñar-rubia, Embalse de Peñarrubia, 09-VI-1983, J. Iranzo (VAL 121255); Carbajal de la Legua, río Besnega, 30TTN5925, 850 m, 27-V-2006, M. Mart. Azorín, M.B. Crespo & C. Pena (ABH 51028). Lérida. La Noguera, Cubells, entre Cubells y la Torre de Fluvià, 31TCG3134, 400 m, 25-V-1989, J. Pedrol (MA 487407); Cubells,

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31TCG2936, 450 m, 22-V-1983, J. Pedrol (MA 418464). Madrid. Villaconejos, 03-VI-1976, Ferránz Díez (MA 202206, SEV 27701); Cerros de Alcalá de Henares, Villalbilla, 21-V-1995, E. Guinea (MA 432619); Rivas de Jarama, 20-V-1919, C. Vicioso (MA 21921); Aranjuez, 30-V, Cutanda (MA 21926). Málaga. Antequera, Sª Chimenea, pr. Cortijo de los Álamos, 30SUF5293, 580 m, 11-V-2005, M.B. Crespo, C. Pena, M. Mart. Azorín & B. Coca (ABH 48456); Ronda, P.N. Sierra de las Nieves, prox. al carril de subida al Puerto de los Pilones, 30SUF1762, 1300 m, 12-VI-1997, D. Navas, P. Navas, Y. Gil & A.V. Pérez-Latorre (MGC 45162); Málaga, La Araña, fábrica de cementos, zona este, 30SUF8264, 50 m, 29-IV-2003, O. Gavira (MGC 56775); Sª Tejeda, bosquete de Zafarraya, 900 m, 01-VI-1982, J.M. Nieto (MGC 20323). Mal-lorca. Bunyola, 08-V-1947, Palau Ferrer (MA 21958). Menorca. Matxani, Binixabonet, Santa Eulalia de Alayor (cf. J.J. Rodríguez Femenias, Flórula de Menorca: 128. 1901). Murcia. Atamaría, 37º35’41”N 00º49’20”W, 110 m, 03-V-2002, C. Aedo (MA 692687); Cartagena, Los Nietos, 17-V-1903, F.A. Ibañez (MA 158391); Jumilla, Sª del Carche, 1050 m, XH6254, 13-VI-1982, F. Alcaraz (MUB 7005); Sª Espuña, 1400 m, XG2694, 26-VI-1986, A. Robledo Mira (MUB 38809). Palencia. Arenas de Valdeginate, cerca de la ermita, 05-VI-2001, J.J. Lastra (FCO 26514, FCO 26515); Antigüedad, La Nava, 921 m, 30TVM0942, 09-VI-2007, L. Delgado & M. Santos Vicente (SALA 123154). Salamanca. Gallegos de Argañán, 03-VI-1976, E. Rico (SALA 10484); Bóveda del Río Almar, 30TTL1325, 02-VI-1983, Fdez.-Arias & Ruiz Téllez (SALA 69039); Estación de Villares de Yeltes, 23-VI-1978, F. Amich (SALA 15696); Almenara de Tormes, 10-VI-1977, J. Sánchez (SALA 17708). Segovia. Hontalbilla, 15-VI-1983, T. Romero (SALA 40352); Los Valles de Fuentidueña, 17-VI-1984, T. Romero (SALA 40353); Siguero, Aldealapeña, 1080 m, 19-VI-1983, T. Romero (MA 566604). Sevilla. Montellano, 16-V-1986, F.J. Fdez. Díez (SALA 48860); Aznalcázar, Reserva Guadiamar, 29SQA49, 25-IV-1978, S. Castro-viejo & E. Valdés-Bermejo (MA 258883); Morón de la Frontera, Sierra Espartero, ladera norte, 28-IV-1984, C. Barneto, V. Caravaca et al. (SEV 105948); Car-mona, Ermita de Santa María de Gracia, 24-IV-1979, P. Candau, I. Fernández & T. Luque (SEV 124197). Tarragona. La Franqueta, c. Horta, 14-VI-1915, P. Font Quer (MA 21956); Priorat, Ulldemolins, entre Coll de Albarca y Ulldemolins, 31TCF27, 600 m, 30-V-1989, J. Pedrol (MA 487272). Teruel. Maicas, Eras del cabezo San Jorge, 30TXL7737, 960 m, 23-VI-1998, Mercadal (VAL 107350); Torre de Arcas, 31-V-1992,

C. Fabregat & S. López Udías (VAL 78567); El Pinar (vivias), 1884 (MA 21953). Valencia. Chera, acceso al embalse de Buseo, 06-VI-1984, E. Sanchís & J. Alcober (VAL 61133); Algar de Palancia, 30SYK20, 402 m, 20-IV-1992, A. Cambronero (VAL 78399); Corral de Diego, Sª Benicadell, Font Freda, 30SYJ20, 650 m, 29-V-1985, G. Mateo, R. Figuerola & J.R. Nebot (MA 433070, VAL 142595); Sierra de la Valldigna, V-1897, C. Pau (MA 21951). Valladolid. Fuente el Sol, V-1944 (GDA 37596); Bocigas, 05-VI-1985, F.J. Fdez. Díez & X. Giráldez (SALA 41526); Castronuño, 30TUL1084, 13-V-1988, C.J. Valle & G. Balbás (SALA 84219); Encinas de Esgueva, 25-VI-1985, J.L. Fernández Alonso (MA 308260). Vizcaya. Estarrona, WN1946, 500 m, 07-VI-1983, G. Morante (BIO 1978). Zamora. Cañizal, 05-VI-1981, X. Giráldez (SALA 29499); Fuentesaúco, 06-VI-1981, X. Giráldez (SALA 29514); Fuentesecas, Gafos, 15-V-1987, R. García Río (SALA 54504); Cubillos, Caldegatos, 22-V-1987, R. García Río (SALA 54505). Zaragoza. Peña-fl or, cerca de San Cristóbal, 30TXM8526, 290 m, 24-V-1996, Mercadal (VAL 99587); Sigues, Venta Garrica, cerca soto río Esca, 30TXN6220, 500 m, 31-V-1986, L. Villar (VAL 16659). PORTUGAL. Algarve. Albu-feira, 23-IV-1968, A. Segura Zubizarreta (MA 355952); Lugar da Fornalha, estrada para Estoi, a 2 km de Mon-carapacho, 27-IV-1986, A. Moura (MA 432101); Loulé, cerro da Zorra, ponsio da encosta a Norte, 16-V-1979, Malato-Beliz & J.A. Guerra (MA 285459); Burgau, 26-V-1978, J.A. Devesa, J. Pastor & S. Talavera (SEV 38306). Alto Alentejo. Elvas, 29-IV-1966, A. Segura Zubizarreta (MA 355999); Reguengos de Monsaraz, cruzamento para Carrapatelo, 05-V-1982, Malato-Beliz & J.A. Guerra 16941 (SEV 89287); Viana do Alentejo, Alcaçovas, Herdade das Gigantes, 11-V-1955, J.R. Mendes (LISI 82/1999); Évora, S. Manços, Her-dade do Casao, num pousio, V-1947, A. Murteira (LISI 78/1999). Baixo Alentejo. Beja, Santiago Maior, a entrada para a Base Aérea (lado da E.N. 18), 250 m, 05-V-1987, M.L. Rocha Afonso (LISI 43/1999); Cuba, a 2 km de Cuba, na estrada para Faro do Alentejo, 02-VI-1972, A. Leitao (LISI 24/1999); Ferreira do Alentejo, Peroguarda, Serra do Mira, 03-V-1996, I. Moreira, A. Monteiro, E. Sousa & T. Vasconcellos (LISI 42/1999); Serpa, Herdade do Barrocão, 17-IV-1952, F.G. Palma (LISI 65/1999). Beira Litoral. Can-tanhede, Povoa do Conde, 25-V-1972, F. Bellot, B. Casaseca & S. Castroviejo (SALA 4225); Porto de Mós, Porto de Mós-Pragais, 230 m, 24-V-1982, J. Franco & M. Correia (LISI 17/1999). Estremadura. Sesimbra, cabo Espichel, 01-VI-1971, Malato-Beliz & J.A. Guerra (MA 285466); Bombarral, Carvalhal, qta.

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da Gafa, 75 m, 20-IV-1984, M.D. Espírito Santo (LISI 15/1999); Cadaval, Cercal, perto da Cada das Barreiras Vermellas, Sª Montejunto, 230 m, 11-V-1989, M.D. Espírito Santo & J.C. Costa (LISI 98/1999); Palmela, Casal Novo, 140 m, 04-VI-1963, A.C. Marcelino (LISI 68/1999). Ribatejo. Alcanhoes, V-1941, A. Pereira Coutinho (LISI 81/1999); Cartaxo, Pontével, Casais de Lagartos, 02-VII-1980, T. Vaconcellos & al. (LISI 11/1999); Torres Novas, S. Pedro, Banda de Além, 24-IV-1970, J. Franco & M.M. Fonseca (LISI 48/1999); Vila Franca de Xira (Monte Gordo), 190 m, 04-V-1942, J.M. de Carvalho & F.M. Flores (LISI 80/1999). Trás-os-Montes. Entre Pocinho e Coa, V-1942, G. Barbosa & M. Myre (LISI 79/1999).

2. Ornithogalum pyrenaicum L., Sp. Pl.: 306. 1753 subsp. pyrenaicum

≡ Loncomelos pyrenaicus (L.) L.D. Hrouda ex J.Holub in Folia Geobot. Phytotax. 23(4): 413. 1988 ≡ O. fl avescens Lam., Fl. Franç. 3: 277. 1778, nom. illeg. [syn. subst., et non Jacq., Coll. 3: 229. 1790]

= Loncoxis sulfurea (Waldst. & Kit.) Raf., Fl. Tellur. 3: 58. 1837 ≡ Anthericum sulfureum Waldst. & Kit., Descr. Icon. Pl. Hung. 1: 98. 1802 ≡ Ornithogalum sulfureum (Waldst. & Kit.) Schult. & Schult. f., Syst. Veg. ed. 15 bis [Roemer & Schultes] 7(1): 518. 1829 — Type: not seen (synonym after description and type locality).

= O. granatense Pau in Mem. Mus. Ci. Nat. Barcelona, Ser. Bot. 1(1): 69. 1922 — Lecto-type (here selected): BC 61929!

Ind. loc. “Habitat in Alpibus Helveticis, Geneven-sibus, Pyrenaicis”.

Lectotype. LINN 428.5 [designated by EL-GADI in JAFRI & EL-GADI (eds.), Fl. Libya 57: 50. 1978; but see additional comments by STEARN (1983)].

Ilustrations. PASTOR, Fl. Vasc. Andalucía Occid. 3: 436. 1987; WITTMANN in Stapfi a 13: 24, 25. 1985; Fig. 6.

Description. Geophyte. Bulb (32)34-50(58) × (22)26-36(38) mm, ovoid, generating contractile roots, usually without secondary bulbils, and tapering into a narrow neck 10-20 mm; outer tunics pale brown. Leaves (3)4-7(8), in a basal

rosette, (38)40-75(80) × (0.6)0.7-0.9(1.3) cm, lin-ear or tapering, slightly keeled, pruinose, gla-brous, green somewhat glaucous, withering at the apex, proteranthous. Floral stem (excluding the infl orescence) (46)48-78(85) × 0.4-0.5 cm, erect, smooth, somewhat glaucous and pruinose. Infl o-rescence racemose, very long and pyramidal at the apex, (16)20-45(51) × (0.7)1.5-2.6(3) cm, (excluding fl owers but not their pedicels), with (20)25-55(64) fl owers; bracts (7)8-15(19) × (2)2.5-3.7(4) mm, shorter or equal than pedicels, rarely a bit longer, ovate-lanceolate, much wider at the base, with apex setose-acuminate, mem-branose, with 3 greenish nerves; fl oral pedicels erect-patent, the lower ones (9)13-25(29) mm, the middle ones (8)11-17(19) mm and the upper ones (2)3-7(9) mm; fruiting pedicels 18-25(29) mm, erect and appressed to the stem. Flowers 18-23 mm of diameter, slightly fragrant; tepals yellow-ish in the adaxial side and yellowish with a cen-tral green band in the abaxial side of 1-1.2 mm width, linear to linear-lanceolate, with smooth edges at the beginning of the anthesis, with incurved or undulate edges when mature; outers 7-11(12) × (1.3)2-2.3 mm; inners 7-10(11) × (1.5)2-2.5 mm, being a little shorter and wider than the outers. Stamens 6, 1/2 to 2/3 of the tepal length; fi laments white, abruptly widened in the basal half, outers 5-6 × 1.2-1.4 mm, inners 5.5-6.2 × (1.2)1.4-1.8 mm, being somewhat longer and wider than the outers; anthers dorsifi xed, pale yel-low, 2.5-3 × 1-1.2 mm before dehiscence and 1.5-1.8(2) × 0.7-0.9(1.2) mm after dehiscence. Ovary (1.5)2-2.5(2.8) × (1.2)1.5-2.5(2.7) mm, pale green, ovoid, sphaerical or shortly cylindrical, truncate in the apex, trigonous with 3 obtuse ribs and with septal nectaries; style whitish, fi liform, (2)2.5-3.5(4) mm; stigma slightly trigonous and glandulose. Capsule (7)8-9(9.5) × (5)5.5-6.5(7) mm, ovate-cylindrical, trigonous, pale brown, with trivalvar dehiscence. Seeds (6)10-18(20) per fruit (n = 32, mean = 12.25, SD = 4.70), with 5-7.5 mg weight, (2.3)2.5-3.1(3.3) × (1.4)1.6-2(2.3) mm, angulose and irregularly compressed, and testa irregularly rugose to almost minutely reticu-late, sometimes with smaller granules scattered on the seed edges.

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Fig. 6. Ornithogalum pyrenaicum (Puebla de Guzmán, Huelva, Spain): a: fl ower, frontal view; b: fl ower, dorsal view; c: infl orescence; d: fruiting raceme; e: bract; f: inner (left) and outer (right) stamens; g: gynoecium; h: cap-sule, lateral view; i: capsule, apical view; j: capsule, transversal section; k: bulb and leaves. Scale bars = 1 cm.

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Chromosome number. 2n = 16 (NEVES 1952, 1973, GADELLA et al. 1966 as O. pyramidale, WITTMANN 1985, MORET 1987, PASTOR & DIOS-DADO 1994). 2n = 16 + (O-2B) (CASTROVIEJO 1984, TORNADORE 1985). 2n = 24 (NEVES 1952 as O. pyramidale). According to TORNADORE (1985), O. pyrenaicum is a diploid taxon, with basic chro-mosome number x = 8.

Biology. It fl owers from May to June, or even July in higher areas. Fruiting extends from June to August. Reproduction is mainly by seeds, which are produced in a high number per infl orescence. However, it can also produce a low number of secondary basal bulbils, though vegetative repro-duction is less signifi cant than that by seeds (cf. HILL & PRICE 2000).

Ecology. This species prefers shady and fresh sites, such as forests, grasses or river edges. It usually grows mainly on soils with acidic nature, though it can also be found on other different types.

Distribution. Europe, northern Africa and Middle East. It is distributed from Portugal to Turkey and central and eastern Russia, and from Morocco to southern England (cf. WITTMANN 1985, HILL & PRICE 2000). In the Iberian Peninsula, it is rela-tively common in the northern and western parts (Fig. 7). Moreover, it does not grow in the eastern and southeastern Iberian Peninsula and the Bal-earic Islands, where soils are predominantly of basic nature.

Observations. This species is quite easy to iden-tify, mainly in fresh. However, when the herbar-ium materials are badly conserved or some parts are lacking, it can be misidentifi ed as O. narbon-ense. However, the shape, size and colour of the tepals, or the microstructure of the testa allow an unequivocal identifi cation. PAU (1922) described O. granatense Pau from Villanueva del Rosario (Málaga), on the basis of its shorter and thicker fl oral peduncles and bracts wider at the basis, though explicitly indicating that it was close to O. pyrenaicum. Those characters however fall into

Fig. 7. Map of the studied material of O. pyrenaicum in the Iberian Peninsula and the Balearic Islands. Dots to cor-respond both studied natural populations and herbarium sheets.

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the specifi c variation of the latter, to which O. granatense is here synonymised. IUCN (2001) category: LC.

Selected material. ANDORRA. Sant Julià de Loria, Pont de la Marginada, 31TCH7604, 980 m, 6-VII-1992, C. Aedo, J.J. Aldasoro et al. (MA 512050). SPAIN A Coruña. Arzua, 29TNH6452, 425 m, 09-VI-2000, I. Pulgar (SANT 43175). Asturias. Teverga, Desfi ladero de Fresnedo, prox. Cueva Huerta, 29TQH3978, 670 m, 12-VI-2006, J.C. Cristóbal (ABH 50811). Ávila. Puerto del Tremedal, Presa de Santa Lucía, 30TTK8069, 1500 m, 30-VI-1999, J.C. Cristóbal (ABH 41982); Valle del río Corneja, risco del Avión, 08-VII-1986, J. Estrada (MA 506483); Valle del Amblés, puerto de Villatoro, 1400 m, 12-VII-1979, M. Ladero (MA 258888); Pinar de Hoyocasero, 30TUK3273, 1250-1325 m, 01-VII-1985 (MA 427548); Aliseda de Tormes, 09-VII-1975, E. Valdés-Bermejo (MA 433696). Badajoz. Magacela, TJ60, 20-V-2001, P. Escobar García (MGC 61297). Barcelona. Fogars de Montclús, Santa Fe del Montseny, El Vallès Oriental, DG5524, 1160 m., 02-VI-2001, G. Mercadal (HGI 17277). Burgos. Hoyos del Tozo, 30TVN2328, 900 m, 15-VI-1985, J.A. Alejandre (MA 338378); Berberana, VI-1928, M. Sosa (MA 21904); Huidobro, 30TVN4435, 18-VI-1987, P. Galán Cela & A. Martín (MA 466154); Cercanías de Dobro, 30TVN4639, 04-VII-1987, P. Galán Cela & A. Martín (MA 466145); Humada, San Martín de Humada, Mte. Portillo, 30TVN1521, 1150-1226 m, 29-VII-1987, B.Fz. De Botoño & J.A. Alejandre (MA 422945). Cáceres. Hervás, pista de Heidi, 30TTK5660, 750 m, 27-V-1987, R. González & J.A. Sánchez Rodríguez (SALA 101681); San Martín de Trevejo, 29TPE8858, 06-I-1982, A. Valdés Franzi (SALA 74244); Barrado, 30TTK5441, 04-VI-1987, A. Amor (SALA 76861); Robledillo de Gata, Monte de la Boya, 29TQE17, 28-VI-1988, J.A.Devesa & M.C. Viera (MA 507400); Alrededores de Guadalupe, 17-VI-1948, A. Caballero (MA 21899). Cantabria. La Lomba, Hermandad de Campóo de Suso, 30TUN96, 1000 m, 10-IV-1984, C. Aedo (MA 621549); Espinama, 22-VII-1954, Casaseca (SANT 8933). Ciu-dad Real. Puebla de Don Rodrigo, Bco. del Pocito, 30SUJ6430, 750 m, 10-V-1992, Carrasco, Garrido & Martín-Blanco (MACB 71021); Piedrabuena, Hoya de Piedrabuena, 30SUJ9621, 570 m, 14-V-1994, Carrasco & Martín-Blanco (MA 621167); Fuencaliente, valle del río Cereceda, 30SUH8753, 640 m, 25-V-1997, R. Garcín & J. Barios (MA 596542). Córdoba. Sierra Morena, barranco del Martín Gonzalo, UH8729, 680 m, 17-VI-1993, M. Melendo (GDAC 41750); Entre Ada-muz y Villanueva de Córdoba, Venta del Puerto, 14-V-

1984, J.A. Devesa & B. Valdés (SEV 120467). Gerona. Sant Julià del Llor, en un antic codolar del riu Ter, La Selva, DG6946, 140 m, 15-V-2003, M. Corominas & L. Villar (HGI 18005). Guadalajara. Corduente, barranco de la Virgen de la Hoz, 30TWL8520, 1040 m, 17-VI-1998, J.M. Herranz (MA 620153); Piqueras, 1700 m, 03-VII-1957, A. Segura Zubizarreta (MA 355409). Huelva. Puebla de Guzmán, orilla del río Malagón, 29SPB5873, 24-V-2004, M. Martínez Azorín, M.B. Cre-spo & C. Pena (ABH 50135). Huesca. Fiscal, VI, C. del Campo (MA 21902). Jaén. Andújar, Nava de la Col-mena, 30SVH0120, 02-VI-1994, F. Gómez Manzaneque & C. Morla (MA 652636); Despeñaperros, in mont. marianis, el Bco. de Valdefl ores, 08-VI-1924, Font Quer (BC 76671, BC 76669). La Rioja. El Rasillo de Cam-eros, VI-1928, Zubia (MA 21905). León. Andiñuela, 11-VII-1947, F. Bernis (MA 21908); Ponferrada, mon-tes Aquilinos, Los Apóstoles, 29TPH9702, 1600 m, 18-VII-1982, G. Nieto Feliner (MA 280062); Oseja de Sajambre, 17-VII-1927, C. Pau (MA 21909); Portilla de la Reina, 25-VII-1972, J. Andrés (MGC 12540). Lé -rida. Espot, pirineos centrales, 31TCH41, 1350 m, 20-VI-1981, A. Carrillo & J.M. Ninot (SALA 33293, SEV 114020, VAL 142592). Lugo. Villardiaz-Fonsagra, 20-VII-1953, E. Carreira (MA 167872); Doiras, Sierra de Ancares, 29TPH6639, 800 m, 28-VI-1982, S. Cas-troviejo, P. Coello, P. Galán & G. Nieto (MA 417286, MA 432533); Monforte de Lemos, Distriz, 29TPH1807, 300 m, 07-VI-2008, J. Amigo (SANT 59109). Madrid. La Pedriza Anterior, 21-VI-1973, S. Rivas Martínez (SALA 28348); Somosierra, camino del monte, 1450 m, 19-VII-1984, T. Romero (MA 566602, SALA 40354); Cercedilla, Sierra de Guadarrama, VI-1914, C. Vicioso (MA 21897); Rozas de Puerto Real, 30TUK7263, 08-VI-1986, F.G. Manzaneque (MA 556909); El Esco-rial, El Castañar, 16-VI-1907, M. Rodríguez López-Neura (MA 349496); Buitrago, 21-VI-1918, C. Vicioso (MA 21894). Málaga. Villanueva del Rosario, VI-1919, Gros (BC 61929). Navarra. Ulzama, entre Alcoz e Iraizoz, río Ulzama, puente de la carretera, 30TXN0861, 550 m, 28-VI-1993, I. Biurrun (BIO 20772). Orense. Carballeda, Teixal de Peña Trevinca, 1300-1500 m, 19-VII-1984, S. Ortiz (SANT 16632). Palencia. Velilla del Rio Carrión, 30TUN4945, 1140 m, 10-VII-1995, C. Navarro et al. (VAL 104621); Cervera de Pisuerga, Peñas Negras, 06-VIII-1914, P. Font Quer (MA 21903). Pontevedra. A Golada, Eidián, Galegos, 29TNH8044, 320 m, 10-VI-1999, J. Amigo & R.I. Louzán (SANT 41632); Vila de Cruces, Piloño, Pastoriza de Raindo, 29TNH5939, 170 m, 09-VI-1999, J. Amigo & R.I. Louzán (SANT 41677). Salamanca. Sierra de la Peña de Francia, 20-VI-1972, F.J. Fdez. Díez (SALA 12484);

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Aldeadávila, 16-VI-1977, F. Amich (SALA 15697); Masueco, ribera del río Uces, 16-VI-1976, F. Amich (SALA 15698); La Fregeneda, 27-V-1977, F. Amich (SALA 15699); La Alberca, 27-VI-1946, A. Caballero (MA 21910). Soria. Villar del Ala, El Valle, 30TWM3540, 1150 m, 14-VII-1996, Montamarta (VAL 99185); Valonsadero, 03-VII-1968, A. Segura Zubi-zarreta (MA 355365). Toledo. Sierra de San Vicente, 1200 m, 18-V-1992, M.A. Carrasco, J.L. Castillo, C.J. Martín-Blanco & S. Pajarón (MA 621679). Vizcaya. Arceda-Avellaneda, barranco de Musita, 30TWN4836, 790 m, 15-VII-1991, J.M. Olano (BIO 23541); Lezaum, WN6150, 1000 m, 04-VII-1981, J.A. Alejandra (BIO 1977). Zamora. Vega del Castillo, Entre las Puentes, 07-VI-1988, R. García Río (SALA 54503); San Martín de Castañeda, 29TPG8866, 1000 m, 26-VI-1987, P. García & A. Roa (MA 509830); Tábara, monte La Folguera, 30TTM5236, 770 m, 15-VI-1996, P. Bariego Hernández (MA 651712). PORTUGAL Algarve. Serra de Monchique, Cruz da Foia, velho souto de tal-hadio, 21-VI-1979, Malato-Beliz & J.A. Guerra (MA 285470). Alto Alentejo. Reguengos de Monsaraz, cru-zamento para Carrapatelo, 05-V-1982, Malato-Beliz & J.A. Guerra (MA 411467); Povra e Maeadas, río Sever, 10-VI-1956, Malato-Beliz & J.A. Guerra (MA 285473); Castelo de Vide, Atalaia, 24-VI-1956, Malato-Beliz (MA 285472); Montalvao, Dourados, barreiras do rio Sever, 03-VI-1951, Beliz & Ruivo (MA 285474). Baixo Alentejo. Arredores de Barrancso, Herdade das Russia-nas, 13-V-1959, Malato-Beliz & J.A. Guerra (MA 285471). Beira Baixa. Castelo Branco, Ladoeiro, rib-eras del río Ponsul, 29SPD4213, 180 m, 30-IV-1994, E. Rico, F. Amich et al. (MA 717032); Penamacor, Mei-moa, Valdedrinha de Baixo, num carvalhal negral, Serra Malcata, PE65, 590 m, 13-VI-1987, M. Lousa, M.L. Rosa & J.P. Luz (LISI 99/1999); Fundao, Alcaide, vert-ente oriental do Cabeço de S. Macário, 18-VII-1955, J.A. Franco (LISI 142/1999). Douro Litoral. Vila Nova de Gaia, Cabadelo, numa bouça, 13-VI-1941, J. Castro (MA 188632). Estremadura. Sintra (arredores) em famares proximo do Algueirao, 30-IV-1944, Bento Rainha (GDA 37602, GDA 37603); Sª Mª de Azeitas, arred. Alambre, margen da ribeira, 25-V-1943, Silva, Gedraet & Fontes (SANT 2893); Cascais, Alcabideche, Qta. do Marques, 80 m, 06-V-1963, V.E. Moreira Lopes (LISI 141/1999). Trás-os-Montes. Casais do Douro, Batevias, 10-VI-1941, G. Pedro (MA 412250). Urrós, río Duero, 29TQF1578, 350 m, 20-V-1997, Cristóbal, Crespo, Camuñas, Juan & Soler (ABH 38762); Lamego, mata junto a Lahin, 12-VI-1943, F. García & M. Myre (LISI 148/1999); Vila Flor, mata dos Castelhanos, 17-VI-1942, G. Barbosa & M. Myre (LISI 152/1999).

ACKNOWLEDGEMENTS

Curators of ABH, BIO, BC, BCN, COI, FCO, GDA-GDCA, HGI, HUAL, K, LISI, MA, MACB, MGC, P, SALA, SANT, SEV and VAL, are gratefully thanked for the loan of herbarium material. José R. Verdú and Catherine Numa (University of Alicante) helped with map drawing. Enrique Rico (University of Salamanca) sent bulbs of O. pyrenaicum

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