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A New Autumn-Flowering Species of Allium (Alliaceae) from Croatia

Article in Folia Geobotanica · March 2009 DOI: 10.1007/s12224-009-9032-2

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A New Autumn-Flowering Species of Allium (Alliaceae) from Croatia

Sandro Bogdanović & Salvatore Brullo & Gianpietro Giusso del Galdo & Cristina Salmeri

# Institute of Botany, Academy of Sciences of the 2009

Abstract Here we describe Allium telmatum Bogdanović, Brullo, Giusso & Salmeri, a new species of Allium sect. Codonoprasum from North Dalmatia (Croatia). Its chromosome number (2n=32), karyotype, leaf anatomy, ecology and taxonomical relationships are examined. Several features, such as phenology (flowering in autumn), occurrence in coastal salt marshes, tetraploid chromosome number, and morphology, indicate that it is most closely related to the Tyrrhenian species Allium savii.Onthe basis of our herbarium survey, we present here a distribution map of the autumn- flowering species of Allium sect. Codonoprasum in the Mediterranean area.

Keywords Allium telmatum . Allium sect. Codonoprasum . Dalmatia . Karyology . Leaf anatomy .

Introduction

The genus Allium L. comprises about 750 species naturally distributed throughout the Northern Hemisphere (Stearn 1978, 1992). It is mostly limited to seasonally dry regions, with a main center of diversity in the Middle East and Central Asia, and a

S. Bogdanović Department of Botany and Botanical Garden, Division of Biology, Faculty of Science, University of Zagreb, Marulićev trg 9a, 10000 Zagreb, Croatia e-mail: [email protected]

S. Brullo (*) : G. Giusso del Galdo Dipartimento di Botanica, Università di Catania, via A. Longo 19, 95125 Catania, Italy e-mail: [email protected] G. Giusso del Galdo e-mail: [email protected]

C. Salmeri Dipartimento di Scienze Botaniche, Università di Palermo, via Archirafi 38, 90123 Palermo, Italy e-mail: [email protected] 84 S. Bogdanović et al. smaller one in North America. The treatment of the genus Allium is quite complex, both from the taxonomical and nomenclatural viewpoints, because many subgenera and sections are recognized (Hanelt et al. 1992), and lots of species names and synonyms exist (Gregory et al. 1998). Taxa widely cultivated as a vegetable belong to Allium subgen. Allium (garlic, leek) and Allium subgen. Rhizirideum (G. Don ex Koch) Wendelbo (onion, chives, shallot), while many ornamental species and cultivars are found in Allium subgen. Melanocrommyum (Webb & Berthel.) Rouy. Relevant diagnostic characters include bulb morphology, leaf number, leaf shape and size, structure, flower morphology (, and ovary with nectaries), as well as chromosome number, phenology and phytogeographical patterns. More recently, also chloroplast and nuclear DNA (ITS) data have been used to support the monophyletic origin of Allium, and to refine its taxonomy (Mes et al. 1999; Somoylov et al. 1999; Raamsdonk et al. 2003; Friesen et al. 2006; Nguyen et al. 2008). The genus Allium comprises 56 taxa (species and subspecies) distributed in Croatia (Nikolić 2008). Within the subgenus Allium, A. sect. Codonoprasum Reichenb. is one of the most interesting and hitherto less investigated section in this area, which includes several species, such as A. paniculatum L., A. flavum L., A. dentiferum Webb & Berthel., A. pallens L., A. oleraceum L., A. tenuiflorum Ten., and A. croaticum Bogdanović et al. All these species are well differentiated in many morphological and karyological features, and usually flower from late spring to early summer (May−July) (Jauzein and Tison 1999; Brullo et al. 2002, 2003a, 2008; Bogdanović et al. 2008). During our field investigations, a peculiar population of Allium growing in salt marshes of the island of Vir (Croatia) was found. This geophyte flowers in autumn and grows on damp soils characterized by a certain accumulation of salts. This ecological requirement is rather rare among the autumn-flowering Allium species, observed so far only in Allium savii Parl. According to Brullo et al. (1994), A. savii occurs in the Tyrrhenian area, where it has a scattered distribution ranging from Latium and Tuscany in Italy, through , Corsica to southern France, and recently reported also from Menorca (Fraga 2002). The Croatian population appears morphologically quite similar to A. savii, mainly in its habit, spathe, inflorescence and some flower characters, while it differs in the number, shape and anatomy of leaves, size and shape of tepals, stamens and capsule. Based on herbarium surveys and literature data, this population has never been collected, and is treated here as a new species, Allium telmatum.

Material and Methods

A total of 30 living mature from five sites on the island of Vir in Croatia (Table 1) were collected and cultivated in the Botanical Garden of Catania (Italy). Qualitative and quantitative morphological characters, considered as diagnostic in Allium, were measured and scored on fresh material (Table 2). Voucher specimens are deposited in CAT, FI and ZA. Leaf anatomy was studied on cultivated plants from the type locality (Table 1), using leaf blades of maximum size in their optimal vegetative development, usually before the A new autumn-flowering species of Allium (Alliaceae) from Croatia 85

Table 1 List of collection sites of Allium telmatum from the island of Vir (Croatia). The first one corresponds to the type locality

Locality and collection details Nr. of sampled plants Nr. of plants used for chromosome counts

Rt Stinice, near Virić, 44°16´50˝ N, 15°07´33˝ E, s. l. 8 3 Rt Stinice, near Virić, 44°16´55˝ N, 15°07´35˝ E, s. l. 6 - Rt Stinice, near Virić, 44°17´26˝ N, 15°07´92˝ E, s. l. 6 2 Rt Stinice, near Virić, 44°17´16˝ N, 15°07´10˝ E, s. l. 5 - Rt Stinice, near Virić, 44°17´57˝ N, 15°07´50˝ E, s. l. 5 -

All specimens were collected on 4 Sept 2007 by Bogdanović, Brullo, Giusso del Galdo, s.n. flowering stage. The leaf tissue was fixed in a mixture of ethanol, chloroform and acetic acid (3:3:1) according to the protocol by Karpechenko (1927), and embedded in paraffin; the transverse sections were stained with ruthenium red and light green. Karyological analyses were performed on mitotic plates from root-tip cells of cultivated bulbs, pre-treated with 0.3% colchicine water solution, fixed in ethanol- acetic acid mixture (3:1), and stained according to the Feulgen technique (Feulgen and Rossenbeck 1924). Metaphase observations and chromosome measurements were conducted using the image analysis systems IKAROS 4.6 (Metasystem) and Zeiss AxioVision 5.1. Karyotyping was done with the specialized software Cromolab© 1.1 (Brullo 2002–2003), which was used to recognize homologous chromosomes, order them by size, classify them according to morphology, and assemble the karyotype formula based on the centromere position (Levan et al. 1964; Tzanoudakis 1983). Ten mitotic plates from five individuals were used for assembling the karyotype.

Results

Description of a New Allium Species

On the basis of our morphological investigation (Table 2, Figs. 1, 2), leaf anatomy features (Fig. 3), and chromosome numbers (Fig. 4), we describe here the following species.

Allium telmatum Bogdanović, Brullo, Giusso & Salmeri, sp. nov.

Type Croatia, North Dalmatia, island of Vir, salt marsh at Rt Stinice near Virić, 44° 16´50˝ N, 15°07´33˝ E, cultivated specimen, 4 Sept 2007, Bogdanović, Brullo, Giusso del Galdo, s.n. (Holotype: CAT; isotypes: CAT, FI, ZA).

Descriptio Ab Allio savii bulbo 15−25 mm longo, 10−15 mm lato, scapo 30−40 cm alto, inserto extra bulbum, foliis 3−4, max. 15 cm longis, 2−2.5 mm latis, 5-costatis, spathae valva maiore usque ad 12 cm longa, valva minore usque ad 7 cm longa, pedicellis usque ad 40 mm longis, tepalis 5.5−6 mm longis, interioribus 3 mm latis, 86 S. Bogdanović et al.

Table 2 Morphological characters differentiating between Allium telmatum and A. savii

Character A. telmatum A. savii (n=30) (n=25)

Bulb size (mm) 15−20×10−15 12−15×10−12 Outer tunics coriaceous fibrous Stem length (cm) 30−40 45−55 Bulb stem insertion external internal Leaf number 3−45−6 Leaf ribs 56−7 Leaf max. length (cm) 15 25 Leaf width (mm) 2−2.5 1−2 Leaf palisade tissue 2-layered 1-layered Leaf vascular bundles usually 19 usually 10 Larger spathe valve length (cm) 6−12 8−16 Smaller spathe valve length (cm) 2.5−75−11 Pedicel length (mm) 10−40 10−30 Number of flowers 10−28 20−28 shape elliptical oblong-elliptical Tepal length (mm) 5.5−66−7 Outer tepals width (mm) 3.2−3.5 2.5−3 Inner tepals width (mm) 3 2.5−3 Stamens exerted included Filament colour purplish above, white below white Outer length (mm) 2.2−32−2.2 Inner stamen length (mm) 3.6−4 2.5−3.3 Annulus height (mm) 1−1.2 1.5 Anther colour white-yellowish, tinged with pink white Anther size (mm) 1.5−1.6×0.8−0.9 1.4−1.5×0.7−1 Anther apex rounded apiculate Ovary size (mm) 3.5−3.7×2 4×2−2.2 Capsule size (mm) 4.5−5×3.7−4.2 4.5−5.5×5−5.5

For quantitative characters showing some variation, minimum and maximum values are given. Characters best discriminating between the species are set in bold. n – number of investigated plants. Data for A. savii were taken from Brullo et al. (1994). exterioribus 3.2−3.5 mm latis, staminibus interioribus exertis e perigonio, filamentis purpurascentibus superne et albidis inferne, exterioribus 2.2−3 mm longis, interioribus 3.6−4 mm longis, annulo 1−1.2 mm alto, antheris albido-luteolis e roseo suffusis, rotundatis apice, ovario 3.5−3.7 mm longo, capsula 4.5−5×3.7−4.2 mm, differt.

Description Bulb ovoid, 15−25×10−15 mm, with outer tunics coriaceous, black- greyish, the inner ones membranaceous, whitish. Stem 30−40 cm tall, cylindrical, glabrous, erect, half of its length covered by leaf sheaths, inserted externally to the bulb. Leaves 3−4, green, 5-ribbed (pentagonal in the cross section), 10−15 cm long, A new autumn-flowering species of Allium (Alliaceae) from Croatia 87

Fig. 1 Diagnostic morphological features of Allium telmatum. a – habit; b – spathe valves; c – perigon; d – open perigon with stamens; e – anther; f – ovary; g – capsule. Drawings are based on specimens from the type locality

2−2.5 mm wide. Spathe persistent, with two valves unequal, longer than the inflorescence, normally unilateral (sometimes fused at the base on one side), the larger 7-nerved and 6−12 cm long, the smaller 5-nerved and 2.5−7 cm long. Inflorescence lax and fastigiate, 10−28-flowered; pedicels unequal, 10−40 mm long. 88 S. Bogdanović et al.

Fig. 2 Inflorescence of Allium telmatum

Perigon campanulate, with tepals equal, white-pinkish, tinged with purplish, elliptical, apiculate at the apex, 5.5−6 mm long, the inner ones 3 mm wide, the outer ones 3.2−3.5 mm wide, midrib purplish. Stamens, at least the inner ones, exerted from the perigon with simple filament, purplish above and white below, unequal, the outer ones 2.2−3 mm long, the inner ones 3.6−4 mm long, connate below into an annulus, 1−1.2 mm high; anthers white-yellowish, tinged with pink, oblong-elliptical, 1.5−1.6×0.8−0.9 mm, rounded at the apex. Ovary obovoid, narrowed at the base, green, tuberculate above, 3.5−3.7×2 mm; style white, 0.8−1.2 mm. Capsule trivalved, obovoid, 4.5−5×3.7−4.2 mm, green.

Etymology The specific epithet refers to the Greek word telma, which means marsh.

Phenology Allium telmatum has an autumnal flowering period, ranging from early September to early November. This ontogenetic cycle was also observed on A new autumn-flowering species of Allium (Alliaceae) from Croatia 89

Fig. 3 Leaf cross section of Allium telmatum cultivated plants. A. telmatum also shows a vegetative reproduction mode through bulbils, locally forming scattered dense tufts.

Habitat and Distribution Allium telmatum grows in coastal salt marshes at the margin of periodically submerged sites. It prefers more-or-less damp muddy-limose soils, where it grows together with various helophytes and halophytes, such as

Fig. 4 Metaphase plate a and idiogram b of Allium telmatum (2n=32). Counted by Bogdanović & Salmeri 90 S. Bogdanović et al.

Limonium narbonense Mill., L. virgatum (Willd.) Fourr., Aster tripolium L., Inula crithmoides L., Juncus acutus L., J. maritimus Lam., Schoenus nigricans L. and Scirpus holoschoenus L. From the phytosociological point of view, this plant community can be referred to as Juncetum maritimo-acuti Horvatić 1934, an association of Juncetea maritimi Br.-Bl. 1952 (cf. Horvatić 1963). Allium telmatum occurs in a few localities of North Dalmatia (Croatia). It has been recorded so far from the islands of Vir and Pag, and at Plemići Bay near Miočići (Fig. 5). Except for the studied populations (listed in Table 1, all from the Vir island), the following three localities have also been documented: 1. North Dalmatia, island of Vir, east of the bridge, 44°17´8˝ N, 15°08´10˝ E, s.l., 21 Sept 2008, Boršić & Bogdanović s.n. (CAT, ZA). 2. North Dalmatia, island of Pag, Vlašići, Ričina Bay, 44°19´16˝ N, 15°12´40˝ E, s. l., 20 Sept 2008, Boršić & Bogdanović s.n. (CAT, ZA). 3. North Dalmatia, Miočići, Plemići Bay, 44°16´49˝ N, 15°18´43˝ E, s.l., 20 Sept 2008, Boršić & Bogdanović s.n. (CAT, ZA).

Leaf Anatomy The leaf cross section of A. telmatum shows a pentagonal outline. A well-developed and slightly denticulate cuticle covers the epidermis. The stomata are numerous and are distributed along the whole perimeter. The palisade tissue is regular and compact, and arranged in two-layered cylindrical cells; the outer layer usually has bigger cells. The spongy tissue is compact, with more-or-less irregular cells outward, and fistulous in the inner part. In the peripheral part of the spongy tissue several secretory canals occur. The vascular bundles are 16−19, of which 10−11 are abaxial (three are bigger) and 7−8 are adaxial (two are bigger) (Fig. 3).

Karyology A. telmatum is tetraploid with 2n=4x=32 (Fig. 4a). The karyotype has an apparently diploidized arrangement (Fig. 4b), as the chromosomes cannot be

Fig. 5 Geographical distribution of the autumn-flowering species of Allium sect. Codonoprasum in the Mediterranean area (based on our own herbarium survey): A. oporinanthum ( ); A. savii (●); A. anzalonei ( ); A. telmatum ( ); A. euboicum ( ); A. rausii (◆); A. platakisii (♢); A. aegilicum ( ); A. tardans (■); A. archeotrichon (▲); A. autumnale ( ); A. tardiflorum ( ) A new autumn-flowering species of Allium (Alliaceae) from Croatia 91 grouped into sets of four similar chromosomes (quartets), but they assemble in pairs of two homologues. The karyotype is characterized by eight more-or-less metacentric pairs of chromosomes, six pairs close to the submetacentric type (arm ratio exceeding 1.35), and two submetacentric pairs. Absolute chromosome length ranges from 15± 0.15 µm to 7.2±0.06 µm, while the arm ratio varies from 1 to 1.71. The karyotype formula can be inferred as: 2n=4x=32: 4 M + 12 m + 12 msm + 4 sm.

Discussion

The newly described Allium telmatum is closely related to A. savii; both have the same ecology, chromosome number and some morphological features. In fact, both species grow on coastal damp soils, they are tetraploid (2n=32), and show a very similar habit, with the same type of inflorescence and flower shape. However, they differ in some morphological characters, chiefly regarding the bulb size, stem insertion, leaf shape and size, spathe valves, tepals, stamens, ovary and capsule (Table 2). Other significant differences regard the leaf anatomy and the karyotype. A. savii has leaves characterized by 1-layered palisade tissue with 10 vascular bundles (sometimes more), the spongy tissue is quite compact, with big cells in the central part, or sometimes slightly fistulous, while in A. telmatum thepalisadetissueis2-layered,the vascular bundles are usually 19 (rarely less), and the spongy tissue is largely fistulous in the central part. As for the karyotype, A. savii has prevalently metacentric chromosomes, four chromosomes have microsatellites on their long arm, and three pairs are submetacentric (Brullo et al. 1994), while A. telmatum shows only two submetacentric pairs, and no evident microsatellites are observed. According to literature data (Kollmann et al. 1991; Tzanoudakis and Kypriotakis 1993; Brullo et al. 1994, 1997a,b,c, 1999, 2003b; Tzanoudakis 2000), a group of autumn-flowering species with a peculiar ontogenetic cycle can be distinguished within the Allium sect. Codonoprasum. It includes Allium savii Parl. (Italy, Corse, Sardinia, southern France, Menorca), A. autumnale Davis (Cyprus), A. tardans Greuter & Zahar. (), A. tardiflorum Kollmann & Schmida (), A. euboicum Rech. f. (Evvoia, Greece), A. anzalonei Brullo, Pavone & Salmeri (Italy), A. oporinanthum Brullo, Pavone & Salmeri (southern Spain and southern France), A. platakisii Tzanoud. & Kypr. (island Pondikonisi, near Crete), A. rausii Brullo et al. (NE Greece), A. aegilicum Tzanoud. (island Antikithira, Greece) and A. archeotrichon Brullo, Pavone & Salmeri (island Rhodos). The map showing distribution based on our herbarium survey is presented in Fig. 5. These species are characterized by a long-lasting vegetative phase, almost lacking the phase of dormancy, and by a spathe with erect valves, usually unilateral and partially fused, enveloping a closely fastigiate and lax inflorescence. They show also peculiar ecological requirements. They grow in habitats where they may vegetate even during the summer, tolerating very well the drought. Some species grow mainly in the underwood of evergreen or deciduous forests (A. autumnale, A. euboicum, A. rausii, A. tardiflorum and A. anzalonei), while others are found on shady rocky places or ledges (A. tardans, A. platakisii, A. archeotrichon, A. aegilicum and A. oporinanthum), and A. savii and A. telmatum prefer damp soils. 92 S. Bogdanović et al.

The autumn-flowering species of Allium sect. Codonoprasum are known only from a single or a few isolated and remote sites, indicating their relict character. Another peculiarity is their overall geographical distribution in the Mediterranean Basin. Besides their fragmented distribution, it is possible to distinguish two main speciation centers (Fig. 5): one is probably located in the western Mediterranean area, where A. oporinanthum, A. anzalonei, A. savii and A. telmatum can be found, while the second one is limited to the eastern Mediterranean area, where the remaining species grow. According to Brullo et al. (1997a), the populations occurring in these two centers also show different ploidy levels; the eastern species are all diploid (2n=16), while the western ones are tetraploid (2n=32). On the basis of morphological, karyological, ecological and chorological features, we may hypothesize that the group of the autumn-flowering species of Allium sect. Codonoprasum had probably arisen from a common diploid ancestor of the Tertiary flora. This ancestor may have been widespread in the Mediterranean Basin, growing in territories characterized by tropical aseasonal climatic conditions. The eastern diploid species are probably the oldest, and can be considered as palaeoendemics currently restricted to refugial sites. Similarly, the symmetrical and clearly diploidized karyotype of the tetraploids (none of the examined populations showed tetravalents) from the western Mediterranean suggests that they are palaeopoly- ploids, probably originating from interspecific hybridizations. In contrast to the large amount of data and knowledge gathered already for the genus Allium, no extensive investigations have been done in the Allium sect. Codonoprasum (see e.g., Friesen et al. 2006). Molecular phylogenetic studies are definitely needed to resolve taxonomic relationships and the evolutionary history of this group.

Acknowledgements The authors wish to thank Prof. Jasenka Topić (University of Zagreb) for indicating the existence of additional sites of Allium telmatum.

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Received: 11 March 2008 / Revised: 20 November 2008 / Accepted: 19 December 2008 / Published online: 31 March 2009

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