Research Article |
Corresponding author: Cristina Salmeri ( cristinamaria.salmeri@unipa.it ) Academic editor: Lorenzo Peruzzi
© 2019 Salvatore Brullo, Cristian Brullo, Salvatore Cambria, Gianpietro Giusso del Galdo, Cristina Salmeri.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Brullo S, Brullo C, Cambria S, Giusso del Galdo G, Salmeri C (2019) Allium albanicum (Amaryllidaceae), a new species from Balkans and its relationships with A. meteoricum Heldr. & Hausskn. ex Halácsy. PhytoKeys 119: 117-136. https://doi.org/10.3897/phytokeys.119.30790
|
A new species, Allium albanicum, is described and illustrated from Albania (Balkan Peninsula). It grows on serpentines or limestone in open rocky stands with a scattered distribution, mainly in mountain locations. Previously, the populations of this geophyte were attributed to A. meteoricum Heldr. & Hausskn. ex Halácsy, described from a few localities of North and Central Greece. These two species indeed show close relationships, chiefly regarding some features of the spathe valves, inflorescence and floral parts. They also share the same diploid chromosome number 2n =16 and similar karyotype, while seed testa micro-sculptures and leaf anatomy reveal remarkable differences. There are also several morphological features that allow them to be differentiated at specific level. The inclusion of both species into a newly described section Pseudoscorodon of the subgen. Allium is proposed. An analytic key to the species, included in the new section, is also provided.
Allium, Amaryllidaceae, Albania, chromosome, new section, taxonomy
One of the richest and largest genera of Monocotyledons is Allium L. and it is almost exclusively widespread in the northern hemisphere, where it is represented by ca. 1200 taxa (
In the context of cytotaxonomical research on the genus Allium in the Mediterranean territories, especially regarding the subgen. Allium (
Plant morphology was analysed on 20 living mature plants. Qualitative and quantitative morphological characters, considered as diagnostic in Allium, were analysed and scored (Table
For the karyological study, living bulbs were collected and potted at the Botanical Garden of Catania University. Root tips were pre-treated with 0.3% (w/v) colchicine water solution for 3 h at room temperature and then fixed overnight in fresh Farmer’s fixative (3:1 v/v, absolute ethanol: glacial acetic acid). Root tips were hydrolysed in 1N HCl at 60 °C for 7 min, washed and stained with Feulgen for 1 h. Microphotographs of good quality metaphase plates were taken with a Zeiss Axioskop2 light microscope equipped with an Axiocam MRc5 high resolution digital camera. Chromosome number and karyotype details were analysed from 10 well spread metaphase plates from 5 individuals, the mean values being used for the karyotype characterisation. Metaphase chromosomes were measured using the image analysis system Zeiss Axiovision 4.8, while karyotyping was performed by CROMOLAB 1.1 software
Leaf anatomy was studied on living materials coming from the type locality and cultivated in the Botanical Garden of Catania University. Leaf blades of maximum size, in their optimal vegetative development, usually before the flowering stage, were taken from the middle part and fixed in Carnoy. Leaf cross sections were double stained with ruthenium red and light green, analysed and photographed with a light microscope (Zeiss Axioskop2 and Axiocam MRc5 digital camera).
Seed testa micro-morphology was analysed on mature and dry material taken from individuals coming from the type locality, using a scanning electron microscope (SEM) Zeiss EVO LS10, according to the protocol reported by
Characters | A. albanicum | A. meteoricum |
---|---|---|
Bulb size (mm) | 8–10 × 5–10 | 10–14 × 8–12 |
Bulb outer coat colour | brownish | blackish-brown |
Stem height (cm) | 14–28(-30) | 10–25 |
Stem diameter (mm) | 1 | 1–1.2 |
Stem coverage by leaf sheaths | 1/4 | 1/2 |
Leaf number | 3 | 3–4 |
Leaf length (cm) | up to 10 | up to 12 |
Spathe valves length (mm) | subequal, 8–12 | unequal, 7–11 |
Spathe valve appendage length (mm) | 1–2.5 | 1–4 |
Spathe valves arrangement | fused up to 1/2 | free |
Larger spathe valve nerves (no.) | 3–5 | 5 |
Smaller spathe valve nerves (no.) | 3 | 3–5 |
Pedicel length (mm) | 6–25 | 6–15 |
Tepal colour | white tinged with pink | purplish-pink |
Tepal midvein colour | greenish-purple | purplish |
Tepal length (mm) | 5.5–6.5 | 6–7.5 |
Tepal apex | eroded | rounded |
Stamen filament colour | yellowish above, white below | white |
Outer stamen filament length (mm) | 1.7–2.1 | 2.7–3.3 |
Inner stamen filament length (mm) | 2.5–3.2 | 3.5–4 |
Anther colour | greenish-pale yellow | yellow |
Anther apex | rounded | apiculate |
Annulus height (mm) | 0.5–0.6 | 0.7–0.9 |
Ovary colour | yellow | green |
Ovary apex | slightly wrinkled | smooth |
Ovary nectariferous pores height | about 1/2 ovary | about 1/4 ovary |
Style length (mm) | 2.7–2.8 | 1.5–2 |
Capsule length (mm) | 4–4.5 | 3–3.5 |
Capsule shape | subglobose-obovate | subglobose |
Seed size (mm) | 3.5–4.0 × 2.4–2.5 | 2.2–2.5 × 1.9–2.0 |
Karyomorphometric parameters and symmetry indices for Allium albanicum and A. meteoricum. Mean values were calculated from 10 good metaphase plates from individuals of the type locality.
Allium albanicum | ||||||||
Pairs | LA (µm) | SA (µm) | TAL (µm) | TRL% | AR | CI | CA | Type |
I | 5.00 ± 1.43 | 3.74 ± 1.07 | 8.74 ± 2.48 | 7.87 ± 0.59 | 1.34 | 42.82 | 0.14 | msm |
II | 4.52 ± 0.94 | 3.38 ± 1.05 | 7.91 ± 1.71 | 7.16 ± 0.34 | 1.34 | 42.80 | 0.14 | msm |
III | 4.03 ± 0.87 | 3.48 ± 0.73 | 7.53 ± 0.58 | 6.84 ± 0.34 | 1.16 | 46.20 | 0.07 | m |
IV | 4.67 ± 0.99 | 1.99 ± 0.38 | 6.79 ± 1.31 | 6.19 ± 0.27 | 2.35 | 29.30 | 0.40 | smsat |
V | 3.56 ± 0.40 | 3.09 ± 0.34 | 6.78 ± 0.66 | 6.26 ± 0.59 | 1.15 | 45.57 | 0.07 | msat |
VI | 3.48 ± 0.77 | 2.93 ± 0.61 | 6.44 ± 1.34 | 5.86 ± 0.42 | 1.19 | 45.56 | 0.09 | m |
VII | 3.33 ± 0.75 | 2.10 ± 0.38 | 5.43 ± 0.38 | 4.91 ± 0.81 | 1.58 | 38.69 | 0.23 | msmsat |
VIII | 2.87 ± 0.42 | 2.33 ± 0.42 | 5.33 ± 0.77 | 4.90 ± 0.47 | 1.23 | 43.68 | 0.10 | msat |
TCL: 109.88 ± 21.7 µm; MCL: 6.87 ± 1.2 µm; d-value: 16.83; DRL%: 3.28; S%: 57.95;MAR: 1.37; MCI: 41.83; CvCL: 17.12; CvCI: 13.00; MCA: 15.63 | ||||||||
Allium meteoricum | ||||||||
Pairs | LA (µm) | SA (µm) | TAL (µm) | TRL% | AR | CI | CA | Type |
I | 4.05 ± 0.21 | 3.23 ± 0.30 | 7.27 ± 0.31 | 7.78 ± 0.02 | 1.26 | 44.35 | 0.11 | m |
II | 3.87 ± 0.44 | 3.06 ± 0.73 | 6.94 ± 1.01 | 7.42 ± 0.35 | 1.26 | 44.19 | 0.12 | m |
III | 3.79 ± 0.43 | 2.74 ± 0.23 | 6.53 ± 0.34 | 6.99 ± 0.37 | 1.38 | 41.98 | 0.16 | msm |
IV | 4.15 ± 0.16 | 1.69 ± 0.11 | 6.16 ± 0.27 | 6.59 ± 0.29 | 2.45 | 27.49 | 0.42 | smsat |
V | 2.90 ± 0.35 | 2.58 ± 0.11 | 5.48 ± 0.11 | 5.87 ± 0.12 | 1.13 | 47.06 | 0.06 | msat |
VI | 2.90 ± 0.23 | 2.50 ± 0.34 | 5.40 ± 0.57 | 5.78 ± 0.61 | 1.16 | 46.27 | 0.07 | m |
VII | 2.66 ± 0.11 | 1.85 ± 0.11 | 4.52 ± 0.23 | 4.83 ± 0.24 | 1.43 | 41.07 | 0.18 | msm |
VIII | 2.42 ± 0.23 | 2.02 ± 0.42 | 4.44 ± 0.57 | 4.74 ± 0.51 | 1.20 | 45.45 | 0.09 | m |
TCL: 93.48 ± 21.7 µm; MCL: 5.84 ± 1.06 µm; d-value: 14.13; DRL%: 3.5; S%: 55.31;MAR: 1.36; MCI: 42.21; CvCL: 18.16; CvCI: 14.48; MCA: 15.21 |
Allium meteoricum auct. fl. Albania non Halacsy, Consp. Fl. Graec. 3(1): 250. 1904, Syn.
ALBANIA. Devoli river, near Berat, serpentines, ca. 700 m elev., 40°43'12.00"N, 20°32'18.00"E, 26 June 2017, S. Cambria s.n. (Holotype: CAT; Isotypes: CAT, FI, G).
Allio meteoricum similis sed bulbis minoribus tunicis exterioribus brunneis, scapo ad 1/4 longitudinem vaginis foliorum tecto, spathae valvis in dimidio inferiore connatis, appendice usque ad 2,5 mm longa, majore 3–5 nervata, minore 3 nervata, tepalis albo-roseis, minoribus, apice erosis, filamentis staminorum minoribus, luteis superne, annulo breviore, antheris viridulis- pallide luteis, apice rotundatis, ovario luteo leviter apice rugoso, poris nectariferis majoris, capsula majore subgloboso-obovata, differt.
Bulb ovoid, 8–10 × 5–10 mm, with outer tunics coriaceous, brownish, the inner membranous, whitish. Stem 14–28(30) cm tall, cylindrical, flexuous, 1–1.5 mm in diameter, glabrous, erect, covered for 1/4 of its length by the leaf sheaths. Leaves 3, rather flat, glabrous, green, ribbed, up to 10 cm long and 1–2.2 mm wide, denticulate at margins. Spathe persistent, with 2 valves subequal, 8–12 mm long, shorter than the inflorescence, fused to half of their length, with an appendage 1–2.5 mm long, the larger 3–5-nerved, the smaller 3-nerved. Bostryces 12. Inflorescence laxly hemispheric, 2–3 cm in diameter, many flowered, with unequal pedicels 6–25 mm long. Perigon cylindrical-urceolate, with tepals of equal length, white tinged with pink, mid-vein greenish-purple, the inner ones linear-elliptical, the outer ones sublanceolate, rounded and slightly eroded at the apex, 5.5–6.5 mm long and 1.7–2 mm wide. Stamens included, with simple filament yellowish above and whitish below, the outers 1.7–2.1 mm long, the inners 2.5–3.2 mm long, below connate into an annulus 0.5–0.6 mm high. Anthers greenish-pale yellow, elliptical, 1–1.1 × 0.6 mm, rounded at the apex. Ovary subglobose-ovoid, yellow, slightly wrinkled at the apex, 1.5–1.7 × 1.4–1.7 mm, with large nectariferous pores, long about half the ovary. Style white, 2.7–2.8 mm long, stigma capitate. Caspule trivalved, subglobose-obovate, 4–4.5 mm, with evident nectariferous pores.
Flowering and fruiting from June to July.
The epithet refers to the Latin “Albanicum”, coming from Albania, the country where the species grows.
The investigated specimens of A. albanicum from the type locality revealed a diploid chromosome number with 2n = 16. The karyotype obtained from somatic metaphase plates (Fig.
The leaf cross section of A. albanicum shows a flat outline, with some dorsal ribs. The epidermis is formed by small cells covered by a well-developed cuticle externally more thickened. Stomata are numerous and distributed along the whole leaf perimeter. The palisade tissue is regular and compact, arranged in one layer of long cylindrical cells, more developed on the adaxial face. The spongy tissue is rather compact and slightly lacunose, in the peripheral part many secretory canals occur. The maximum number of vascular bundles is 20, 11 of which are very small and are localised on the adaxial face, while on the abaxial face, there is one large central vascular bundle and 4 smaller ones for each side (Fig.
Comparison of karyotype morphometric data between Allium albanicum and A. meteoricum. A Variation of centromeric index for each chromosome pair B Variation of the main karyomorphometric parameters and symmetry indices (LC longest chrom., SC shortest chrom.; MCL mean chromosome length; MLA mean long arm; MSA mean short arm; other abbreviations see Table
As emphasised by numerous authors (
SEM micrographs of the seed coat of Allium albanicum. A Seed (dorsal face, 30×) B Seed (ventral face, 30×) C Seed coat (central part of dorsal face, 600×) D Seed coat (central part of ventral face, 600×) E Seed coat (central part of dorsal face, 1200×) F Seed (central part of ventral face, 1200×). Photos from material of type locality (CAT).
SEM micrographs of the seed coat of Allium meteoricum. A Seed (dorsal face, 30×) B Seed (ventral face, 30×) C Seed coat (central part of dorsal face, 600×) D Seed coat (central part of ventral face, 600×) E Seed coat (central part of dorsal face, 1200×) F Seed (central part of ventral face, 1200×). Photos from material of type locality (CAT).
The investigated population of A. albanicum, previously reported as sub A. meteoricum (
Phenological features of Allium albanicum and A. meteoricum. A Growing habitat of A. albanicum in the locus classicus (Albania) B Individuals of A. albanicum from the locus classicus C A. albanicum cultivated material in Botanical Garden of Catania D Leaf of A. albanicum, cultivated material E, F Individual of A. meteoricum, from Meteora (Greece). Photos by S. Cambria.
ALBANIA. In humidis collinis serpentinum ad Renci distr. Scutari, 11 June 1897, Baldacci 85a (BM, G); In humidis collinis serpentinii ad Renci distr. Scutari, June 1897, Baldacci 355 (WU); Nordost Albanien, auf Felsen in der subalp. Region des Pastrik ca. 1200 m elev., 31 July 1914, Dorfler 593 (WU); Nord Albania, Umgebung von Shkodra Abhänge des kleinen Bordans alt. Serpentin, 8 June 1916, Janchen s. n. (WU); Hasi Pastrik an Felsen des westlichen Ausläufers, ca. 1200 m elev., 22 July 1918. Dorfler 908 (BM, K, W, WU).
GREECE, Thessalia superior in collibus circa monasteria Meteora supra Kalabaka (Aeginium veterum), substrata diluviali e saxis conglomeratis, 15/16 July 1885 Heldreich s. n., sub Allium meteoricum Heldreich & Haussknecht sp. nova (WU Herbarium Halacsy); this specimen, already quoted by
For its general habit and some features such as flat leaves, spathe valves very short, 3–5 nerved, briefly appendiculate, umbel laxly subglobose, perigon cylindrical-urceolate, stamens not exserted, ovary with evident nectariferous pores, the populations of A. albanicum were previously referred to as A. meteoricum (Halacsy 1904,
In light of in-depth taxonomical investigations carried out on living and herbarium material, the analysed Albanian populations are well differentiated from those of A. meteoricum coming from the locus classicus, formerly studied by
According to previous research data (
Based on literature (
In particular, the sect. Scorodon s.str., typified by A. moschatum L., now belongs to the subgen. Polyprason Radić, which groups rhizomatous species (
Allium obtusiflorum DC in
Bulbus solitarius vel bulbilliferous, sine basali rhizomate, folia glabra vel pilosa, numquam filiformes, plerumque spathae valvae umbella breviores, persistentes, saltem 3-nervatae, staminum filamenta complanata inferne, interiores saepe 1–2 cuspidibus praedita, ovarium nectariferis poris bene evolutis, plica membranacea praeditis, partim nectariferum porum tegente.
Bulb solitary or bulbilliferous, leaves glabrous to hairy, never thread-like, spathe valves persistent and usually shorter than the inflorescence, at least 3-nerved, stamen filaments flattened and widened in the lower part, the inner ones often uni-bicuspidate, ovary with well-developed nectariferous pores, bordered by a membranous plica, partly covering the nectariferous pore.
Based on current knowledge (
Based on the descriptions and related iconographies, all of these species share the set of discriminant features that characterise the new section and distinguish it very well from all the known sections of the subgenus Allium (
In order to highlight the morphological similarities and differences amongst the species of the new section, the following analytic key is provided.
1 | Leaves hairy | 2 |
− | Leaves glabrous or subglabrous | 7 |
2 | Tepals and stamen filaments greenish-yellow | 3 |
− | Tepals and stamen filaments white to pink or purplish | 5 |
3 | Tepals thickened at the base, 2.5−3.2 mm wide. Inner stamen filaments with 1−2 cusps at the base. Capsule 5−5.5 mm long | A. valdecallosum |
– | Tepals not thickened at the base, 1.5−2.5 mm wide. Stamen filaments all simple. Capsule 3.5−4(4.5) mm long | 4 |
4 | Leaf blade 0.5−1 mm wide. Umbel fastigiate, 3−4.5 mm long. Stamen filaments exserted | A. chrysonemum |
− | Leaf blade 1.5−2 mm wide. Umbel expanded, 4−6 mm long. Stamen filaments included | A. rouyi |
5 | Spathe valves much shorter than umbel. Perigon 4.5−6 mm long. Stamen filaments all simple, exserted | A. reconditum |
− | Spathe valves slightly shorter than umbel (sometimes subequal). Perigon 6−8 mm long. Stamen filaments included, the inner ones with two cusps in the middle part | 6 |
6 | Leaves almost totally densely hairy. Perigon cup-shaped, white to white-pink with tepals 8−8.2 mm long and 3.2−3.3 mm wide | A. seirotrichum |
− | Leaves sparsely hairy in the sheath. Perigon cylindrical-urceolate, pink-lilac with tepals 5−7 mm long and 1−2 mm wide | A. trichocnemis |
7 | Tepals 3.5−5 mm long | 8 |
− | Tepals more than 5 mm long | 12 |
8 | Outer bulb tunics breaking into parallel fibres, pale brown. Inner stamen filaments with two basal cusps | A. thessalicum |
− | Outer bulb tunics coriaceous, brown to dark brown. Inner stamen filaments without basal cusps | 9 |
9 | Spathe valves free. Umbel with flexuous pedicels | 10 |
− | Spathe valves connate at the base. Umbel with erect or suberect pedicels | 11 |
10 | Leaves 4−6. Inflorescence dense and compact. Spathe valves both 3-nerved, 5−7 mm long. Anthers purple-violet. Ovary with apical purplish-brown spots. Nectariferous pores about ½ of the ovary length | A. obtusiflorum |
− | Leaves 2−4. Inflorescence lax. Spathe valves (1)2−4-nerved, 5−20 mm long. Anthers yellowish. Ovary without apical spots. Nectariferous pores about 1/10th of the ovary length | A. maniaticum |
11 | Leaves (3)4−5. Pedicels 2−8 mm long. Tepals whitish-pink. Anthers purplish-violet. Ovary 1.3−2 mm log, with a purplish-brown apical spot | A. runemarkii |
− | Leaves 3. Pedicels 5−20 mm long. Tepals purplish-pink. Anthers pale yellow. Ovary 1.2−1.3 mm long, without apical spot | A. erythraeum |
12 | Tepals linear, 5−5.5 × 0.8−1 mm. Ovary 1–1.2 mm long. Capsule max. 3 mm long | A. franciniae |
− | Tepals linear-elliptical to sublanceolate or oblong-elliptical, 5.5−8 × 1.7−2.5 mm. Ovary 1.5−2 mm long. Capsule 3−5 mm long | 13 |
13 | Spathe valves unilateral, long fused. Inflorescence fastigiate and unilateral | 14 |
− | Spathe opposite, free or partially fused. Inflorescence expanded, never unilateral | 15 |
14 | Stem 15−25 cm long. Inflorescence 12−20-flowered. Tepals purplish at the apex. Ovary 1.6−1.8 mm long | A. rhodiacum |
− | Stem 5−12 cm. Inflorescence 2−12-flowered. Tepals concolorous. Ovary 1.2−1.5 mm long | A. chalkii |
15 | Stem flexuous. Spathe valves subequal, fused to half of their length. Tepals white-pink. Ovary with very large nectariferous pores | 16 |
− | Stem rigid. Spathe valves unequal, free. Tepals purplish-pink. Ovary with small nectariferous pores | 17 |
16 | Stem 3-leaved, 14−28(30) cm tall. Spathe valves 8−12 mm long, 3−5-nerved. Style 2.7–2.8 mm long | A. albanicum |
− | Stem 1-leaved, 9−15 cm tall. Spathe valves 5−7 mm long, 1−3-nerved. Style 1 mm long | A. lagarophyllum |
17 | Outer bulb coats blackish-brown. Anthers yellow. Staminal annulus 0.7−0.9 mm high. Capsule 3−3.5 mm long | A. meteoricum |
− | Outer bulb coats purplish-brown. Anthers purplish-pink. Staminal annulus 1.5 mm high. Capsule 4−5 mm long | A. grosii |