Allium Cithaeronis Bogdanović, C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), a New Species from Greece

Abstract

Bogdanović, S., C. Brullo, S. Brullo, G. Giusso del Galdo, C. M. Musarella & C. Salmeri (2011). Allium cithaeronis Bogdanović, C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), a new species from Greece. Candollea 66: 377–382. In English, English and French abstracts. A new species of Allium sect. Scorodon Koch, Allium cithaeronis Bogdanović, C. Brullo, Brullo, Giusso, Musarella & Salmeri, is described and illustrated from Greece. Its chromosome number (2n = 16), karyogram, leaf anatomy, ecology, and taxonomical relationships are examined. For some morphological features, such as exserted stamens from perigon and tuberculate lobes in the upper part of the ovary, it represents a very isolated species within the section Scorodon.

Full text

Online ISSN: 2235-3658 Candollea 66(2): 377-382 (2011) © CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 2011
Allium cithaeronis Bogdanovic
´, C. Brullo, Brullo, Giusso,
Musarella & Salmeri (Alliaceae), a new species from Greece
Sandro Bogdanovic
´,Cristian Brullo, Salvatore Brullo, Gianpietro Giusso Del Galdo,
Carmelo Maria Musarella & Cristina Salmeri
Abstract
BOGDANOVIĆ, S., C. BRULLO, S. BRULLO, G. GIUSSO DEL GALDO,
C. M. MUSARELLA & C. SALMERI (2011). Allium cithaeronis Bogdanović,
C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), a new species
from Greece. Candollea 66: 377-382. In English, English and French abstracts.
A new species of Allium sect. Scorodon Koch, Allium cithaero-
nis Bogdanović, C. Brullo, Brullo, Giusso, Musarella &
Salmeri, is described and illustrated from Greece. Its chromo-
some number (2n = 16), karyogram, leaf anatomy, ecology,
and taxonomical relationships are examined. For some mor-
phological features, such as exserted stamens from perigon
and tuberculate lobes in the upper part of the ovary, it repre-
sents a very isolated species within the section Scorodon.
Key-words
ALLIACEAE – Allium cithaeronis – Allium sect. Scorodon –
Taxonomy
Résumé
BOGDANOVIĆ, S., C. BRULLO, S. BRULLO, G. GIUSSO DEL GALDO,
C. M. MUSARELLA & C. SALMERI (2011). Allium cithaeronis Bogdanović,
C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), une nouvelle
espèce de Grèce. Candollea 66: 377-382. En anglais, résumés anglais et
français.
Une nouvelle espèce d’Allium sect. Scorodon Koch, Allium
cithaeronis Bogdanović, C. Brullo, Brullo, Giusso, Musarella
& Salmeri, est décrite et illustrée de Grèce. Le nombre chro-
mosomique (2n= 16), le caryogramme, l’anatomie foliaire,
l’écologie et les affinités taxonomiques sont donnés. En rai-
son de ses caractéristiques morphologiques particulières
comme les étamines saillantes du périgone et l’ovaire présen-
tant des lobes tuberculeux dans la partie supérieure, ce taxon
représente une espèce très isolée dans la section Scorodon.
Addresses of the authors: SB: Department of Agricultural Botany, Faculty & Agriculture, University of Zagreb, Svetosimunska 25, 10000 Zagreb. Email: sbogdanovic@agr.hr
CB, SB, GGDG, CMM: Department of Botany, University of Catania, via A. Longo 19, 95125 Catania, Italy.
CS: Department of Botanical Science, University of Palermo, via Archirafi 38, 90123 Palermo, Italy.
Submitted on November 23, 2009. Accepted on August 29, 2011. Edited by P. Bungener
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378 – Candollea 66, 2011378 – Candollea 66, 2011
Introduction
During field investigations on the orophilous flora of Greek
mountains, a very peculiar population of Allium L. was found
on the top of MontKithairon in Boeotia (C Greece). This plant
forms dense tufts localized among the dwarf shrub vegetation
colonizing the summit plateau (about 1400 m). It is a small geo-
phyte, with superficial clustered bulbs, characterized by short
spathe valves, few-flowered and compact umbel, simple and
exserted stamens, ovary with visible nectaries. On the basis of
these features, this plant can be referred to Allium sect. Scorodon
Koch, which is typically represented by species with punctiform
or very circumscribed geographical distribution. According to
literature (STEARN, 1978, 1980, 1981; GARBARI & al., 1979;
MICELI & GARBARI, 1979; PHITOS & TZANOUDAKIS, 1981;
TZANOUDAKIS, 1983, 1992; TZANOUDAKIS & VOSA, 1988;
BRULLO & TZANOUDAKIS, 1989; BRULLO & al., 1992, 1993,
1994, 2001; TZANOUDAKIS & KOLLMANN, 1991; HANELT & al.,
1992; TRIGAS & TZANOUDAKIS, 2000, FRIESEN, 2001), this sec-
tion is represented in Greece by several species, mostly endemic
and found in coastal and inner stands. From herbarium and field
investigations, the MontKithairon population is morphologi-
cally well differentiated from the other ones of Allium sect.
Scorodon s.l., hitherto known from Greece. Therefore, it is
described as a species new to science.
Material and Methods
The investigation is based on about 40 living flowering
bulbs collected on the top of MontKithairon (C Greece) and
cultivated in the Botanical Garden of Catania (Italy). The mor-
phological analysis of diacritic characters was made on fresh
material; the voucher specimens are deposited in ATH, CAT,
G, FI, UPA, and ZA. In addition, several specimens of Allium
sp. pl., preserved in many European Herbaria (B, BM, C, CAT,
FI, G, HUJ, ISTE, K, M, P, PR, PRC, RO, W, WU, ZA) and
belonging to other species of Allium sect. Scorodon, have been
examined and compared.
The leaf anatomy was studied on leaf blades of maximum
size coming from the living original collection, sectioned using
a freezing microtome.
The karyological analyses were made on mitotic plates from
root-tip cells of cultivated bulbs, pre-treated with 0.3% colchicine
water solution, fixed in ethanol-acetic (3:1) and stained accord-
ing to the Feulgen technique. Metaphases observations and chro-
mosome measurements were made using the image analysis
systems IKAROS 4.6 (Metasystem) and AXIOVISION 5.1
(ZEISS). Karyotyping was done with the software Cromolab©
1.1. (BRULLO, 2002-2003) specialized in recognizing chromo-
some pairs, ordering them by size and classifying by morphol-
ogy, and assembling the karyotype formula based on the cen-
tromere position (LE VA N & al., 1964; TZANOUDAKIS, 1983).
Results
Allium cithaeronis Bogdanović, C. Brullo, Brullo, Giusso,
Musarella & Salmeri, spec. nova (Fig. 1)
Holotype: GREECE. Sterea Ellas: MontKithairon
in stazioni cacuminali a ca. 1400 m. di quota, 38°11
63 N, 23°14 46 E, 3.VII.2007, Brullo, Musarella
50.07 (holo-: CAT; iso-: ATH, CAT, FI, G, UPA, ZA).
Bulbus 10-15 ⫻7-10 mm, tunicis exterioribus coria-
ceis, brunneis. Scapus 6-11 cm altus. Folia semi-
cylindrica, viridia, 40-60 ⫻2-2.5 mm. Valvae spathae
inaequales, major 1-2.2 cm longa, minor 0.8-1.3 cm
longa. Inflorescentia compacta, 6-15 floribus. Tepala
elliptica, 4-5 ⫻1.8-2 mm, albo-rosea, apiculata.
Stamina e perigonio exserta, filamentis albidis,
subaequalibus, 3.5-4 mm longis, antheris luteo-
pallidis, ca. 1 mm longis. Ovarium subcylindricum,
2.5-3 ⫻1.4-1.5 mm, superne dilatatum in 6 lobos
tuberculatos, tumulis nectariorum bene visillibus.
Capsula subglobosa, 3-3.5 mm diametro.
Bulb ovoid, 10-15 ⫻7-10 mm, with membranaceous
whitish inner tunics, the outer ones coriaceous, dark brown.
Stem 6-11 cm high, cylindrical, glabrous, erect, covered by the
leaf sheaths for 1/2-2/3 of its length. Leaves 4, semicylindri-
cal, glabrous, green, smooth, 4-6 cm long, 2-2.5 mm wide.
Spathe persistent, with two valves unequal, subequal or longer
than the inflorescence, the larger 6-7-nerved, 1-2.2 cm long,
the smaller 3-5-nerved, 0.8-1.3 cm long. Inflorescence com-
pact, 1-2 cm in diameter, 6-15 flowered, with subequal pedicels
4-6 mm long. Perigon campanulate, with equal tepals, white-
pinkish tinged with purple, green-purplish midrib, elliptical,
apiculate at the apex, 4-5 mm long and 1.8-2 mm wide. Sta-
mens exserted from perigon, with simple white filaments, sube-
qual, 3.5-4 mm long, below connate into an anulus 1 mm high.
Anthers pale yellow, elliptical, 1 ⫻0.8 mm, rounded at the
apex. Ovary green-yellowish, subcylindrical broadened at the
top in 6 tuberculate lobes, 2.5-3 mm long, 1.4-1.5 mm wide,
with basal mounds of nectaries well visible. Style white, 05.-
0.7 mm long. Capsule green, trivalved, subglobose, 3-3.5 mm
in diameter, with 6 small ovoid appendages on the top.
Etymology. – The specific epithet refers to “Cithaeron” the
old name of MontKithairon in Boeotia (Greece).
Habitat and Distribution. – Allium cithaeronis is localized on
the top of MontKithairon (about 1400 m), an isolated mountain
near Erythres (C Greece). This species has a flowering peak in
June, and it is member of a dwarf shrubby vegetation character-
ized by many prostrate or caespitose orophytes, such as Thymus
parnassicus Halacsy, Crepis incana Sm., Asperula rigidula
Halacsy, Cerastium candidissimum Correns, Silene radicosa Boiss.
& Heldr., Minuartia attica Vierh., Galium thymifolium Boiss. &
Heldr., Nepeta argolica Chaub., Festuca janpertii (St-Yves)
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Allium cithaeronis Bogdanovic, C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), a new species from Greece
– 379
Fig. 1 – Allium cithaeronis Bogdanovic
´
, C. Brullo, Brullo, Giusso, Musarella & Salmeri. A. Habit; B. Perigons; C. Open perigon with stamens; D. Anther ; E. Ovaries ; F. Capsules ;
G. Spathe valves.
[MontKithairon, Brullo & Musarella s.n. (CAT)] [Drawn by Salvatore Brullo]
1 cm
1 mm
5 mm
1 mm
A
BB
E
B – C – F –G
D
FG
C
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380 – Candollea 66, 2011
Markgr.-Dannen., Anthemis cretica L., Pterocephalus perennis
Coult., Achillea holosericea Sibth. & Sm., Paronychia macedonica
Chaudhri, etc. This vegetation is extremely peculiar and interest-
ing from the phytogeographical viewpoint. Unfortunately, this
vegetation appears nowadays quite degraded, especially for the
intensive overgrazing.
Karyology. – Allium cithaeronis is a diploid specie with a
somatic chromosome number 2n= 16 (Fig. 2A), which is rather
common within the genus Allium. The chromosome comple-
ment is regular and homogeneous, mostly characterized by meta-
centric pairs. Only two pairs tend towards the submetacentric
type (arm ratio exceeding 1.30) and are indicated as “msm”
according to TZANOUDAKIS (1983). Short satellites were well
evident just in few metaphases, occurring on the long arm of
one msm pair and on the short arm of the smallest metacentric
pair (Fig. 2B). The karyotype formula can be resumed as 2n=
2⫻= 16: 10 m + 2 msat + 2 msm + 2msmsat. The absolute chro-
mosome length varies from 14.40 ±1.83 µm for the longest
chromosome, to 8.15 ±1.26 µm for the shortest one, while the
relative length ranges from 8.05 ±0.37% to 4.55 ±0.17%. Many
different indices were also calculated to evaluate the symmetry
degree of the chromosome complement; all values remarked a
high homogeneity in the karyotype of A. cithaeronis (Table 1).
Leaf anatomy. – The leaf cross section of A. cithaeronis
shows a subelliptical outline. The epidermis is covered by a
well developed smooth cuticle. The stomata are numerous and
distributed along the whole perimeter. The palisade tissue is
regular and compact and consists of one-layered cylindrical
cells. The spongy tissue is compact or with larger intercellu-
lar spaces and bigger cells in the centre. In the peripheral
part of the spongy tissue several secretory canals occur. The
vascular bundles are 13, of which 7 are abaxial (4 of which
bigger) and 6 are adaxial (Fig. 3).
Discussion. – As already stated before, due to some mor-
phological features, mainly regarding spathe valves, stamens
and ovary, A. cithaeronis should be included in the A. sect.
Scorodon Koch s.l. Several authors (VALSECCHI, 1974;
CHESHMEDZHIEV, 1975; HANELT & al., 1992; FRIESEN & al.,
2006; BRULLO & al., 1995, 2008) have already highlighted that
this section represents a not natural group, since it includes
several taxa morphologically rather heterogeneous. Actually,
some species, previously referred to this section, have been
recently included in other distinct sections, such as Allium sect.
Brevispatha Valsecchi (type: A. parciflorum Viv.), Allium sect.
Cupanioscordon Cheshm. (type: A. cupanii Rafin.), Allium
sect. Eremoprasum (Kamelin) F. O. Khass., R. M. Fritsch &
N. Friesen (type: A. sabulosum Stev.), Allium sect. Avulsea
F. O. Khass. (type: A. rubellum M. Bieb.), etc. As concerns
Allium sect. Scorodon s. str., it has been attributed to the sub-
genus Polyprason Radić, while the just mentioned sections
belong to the subgenus Allium (see FRIESEN & al., 2006).
Unfortunately, we did not do any molecular analyses on A. chi-
taeronis, but we deem that, basing on morphological evidences,
this species should be, at least for the time being, included in
Allium sect. Scorodon s. str. (type: A. moschatum L.). In fact,
this section is characterized by spathe valves acuminate or
cuspidate, not much developed and usually shorter than umbel,
stamen filaments simple or with two short teeth in the
lower part and ovary with well visible nectaries. However,
A. cithaeronis shows a character rather unusual for the species
of this section, such as stamens clearly exserted from the
perigon. According to STEARN (1980) and KOLLMANN (1984),
only some Anatolian and Central Asian alliums of this section
show this morphological feature: A. microspathum Ekberg,
A. wendelboanum Kollmann, A. sabulosum Bunge, and
A. sieheanum Kollmann. Many diacritic features regarding size
and shape of the stems, leaves, spathe valves, umbels, flowers
and capsule allow to well distinguish them from A. cithaero-
nis. In particular, the ovary of the above mentioned species is
always without broadened lobes in the upper part.
Allium cithaeronis shows also some relationships with
certain species of the Allium sect. Codonoprasum Reichenb.,
particularly with those ones belonging to the A. stamineum
Boiss. group. These affinities chiefly regard the habit and the
stamens exserted from the perigon, but the spathe valves much
longer than umbel, and ovary without visible mounds of nec-
taries allow to distinguish the species of A. stamineum group
from A. cithaeronis (BRULLO & al., 2007).
Fig. 2 – Allium cithaeronis Bogdanovic
´
, C. Brullo, Brullo, Giusso, Musarella & Salmeri.
A. Metaphasic plate (2n = 16); B. Karyogram.
10 µm
10 µm
B
A
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Allium cithaeronis Bogdanovic, C. Brullo, Brullo, Giusso, Musarella & Salmeri (Alliaceae), a new species from Greece
– 381
Table 1. – Measurements and chromosome classification of Allium cithaeronis.
Absolute length (µm) Relative length %
Chrom.Long arm Short arm Total length Long arm Short arm Total length Ratio
Mean ± S.D Mean ± S.D. Mean ± S.D. Mean ± S.D. Mean ± S.D. Mean ± S.D. l/s Cl Type
1 7.54 ± 0.89 6,86 ± 0,98 14.40 ± 1,83 4.22 ± 0.15 3.84 ± 0.28 8.05 ± 0.37 1.10 47.65 m
2 7.17 ± 1,08 6,64 ± 1.01 13.81 ± 2.08 4.00 ± 0.23 4.00 ± 0.23 7.70 ± 0.45 1.08 48.06 m
3 7.21 ± 0.97 6.11 ± 1.06 13.32 ± 2.01 4.03 ± 0.21 3.40 ± 0.14 7.43 ± 0.27 1.18 45.89 m
4 7.02 ± 0.96 5.85 ± 0.83 12.87 ± 1.78 3.92 ± 0.07 3.26 ± 0.09 7.18 ± 0.16 1.20 45.43 m
5 6.60 ± 0.99 5.73 ± 0.80 12.32 ± 1.79 3.68 ± 0.02 3.20 ± 0.03 6.87 ± 0.05 1.15 46.48 m
6 6.44 ± 0.95 5.65 ± 0.97 12.09 ± 1.92 3.59 ± 0.13 3.14 ± 0.12 6.74 ± 0.24 1.14 46.72 m
7 7.06 ± 0.97 4.98 ± 0.53 12.04 ± 1.50 3.94 ± 0.18 2.79 ± 0.13 6.73 ± 0.29 1.42 41.37 msm
8 6.61 ± 1.25 4.61 ± 0.50 11.23 ± 1.66 3.67 ± 0.19 2.59 ± 0.24 6.26 ± 0.24 1.43 41.09 msm
9 5.98 ± 1.52 5.23 ± 0.99 11.21 ± 2.49 3.30 ± 0.41 2.91 ± 0.27 6.21 ± 64.00 1.14 46.65 m
10 5.62 ± 1.27 5.10 ± 1.03 10.71 ± 2.28 3.11 ± 0.35 2.84 ± 0.36 5.95 ± 0.69 1.10 47.58 m
11 6.21 ± 0.98 3.79 ± 0.61 10.00 ± 1.45 3.47 ± 0.37 2.11 ± 0.18 5.58 ± 0.38 1.64 37.88 msmsat
12 5.67 ± 0.78 3.89 ± 0.56 9.65 ± 1.38 3.18 ± 0.41 2.17 ± 0.19 5.40 ± 0.49 1.46 40.25 msmsat
13 5.23 ± 0.70 4.56 ± 0.94 9.79 ± 1.58 2.92 ± 0.18 2.53 ± 0.19 5.45 ± 0.18 1.15 46.58 m
14 4.90 ± 0.85 4.40 ± 1.00 9.30 ± 1.84 2.73 ± 0.09 2.44 ± 0.26 5.17 ± 0.35 1.11 47.33 m
15 4.77 ± 0.93 3.65 ± 0.39 8.48 ± 1.43 2.65 ± 0.11 2.05 ± 0.08 4.72 ± 0.09 1.30 43.08 msat
16 4.50 ± 0.86 3.65 ± 0.50 8.15 ± 1.26 2.50 ± 0.14 2.04 ± 0.19 4.55 ± 0.17 1.20 44.80 msat
Complement length (2n) : 179.36 ± 26.92 µm – Symmetry indices: Stebbins’ Cat.: 1A; AI : 1.09; CVCL: 15.77; CVCI: 6.92 ; A1: 0.18; A2 : 0.16; REC : 76.36; SYi: 81.90; TF% : 44.99
Therefore, based on the current knowledge on the genus
Allium in Greece, A. cithaeronis can be considered a species
taxonomically quite isolated, especially for some morpholog-
ical characteristics. Moreover, it should be stressed that A. sect.
Scorodon s. str. groups many rare species, usually well differ-
entiated and showing an extremely localized distribution, such
as A. meteoricum Halácsy, A. maniaticum Brullo & Tzanoud,
A. calamarophilon Phitos & Tzanoud., A. lagarophyllum
Brullo, Pavone & Tzanoud., A. chalkii Tzanoud. & Kollmann,
A. runemarkii Trigas & Tzanoud (BRULLO & PAVONE, 1983;
TZANOUDAKIS, 1983, 1992; TZANOUDAKIS & VOSA, 1988;
BRULLO & TZANOUDAKIS, 1989; BRULLO & al., 1992, 1993,
1994).
Acknowledgments
We are very grateful to the curators of the Herbaria B, BM,
C, CAT, FI, G, HUJ, ISTE, K, M, P, PR, PRC, RO, W, WU, ZA
for loans and help during our visits for material consultation.
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´
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Leaf cross section.
0.5 mm
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