Download Charybdis glaucophylla (Asparagaceae), a new species from Sardinia

Phytotaxa 69: 16–26
(2012)
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Copyright © 2012 Magnolia Press
ISSN 1179-3155 (print edition)
PHYTOTAXA
Article
ISSN 1179-3163 (online edition)
Charybdis glaucophylla (Asparagaceae), a new species from Sardinia
GIANLUIGI BACCHETTA1*, SALVATORE BRULLO2, SAVERIO D’EMERICO3,
CRISTIANO PONTECORVO1 & CRISTINA SALMERI4
1
Centro Conservazione Biodiversità (CCB), Dipartimento di Scienze della Vita e dell’Ambiente, Università degli Studi di Cagliari,
Viale S. Ignazio da Laconi 11-13, I 09123 Cagliari, Italy, e-mail: [email protected]
2
Dipartimento di Scienze Biologiche, Geologiche ed Ambientali, Università degli Studi di Catania, Via A. Longo 19, I 95125 Catania,
Italy, e-mail: [email protected]
3
Dipartimento di Biologia e Patologia Vegetale, Università di Bari, Via Orabona, 4. I 70125 Bari, Italy.
4
Dipartimento di Biologia ambientale e Biodiversità, Università degli Studi di Palermo, Via Archirafi 38, I 90123 Palermo, Italy,
e-mail: [email protected]
*
author for correspondence
Abstract
Charybdis glaucophylla (Asparagaceae), a new species from Sardinia, is described and illustrated. It is a diploid species
with 2n = 20 chromosomes growing along the rocky coast, sandy dunes and mountain top in the south-western part of the
island. Within the genus, this species results taxonomically well isolated and differentiated due to relevant morphological
and phenological features, such as the leaves wide and short, very rigid and glaucous-pruinose, as well as the very late
foliation (winter). It shows only some relationships with C. pancration for the whitish bulb tunics and the diploid
chromosome complement, and with C. maura and C. aphylla due to the glaucous leaves.
Key words: endemic flora, karyology, Mediterranean Basin, Sardinia, taxonomy, Urginea
Introduction
The genus Charybdis Speta (1998: 58) is typified by Scilla maritima Linnaeus (1753: 308) [≡ Urginea
maritima (L.) Baker (1873: 221); Charybdis maritima (L.) Speta (1998: 60)]. This species is reported
throughout the Mediterranean area by the main floras (Maire 1958, Chouard & Guinochet 1978, El-Gadi
1978, McNeill 1980, Pignatti 1982, Edmondson 1984, Meikle 1985, Feinbrun-Dothan 1986, Pastor 1987,
Amaral Franco & Rocha Alfonso 1994, Juan 2002, Boulos 2005, Tison 2007). C. maritima indeed represents
a species complex well differentiated in several morphologically, karyologically and genetically distinct
populations, as many authors highlighted (Martinoli 1949, Battaglia 1957a, 1957b, 1964, Maugini 1953,
1956, 1960, Maugini & Bini Maleci 1974, Speta 1980, 1998, 2001, Pfosser & Speta 1999, 2004, Boscaiu et al.
2003, Rosselló et al. 2005).
Typical populations of C. maritima s. str. are hexaploid (2n = 60) and occur in the Iberian peninsula and
NW Morocco (Talavera et al. 1995, Pfosser & Speta 2004). They are characterized by bulb whitish tunics,
green, very long, narrowly lanceolate and acute leaves, tepals with purplish midrib. Tetraploid populations (2n
= 40) are usually attributed to C. hesperia (Webb & Berthelot 1848: 339) Speta (1998: 60), C. numidica
(Jordan & Fourreau 1869: 1) Speta (1998: 60) and C. aphylla (Forsskal 1775: 209) Speta (1998: 60) (Pfosser
& Speta 2004). The latter taxa are geographically and morphologically well distinct; C. hesperia is restricted
to the Canary Islands and N Morocco and shows bulb tunics white-greenish and roots whitish, leaves green,
very long, narrowly lanceolate and acute, tepals with dark purplish midrib. C. numidica is distributed in the
CW Mediterranean area and shows reddish bulb tunics and whitish roots, green-glaucous, wide and oblong
16 Accepted by Lorenzo Peruzzi: 4 Oct. 2012; published online in PDF: 16 Oct. 2012
leaves, tepals with green midrib. Finally, C. aphylla is limited to the E Mediterranean area and exhibits whitereddish to brown-red bulb tunics and whitish roots, glaucous, wide, broadly lanceolate leaves, tepals with
green to purplish midrib. Diploid populations (2n = 20) are only known for CW Mediterranean territories and
they are mostly referred to C. pancration (Steinheil 1836: 279) Speta (1998: 60) (Boscaiu et al. 2003) and C.
maura (Maire 1923: 158) Speta (1998: 60) (Battaglia 1964). Charybdis pancration, which is mainly
distributed in Sicily, Calabria, Puglia, Malta, Lampedusa, Tunisia (Pfosser & Speta 2004) and Cabrera
(Rosselló et al. 2005), is differentiated by bulb tunics green-whitish and roots reddish, leaves green, long and
wide, lanceolate, tepals with purplish midrib. On the contrary, C. maura is endemic to N Africa and has greenwhitish bulb tunics and whitish roots, glaucous, very long, narrowly lanceolate and acute leaves, tepals with
purplish midrib. It should be noted that the green tinge of bulb tunics in these species is probably due to the
large epigeal growth of bulbs.
During field investigations in Sardinia, very peculiar populations of Charybdis were found in the southwestern part of the island. They are characterized by whitish bulb tunics and roots, very wide, short, glaucouspruinose and rigid leaves, oblanceolate and apiculate at apex, tepals with pink-purplish midrib. There are also
relevant peculiarities in phenology, since these populations show early flowering and a long dormant period
between flowering and leaf sprout. All studied populations are diploid with 2n = 20, a chromosome count only
found in few taxa or populations within genus Charybdis (Boscaiu et al. 2003). Based on such remarks, the
Sardinian populations can be treated as a species new to science, named C. glaucophylla.
Materials and methods
Morphological investigations were carried out on living specimens sampled in 4 Sardinian localities (see Fig.
4) and cultivated in the Botanical Gardens of Cagliari and Catania. In addition, herbarium specimens from
CAG and CAT were also examined for taxonomic comparison. The material was examined under a Zeiss
Stemi SV 11 Apo stereomicroscope at 6–66x magnification.
TABLE 1. Karyomorphometric parameters and symmetry indices for Charybdis glaucophylla.
Mean values comes from 10 good metaphase plates from different individuals of the type locality.
Pair n.
TAL (µm)
TRL
AR
CI
Type
I
10.27 ± 1.1
9.43 ± 0.1
5.62
15.10
st
II
9.17 ± 0.8
8.40 ± 0.4
5.57
15.45
st
III
6.00 ± 0.8
5.49 ± 0.2
4.22
19.17
st
IV
4.60 ± 0.5
4.21 ± 0.2
3.45
22.46
st
V
4.52 ± 0.5
4.15 ± 0.03
2.91
25.58
smsat
VI
4.07 ± 0.2
3.73 ± 0.1
3.69
21.31
st
VII
4.17 ± 0.62
3.81 ± 0.09
2.85
26.00
sm
VIII
3.98 ± 0.51
3.64 ± 0.10
2.32
30.13
sm
IX
3.97 ± 0.32
3.64 ± 0.08
1.48
40.34
msm
X
3.82 ± 0.43
3.49 ± 0.06
2.63
27.51
sm
TKL : 109.10 ± 2.3; SC–LC range: 3.77–10.37 µm; LC/SC: 2.70;
CVCL: 42.99; CVCI: 30.90
Abbreviations: TAL = total absolute length; TRL = total relative length; AR = arm ratio; CI = centromeric index; Type = chromosome
nomenclature according to Levan et al. (1969) and Tzanoudakis (1983); sat = satellited; TKL = total karyotype length; LC = longest
chromosome; SC = shortest chromosome.
Karyological analyses were performed on mitotic plates from root tip cells of cultivated bulbs, pre-treated
with 0.3% (W/V) colchicine water solution at room temperature, fixed in Carnoy solution for 12 hours, then
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
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hydrolyzed with 1N HCl for 8 min at 60°C and stained according to Feulgen technique. Micrographs of good
quality metaphase plates were taken using a Zeiss Axioskop 2 microscope equipped with a monochrome CCD
camera and an Axiocam MRc5 high resolution digital camera. The somatic chromosome number and
karyotype details were studied in 10 well prepared metaphase plates from different individuals and localities,
the mean values were used for the analysis. Metaphase handlings and chromosome measures were made using
the image analysis systems IKAROS 4.6 (Metasystem) and Zeiss Axiovision 4.6. Karyotyping was worked
out by a specific software Cromolab© 1.1 (Brullo 2002) for the recognition of homologues, couple ordering,
chromosome classification and karyotype formula based on the centromere position (Levan et al. 1964,
Tzanoudakis 1983). The karyotype symmetry was determined calculating both CvCI and CvCL parameters
(Paszko 2006).
Taxonomic Treatement
Charybdis glaucophylla Bacch., Brullo, D'Emerico, Pontec. & Salmeri, sp. nov. (Fig. 1, 2)
Affinis Charybde pancratione sed bulbo 5–8 x 6–10 cm, tunicis et radicibus albis, foliis glauco-pruinosis, numero
(5–)6–9 varians, oblanceolatis, (16–)22–28(–34) x (3–)4.5–8(–10) cm, inflorescentia (10–)20–40(–57) cm longa ad
150–200 flores composita, perigonii lobis 7–7.5 mm longis, ovario 2.6–2.8 mm longo, stigmate capitati differt.
Type:—ITALY. Sardinia: Isola di San Pietro: Cala Vinagra, Carloforte, 63 m a.s.l., 38° 09’ 47,49’’N, 8° 14’ 37,75’’E,
19 July 2004, G. Bacchetta & C. Pontecorvo s.n. (holotype CAT!; isotypes CAG!, CAT!).
Bulb ovoid, 5–8 × 6–10 cm, with outer tunics coriaceous and brown in colour, the inner ones whitish. Leaves
(5–)6–9 in number, glaucous-pruinose, rigid, oblanceolate, (16–)22–28(–34) × (3–)4.5–8(–10) cm, obtuse to
acute, cucullate and apiculate at the apex. Stem 28–35 cm long, greenish, tinged with violet in the upper part.
Raceme cylindrical, greenish, (10–)20–40(–57) cm long, with 150–200 flowers. Pedicels erect-patent, 12–18
mm long, longer than perigon, extending in fruiting plants. Flower buds white, sometimes tinged with pink,
7–8 mm long. Perigon white, stellate, 15–16 mm in diameter; lobes 7–7.5 × 3.4–3.8 mm, oblong to oblongelliptic, the inner ones rounded, the outer ones obtuse, midrib purplish. Stamens subequal or shorter than the
perigon; anthers greenish, 3.0–3.2 mm long; filaments white, subulate, 3.5–4.2 mm long. Ovary ellipsoid,
green, 2.6–2.8 × 1.9–2 mm; style white, 2.2–3.2 mm long; stigma capitate, white, papillose. Fruiting raceme
linear-cylindrical. Capsule trigonous, ellipsoid, 8.5–10 × 6–7.5 mm, truncate at the base. Seeds oblong, black,
shining, 4.3–5 × 2–2.3 mm.
Phenology:—Flowering late July to August, fruiting August to September, foliation January to May.
Karyology:—The somatic chromosome number of Charybdis glaucophylla (Fig. 3A, B) was found to be
2n = 2x = 20 in all studied samples. The karyotype (Fig. 3C) consists of 5 subtelocentric pairs, 2 of which
much longer than others, 4 submetacentric pairs, one of which provided with a very long satellite on the short
arm, and 1 meta-submetacentric pair (arm ratio 1.5). Chromosome measures and symmetry indices (Tab. 1)
show a relatively high degree of karyotype asymmetry both in chromosome relative size (big pairs vs. small
pairs) and in centromere position (many subterminal pairs). Chromosome length varies from 10.37 ± 1 µm of
the longest chromosome to 3.77 ± 0.4 µm of the shortest one, while the relative length ranges from 9.54% ± 1
to 3.45% ± 0.2. The karyotype formula can be expressed in 2n = 2x = 20: 10 st + 6 sm + 2 smsat + 2 msm.
Habitat:—This species is typically linked to rocky and sandy places preferably next to sea and strongly
windy (Mistral), where it grows on different substrata, like sand, limestone, metamorphic and volcanic rocks.
Usually, it is a member of subhalophilous plant communities characterized by some endemic species linked to
rocky coasts, such as Bellium crassifolium Moris (1827: 26), Hyoseris taurina (Pampanini 1948: 138)
Martinoli (1953: 257), Limonium sulcitanum Arrigoni (1981: 233). On sandy dunes this species grows
together with various psammophytes [Ephedra distachya Linnaeus (1753: 1040), Genista arbusensis
Valsecchi (1984: 291), Scrophularia ramosissima Loiseleur-Delongchamps (1807: 381)], hence it appears to
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BACCHETTA ET AL.
be well adapted to the marine salt spray. A small population was found in a mountain stand on metamorphic
rocks at ca. 1000 m elevation. Here it grows in garigues characterized by Teucrium marum Linnaeus (1753:
564), Genista sulcitana Valsecchi (1986: 193) and Helichrysum microphyllum (Willdenow 1803: 1863)
Cambessedes (1827: 272) subsp. tyrrhenicum Bacchetta, Brullo & Giusso (in Angiolini et al. 2005: 272).
FIGURE 1. Diagnostic features of Charybdis glaucophylla. A. Habit. B. Inflorescence. C. Leaves. D. Inflorescence detail. E. Fruits.
Illustration by Salvatore Brullo based on Bacchetta & Pontecorvo s.n. (CAT).
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
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FIGURE 2. A. Flower, side view. B. Flower, upper view. C. Perigon with stamens. D. Bud. E. Anther. F. Ovary. G. Stigma. H. Fruit.
I. Seed. Illustration by Salvatore Brullo based on Bacchetta & Pontecorvo s.n. (CAT).
Distribution:—On the basis of our field investigations, this species is quite rare and scattered just along
the south-western part of Sardinia (Fig. 4). Currently, only four populations are known, three of them [S.
Pietro (Carloforte), Pranu Sartu (Buggerru and Iglesias) and Monte Linas (Gonnosfanadiga)] occur in rocky
habitats and the other one [Scivu (Arbus)] is typical of sandy dunes.
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FIGURE 3. Chromosome complement (2n = 20) of Charybdis glaucophylla. Mitotic metaphase plate from S. Pietro (A) and Pranu
Sartu (B); arrows indicates satellited chromosomes. C. Idiogram.
Etymology:—The name refers to the characteristic waxy and greyish-blue leaves (from Greek glaucos =
greyish-blue and phyllon = leaf).
Conservation:—Despite the four populations are threatened by grazing, population decline was not
observed. However, due to the population small size and the risk that the threat level can quickly increase (i.e.
human activity or stochastic events), following the IUCN Red List Categories and Criteria (2001) we suggest
that Charybdis glaucophylla should be treated as Vulnerable VU = D2, as the total number of mature plants
ranges from 500 to 1000, distributed in less than 20 km2.
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FIGURE 4. Distribution map of Charybdis glaucophylla populations. A. S. Pietro (Carloforte, CI). B. Pranu Sartu (Buggerru-Iglesias,
CI). C. Scivu (Arbus, VS), D. Monte Linas (Gonnosfanadiga, VS).
Observations:—Historically, the Mediterranean populations of Urginea Steinheil (1834: 321)
characterized by big bulb, large leaves and very long inflorescence were attributed to U. maritima (Speta
1998), while the plants with small bulb and leaves, and short inflorescence were referred to U. undulata
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BACCHETTA ET AL.
(Desfontaines 1798: 300) Steinheil (1834: 330), if provided with flat and undulate leaves, or to U. fugax
(Moris 1827: 46) Steinheil (1834: 328), if leaves are filiform. On the basis of genetic analyses, only U. fugax
is actually included in the genus Urginea, while the remaining species are included within Charybdis (Speta
1998, 2001). Karyological studies on these species revealed distinct ploidy levels (from 2x to 6x) within
various populations, especially in C. maritima s.l. (Boscaiu et al. 2003). The occurrence of different
cytotypes, supported by genetic and morphological diversity, helped to redefine the taxonomic treatment of
many populations, restoring neglected species (Kreen et al. 2001, Speta 2001, Pfosser & Speta 2004).
The species currently belonging to the C. maritima group are: C. aphylla, C. hesperia, C. maritima, C.
maura, C. numidica and C. pancration.
As already pointed out, the new species C. glaucophylla may be easily distinguished from the aforesaid
species by its morphological, phenological and ecological features.
Due to the diploid chromosome complement, the whitish bulb tunics and the perigon shape, C.
glaucophylla resembles C. pancration, which, however, is significantly distinct in having larger bulb (12–20
× 9–18 cm), red roots, 9–12 lanceolate green leaves, (33–)38–42(–45) × (6–)8–10(–11) cm, inflorescence
175–190 cm long, provided with 400–450 flowers, perigon lobes 7.5–8.2 mm long, ovary 3.7–4.2 mm long,
and flattened stigma. These species further differ in their life cycle, since C. glaucophylla shows early
flowering (July–August) and four months dormancy between flowering and leaf sprout. Leaves of C.
glaucophylla actually develop in winter (January), while both in C. pancration and other species of C.
maritima group they usually sprout in early autumn, afterwards flowering (Fig. 5, 6).
FIGURE 5. Comparison of phenological stages between Charybdis glaucophylla and other Charybdis taxa. Pale gray = foliation.
Dark gray = flowering. White = dormancy. Dashed = fruiting.
Based on its glaucous leaves, C. glaucophylla is similar to C. aphylla and C. maura, but relevant
characters allow these taxa to be distinguished well enough: C. aphylla is a tetraploid species, with red bulb
tunics and lanceolate leaves; instead, C. maura, despite the same diploid chromosome count and white bulb
tunics as C. glaucophylla, deeply differs in leaf morphology, showing very long leaves, narrowly lanceolate
and acute.
As far as chromosome morphology is concerned, the karyotype of C. glaucophylla appears clearly
bimodal in having two outstanding groups of different mean sizes (big chromosomes vs. small ones). This
morphology agrees fairly well with old reports quoted for some Italian populations of C. maritima s.l.,
especially with reference to the peculiar feature of the submetacentric chromosomes having an intercalary
pseudosatellite (Giuffrida 1950, Maugini 1953, Battaglia 1957a). The same structure was also pointed out in
C. maura populations from Morocco (Battaglia 1957a). Recent reports for C. maritima s.l. cytotypes (Boscaiu
at al. 2003) and C. pancration (Rosselló et al. 2005), however, do not evidence this kind of chromosomes,
maybe because satellites on small chromosomes are often not clearly visible. Actually, karyotypes in
Charybdis taxa and populations are quite similar, indicating the existence of interspecific stability of
chromosomes, against a great variability of morphological and anatomical traits and a strong geographic
pattern of different cytotypes. This process can be explained by karyotype orthoselection mechanism, where
structural chromosome mutation occurs in a certain way resulting in uniformity of basic number and gross
morphology of chromosomes (White 1973, Brendham 1983). Similar results, indeed, have been found in
many other monocots, belonging to Asparagaceae, Xanthorrhoeaceae (Alooideae) and Amaryllidaceae
(Brendham & Doherty 1998, Vosa 2005, Cisternas et al. 2010).
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FIGURE 6. Phenological features of Charybdis glaucophylla. A. Plant in rocky habitat (Pranu Sartu, Buggerru-Iglesias). B. Plant in
sandy habitat (Scivu, Arbus). C. Habit in winter (foliation). D. Flowering (Cala Vinagra, Carloforte). E. Habit in late summer (Pranu
Sartu, Buggerru-Iglesias). (Photos: G. Bacchetta).
As Pfosser & Speta (2004) hypothesized, all diploid Charybdis and Urginea populations most likely
displayed a clustering in the western Mediterranean area as far back as Early Miocene, when the main islands
Sardinia, Corsica, Sicily and the Balearic Archipelago were much further west and closest to the Iberian and
African coasts. This overlapping distribution of diploid taxa and the current geographic pattern of Charybdis
cytotypes seems to suggest an early colonization of W Mediterranean by the diploid populations (which are
probably the ancestral ones), starting from the Iberian peninsula and NW Africa, with a subsequent eastward
migration. Then, geographic isolation and ecological adaptation may have allowed some populations to well
differentiate, while autopolyploidy or hybridization processes gave rise to several distinct cytotypes and
genetic patterns.
As a matter of fact, C. glaucophylla is a diploid species endemic to the Sulcis-Iglesiente territory, which
forms the SW part of Sardinia and is isolated from the rest of the island by the Graben of Campidano. This
area, including the oldest geologic elements of Sardinia dates back to the Paleozoic, representing a well
distinct biogeographic sector that is very rich in rare and endemic taxa, many of which are paleoendemics
(Bacchetta & Pontecorvo 2005, Bacchetta et al. 2007). In this biogeographic sector C. glaucophylla is
sympatric only with tetraploid populations of C. maritima s.l. (Boscaiu et al. 2003), but in the four locations
currently known it is found exclusively, and nearby do not find C. maritima; for this reason C. glaucophylla
can be considered allopatric with respect to the latter.
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Based on these considerations, supported by distinctive morphological features, unusual life cycle, diploid
arrangement, geographic confinement and scattered distribution, C. glaucophylla can be considered a relictual
schizoendemic arisen by gradual diversification as a consequence of a long geographic isolation.
Paratypes:—ITALY. Sardinia: Isola di San Pietro: Canale di Basilio, Carloforte (CI), 39°9'58.29"N,
8°14'52.94"E, 15 August 2002, G. Bacchetta & C. Pontecorvo s.n. (CAG); Cala Vinagra, Carloforte (CI),
39°9'47.66"N, 8°14'30.95"E, 3 September 2005, G. Bacchetta & C. Pontecorvo s.n. (CAG); Canalgrande,
Pranu Sartu, Iglesias (CI), 39°21'19.50"N, 8°23'37.86"E, 2 April 2005, G. Bacchetta & C. Pontecorvo s.n.
(CAG); Penisola a N di Punta Cubedda, Pranu Sartu, Iglesias (CI), 39°21’11.60’’N, 8°23’11.20’’E, 27 August
2005, G. Bacchetta, C. Pontecorvo & T. Carai s.n. (CAG); Scivu-Is Arenas, Arbus (VS), 29 March 2010, G.
Bacchetta s.n. (CAG); Monte Linas, Punta Cammedda, Gonnosfanadiga (VS), 39°26'13.55"N, 8°38'13.45"E,
27 July 2012, G. Bacchetta & C. Pontecorvo s.n. (CAG).
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