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Charybdis glaucophylla (Asparagaceae), a new species from Sardinia

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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.
16 Accepted by Lorenzo Peruzzi: 4 Oct. 2012; published online in PDF: 16 Oct. 2012
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN
1179-3163 (online edition)
Copyright © 2012 Magnolia Press
Phytotaxa 69: 1626 (2012)
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/phytotaxa/
Article
Charybdis glaucophylla (Asparagaceae), a new species from Sardinia
GIANLUIGI BACCHETTA
1*
, SALVATORE BRULLO
2
, SAVERIO D’EMERICO
3
,
CRISTIANO PONTECORVO
1
& CRISTINA SALMERI
4
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: bacchet@unica.it
2
Dipartimento di Scienze Biologiche, Geologiche ed Ambientali, Università degli Studi di Catania, Via A. Longo 19, I 95125 Catania,
Italy, e-mail: salvo.brullo@gmail.com
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: cristinamaria.salmeri@unipa.it
*
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
Phytotaxa 69 © 2012 Magnolia Press 17
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
leaves, tepals with green midrib. Finally, C. aphylla is limited to the E Mediterranean area and exhibits white-
reddish 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 green-
whitish 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 south-
western part of the island. They are characterized by whitish bulb tunics and roots, very wide, short, glaucous-
pruinose 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.
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
Pair n. TAL m) 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 sm
sat
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;
CV
CL
: 42.99; CV
CI
: 30.90
BACCHETTA ET AL.18 Phytotaxa 69 © 2012 Magnolia Press
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
Cv
CI
and Cv
CL
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 oblong-
elliptic, 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 sm
sat
+ 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
Phytotaxa 69 © 2012 Magnolia Press 19
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
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).
BACCHETTA ET AL.20 Phytotaxa 69 © 2012 Magnolia Press
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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CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
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 km
2
.
BACCHETTA ET AL.22 Phytotaxa 69 © 2012 Magnolia Press
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
Phytotaxa 69 © 2012 Magnolia Press 23
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
(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)3842(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).
BACCHETTA ET AL.24 Phytotaxa 69 © 2012 Magnolia Press
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.
Phytotaxa 69 © 2012 Magnolia Press 25
CHARYBDIS GLAUCOPHYLLA, A NEW SPECIES FROM SARDINIA
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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... Many authors recently accepted Charybdis to be different from Urginea and Drimia at generic rank (cf. Speta, 1998a;Pfosser & Speta, 2001Conti et al., 2005;Jeanmonod & Gamisans, 2007;Bacchetta et al., 2012;Ali et al., 2013;Tison & Foucault, 2014;Véla et al., 2016), although others (e.g. Manning et al., 2004;Manning & Goldblatt, 2018;WCSP, 2019) still maintain the broadly-circumscribed Drimia. ...
... 30,40,50, 60 (x = 10) (cf. Battaglia, 1957aBattaglia, ,b,c, 1964Valdés-Bermejo, 1980;Speta, 1980;Pfosser & Speta, 2001;Rico, 2013;Bacchetta et al., 2012;Véla et al., 2016). Véla et al., 2016;Crespo et al., 2020). ...
... Sardinia, Corsica, S Italian Peninsula, Croatia, Malta, Lampedusa, and Tunisia) and some localities in the Iberian Peninsula (Gerona) and the Balearic Islands (Mallorca, Menorca and Cabrera) (Figure 2) (see Pfosser & Speta, 2004;Rosselló et al., 2005;Bacchetta et al., 2012;Véla et al., 2016). Remarks.− ...
Article
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Squilla Steinh. was considered to be an orthographic variant of Scilla L., and therefore the new genus Charybdis Speta was created to include Scilla maritima L. and related taxa occurring in the Mediterranean. Molecular phylogenetic studies recovered Charybdis as distant from Urginea; this finding was also supported by morphology and phytochemistry data. However, after typification of Scilla using S. maritima by Rafinesque, Charybdis became illegitimate under Art. 52 of the Shenzhen Code as its name became superfluous when published. A binding decision was requested from the Nomenclature Committee for Vascular Plants (NCVP) on whether Scilla L. and Squilla Steinh. are sufficiently alike to be considered orthographic variants and, hence, to be confused. Most members of the committee favour treating Squilla as not confusable with Scilla, which leaves the former name available for the current concept of Charybdis. In this context, we reevaluate the taxonomy of the genus, accepting 12 species of which eight are accommodated in Squilla as new combinations. Conversely, one of the species of Charybdis is transferred here to Urginavia. Nomenclatural types (including designation of 13 lectotypes, one neotype and one epitype) and the most relevant synonyms are given for each accepted taxon. An identification key is also presented for Squilla to assist future taxonomic studies in this group. We also include a revision of the taxonomic circumscription of the taxa related to S. undulata.
... Charybdis glaucophylla was considered to be endemic to the Sardo-Corsican biogeographical province (Bacchetta et al., 2012): its known distribution included seven subpopulations in Sardinia and one in Corsica. Until recent times, C. glaucophylla was only known for the Sulcitano-Iglesiente biogeographic sector (San Pietro Island, Pranu Sartu, Is Arenas and Monte Linas), but three small subpopulations have been subsequently found in the mountainous inland at Monte Limbara (N Sardinia) (Fenu et al., 2014). ...
... Charybdis glaucophylla (Bacchetta et al., 2012), has been described from a specimen collected from the Island of Santo Pietro near the SW coasts of Sardinia. Its recent inclusion in the genus Drimia (Raus, 2016) is still controversial. ...
... 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 (Bacchetta et al., 2012). ...
... Speta, also a diploid (2n = 20), only for Sardinia, in terms of its Italian distribution (Martinoli, 1949). Furthermore, Bacchetta et al. (2012) described a new species named Charybdis glaucophylla that occurs throughout Sardinia, and was first found in the Iglesiente. According to Speta (1980), Boscaiu et al. (2003), Pfosser & Speta (2004), and Bacchetta et al. (2012) (Colasante, 1996), I. pseudopumila Tineo, and I. planifolia (Mill.) ...
... Furthermore, Bacchetta et al. (2012) described a new species named Charybdis glaucophylla that occurs throughout Sardinia, and was first found in the Iglesiente. According to Speta (1980), Boscaiu et al. (2003), Pfosser & Speta (2004), and Bacchetta et al. (2012) (Colasante, 1996), I. pseudopumila Tineo, and I. planifolia (Mill.) There are few dicotyledons, and these include some Apiaceae: Thapsia garganica L. and Ferula communis L., with all of the subspecies and varieties described, such as: F. arrigonii Bocchieri (endemic to Sardinia) and F. glauca L. [syn.: F. communis subsp. ...
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A phytosociological study that was mainly aimed at describing the dynamic processes of Mediterranean and sub-Mediterranean perennial secondary grasslands is presented here, using the concepts tested previously for similar grasslands in temperate zones. The study was carried out in two different geographic areas: the Gargano peninsula, and the central Apennines between Marche and Umbria. Thirty-six phytosociological surveys were analyzed using cluster analysis. The study highlights that after the abandonment of agro-pastoral activities, in the ecotone zone between the grassland and the wood mantle, macrophytic, nitrophilous and sub-nitrophilous species spread rapidly. These species have underground organs of propagation (e.g., rhizomes, bulbs) that are mostly toxic to herbivorous animals, such as Asphodelus spp., Charybdis spp., Ferula spp., Thapsia spp., Asphodeline spp.. The communities that have participated in these processes are attributed to the new class Charybdido pancratii-Asphodeletea ramosi, within which the new order Asphodeletalia ramosi is described. This order includes four alliances: Charybdido pancratii-Asphodelion ramosi, Asphodelo ramosi-Ferulion communis, and Asphodelion fistulosi in the Mediterranean area; and Asphodelino luteae-Ferulion glaucae in the sub-Mediterranean area. Overall, six new associations and a new sub-association, feruletosum glaucae of the association Cephalario leucanthae-Saturejetum montanae, are described. Furthermore, the international literature on secondary perennial grasslands of much of the Mediterranean basin (i.e., Europe, north Africa, the Middle East) was studied to define the occurrence in these territories of the species considered to be important in the described dynamic processes. We can conclude that the same processes observed in the present study area occur across the whole Mediterranean basin.
... Speta, also a diploid (2n = 20), only for Sardinia, in terms of its Italian distribution (Martinoli, 1949). Furthermore, Bacchetta et al. (2012) described a new species named Charybdis glaucophylla that occurs throughout Sardinia, and was first found in the Iglesiente. According to Speta (1980), Boscaiu et al. (2003), Pfosser & Speta (2004), and Bacchetta et al. (2012) (Colasante, 1996), I. pseudopumila Tineo, and I. planifolia (Mill.) ...
... Furthermore, Bacchetta et al. (2012) described a new species named Charybdis glaucophylla that occurs throughout Sardinia, and was first found in the Iglesiente. According to Speta (1980), Boscaiu et al. (2003), Pfosser & Speta (2004), and Bacchetta et al. (2012) (Colasante, 1996), I. pseudopumila Tineo, and I. planifolia (Mill.) There are few dicotyledons, and these include some Apiaceae: Thapsia garganica L. and Ferula communis L., with all of the subspecies and varieties described, such as: F. arrigonii Bocchieri (endemic to Sardinia) and F. glauca L. [syn.: F. communis subsp. ...
Article
Full-text available
It is here presented a phytosociological study aimed mainly to describe the dynamic processes of Mediterranean and sub-Mediterranean perennial secondary grasslands, using the same concepts already tested in similar grasslands in temperate zones. The study was carried out in two different geographic areas: the Gargano peninsula and the central Apennines between Marche and Umbria. The xx phytosociological surveys have been analyzed through the cluster analysis method. The study has highlighted that in the ecotone zone between the grassland and the wood’s mantle, after the abandonment of the agro-pastoral activities, macrophytic, nitrophilous and subnitrophilous species spread rapidly. These species, such as Asphodelus sp.pl., Charybdis sp.pl., Ferula sp.pl., Thapsia sp.pl., Asphodeline sp.pl. etc., have underground organs of propagation (rhizomes, bulbs etc.), mostly toxic to herbivorous animals. The communities participating in these processes are attributed to the new class Charybdido pancratii- Asphodeletea ramosi, within which the new order Asphodeletalia ramosi is described. The order includes four alliances: in the Mediterranean area there are the alliances Charybdido pancratii-Asphodelion ramosi, Asphodelo ramosi-Ferulion communis and Asphodelion stulosi while in the sub-Mediterranean area the alliance Asphodelino luteae-Ferulion glaucae occurs. On the whole, six new association and a new subassociation, feruletosum glaucae of the association Cephalario leucanthae-Saturejetum montanae, have been found. Furthermore, the international literature on secondary perennial grasslands of much of the Mediterranean basin (Europe, North Africa and the Middle East) has been studied in order to verify the occurring in these territories of the species considered as important in the described dynamical processes. We have concluded that the same process observed in the study area occurs in the whole Mediterranean basin.
... comm.), in which case the name Squilla would be available for the current concept of Charybdis, as already accepted by Martínez-Azorín et al. (2022). Previous phylogenetic analyses (Pfosser & Speta, 2001, 2004Pfosser et al., 2012) place numerous samples of Squilla (as Charybdis) in a strongly supported clade, supporting acceptance of this group as an independent genus (Speta, 1998b;Pfosser & Speta, 2001, 2004Conti et al., 2005;Jeanmonod & Gamisans, 2007;Bacchetta et al., 2012;Ali et al., 2013;Véla et al., 2016). We found our 22 samples of Squilla to form a strongly supported clade in an isolated position within Urgineoideae, and therefore, we recognize this genus based on the hysteranthous leaves; presence of distinct bracteoles; and flattened and winged seeds, together with their Mediterranean distribution (Martínez-Azorín et al., 2022). ...
Article
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The taxonomy and systematics of Urgineoideae (Hyacinthaceae) have been controversial in recent decades with contrasting taxonomic treatments proposed based on preliminary and partial studies that have focused on morphology and/or solely plastid DNA sequence data. Some authors have recognised only two genera, with a very broadly conceived Drimia, while others have accepted several genera which, although better defined morphologically, were doubtfully monophyletic. Here we present phylogenetic analyses involving four plastid DNA regions (trnL intron, trnL‐F spacer, matK and trnCGCA‐ycf6 intergenic region), a nuclear region (Agt1) and a selection of 40 morphological characters. Our study covers 293 samples and ca. 160 species of Urgineoideae (ca. 80% of its global diversity). Bayesian Inference (BI), Maximum Likelihood (ML), and Maximum Parsimony (MP) analyses were performed to derive the phylogenetic patterns. The combination of data yielded phylogenetic trees with 31 well‐defined clades or lineages, most corresponding to previously described genera, although some have required description or revised circumscription. As with other monocot families, a considerable degree of homoplasy was observed in morphological characters, especially in those groups with unspecialised flowers; nonetheless, consistent syndromes of traditional and novel characters are shown to support clade recognition at genus rank. The forthcoming revised classification of Urgineoideae is outlined here. This article is protected by copyright. All rights reserved.
... Recent Mediterranean floras monographs and/or databases accept Charybdis Speta at generic rank (cf. Conti et al. 2005;Bacchetta et al. 2012;Véla et al. 2016; Bartolucci et al. 2018), whereas others include it in a widely circumscribed Drimia (cf. Dobignard and Chatelain 2010;Tison and Foucault 2014;The Plant List 2020) or Urginea (cf. ...
Article
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The identity of Drimia purpurascens, a name usually synonymised to the African D. elata, is clarified. Morphological data in the protologue allow identification of the former with the Mediterranean D. undata (Urginea undulata), which is usually treated as belonging to Charybdis, a genus morphologically and molecularly separated from both Drimia s.str. and Urginea. However, Charybdis is considered to be illegitimate and hence unavailable for use, and when treated as different from Drimia and Urginea, the name Squilla might be applied. A binding decision was requested from the Nomenclature Committee for Vascular Plants on whether Scilla and Squilla are sufficiently alike to be confused. In the meantime, Drimia is here accepted to include, among others, two morphologically distinct aggregates: the “D. undata group” and the “D. maritima group”. The present contribution focusses on the former, in which three species are recognised that show clear morphological and biogeographical differences. Two specific names are revived: D. purpurascens (for D. undata) and D. serotina (for D. tazensis), the latter established as a new combination. Furthermore, a new species (D. palaestina sp. nov.) is described for the easternmost Mediterranean populations referred to D. undata. Similarly, one additional combination, D. secundiflora, is effected in the “Drimia maritima group”. Nomenclatural types (including 3 new lectotypes, 2 new neotypes and 1 new epitype) are indicated, and synonyms are shown for each accepted taxon. The taxonomic treatment for the “D. undata group” including chromosome numbers, phenology, habitats, distribution, and conservation status is presented. An identification key is also reported.
... Elle occupe les pelouses littorales écorchées un peu halophiles, sur substrat arénacé ou graveleux, et même les sentiers soumis à un léger piétinement ; elle individualise une association synendémique avec Catapodium marinum(Géhu et al. 1989). décrit au sud-ouest de la Sardaigne sous le nom de C. glaucophyllaBacchetta et al. (Bacchetta et al. 2012a). J.-M.Tison (comm. ...
... Furthermore, many taxa have been recently described for Italy as new to science (e.g. Bacchetta et al. 2012aBacchetta et al. , 2012bMelai et al. 2012;Peruzzi and Gestri 2013;Troìa and Azzella 2013;Peccenini and Polatschek 2014;Domina and Soldano 2015;Brullo et al. 2015b;Peccenini and Polatschek 2016;Conti and Bartolucci 2017). Therefore, there is a need of a new checklist to summarize the current state of the floristic and taxonomic knowledge of the Italian vascular flora. ...
Article
An updated inventory of the vascular flora alien to Italy, providing details on the occurrence at regional level, is presented. The checklist includes 1,597 species, subspecies, and hybrids, distributed in 725 genera and 152 families; 2 taxa are lycophytes, 11 ferns and fern allies, 33 gymnosperms, and 1,551 angiosperms. 157 taxa are archaeophytes and 1,440 neophytes. The alien taxa currently established in Italy are 791 (570 naturalized and 221 invasive), while 705 taxa are casual aliens, 4 are not assessed, 7 are of unknown regional distribution, 47 have not been confirmed in recent times, 3 are considered extinct or possibly extinct in the country, and 40 are doubtfully occurring in Italy. This checklist allows to establish an up-to-date number (9,792) of taxa constituting the whole (native and alien) Italian flora.
... Furthermore, many taxa have been recently described for Italy as new to science (e.g. Bacchetta et al. 2012aBacchetta et al. , 2012bMelai et al. 2012;Peruzzi and Gestri 2013;Troìa and Azzella 2013;Peccenini and Polatschek 2014;Domina and Soldano 2015;Brullo et al. 2015b;Peccenini and Polatschek 2016;Conti and Bartolucci 2017). Therefore, there is a need of a new checklist to summarize the current state of the floristic and taxonomic knowledge of the Italian vascular flora. ...
Article
Full-text available
An updated inventory of the native vascular flora of Italy, providing details on the occurrence at regional level, is presented. The checklist includes 8195 taxa (6417 species and 1778 subspecies), distributed in 1092 genera and 152 families; 23 taxa are lycophytes, 108 ferns and fern allies, 30 gymnosperms and 8034 angiosperms. The taxa currently occurring in Italy are 7483, while 568 taxa have not been confirmed in recent times, 99 are doubtfully occurring in the country and 19 are data deficient. Out of the 568 not confirmed taxa, 26 are considered extinct or possibly extinct.
Thesis
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Work for the present thesis was focused on three main levels: 1. Collaboration in the implementation and updating of the database of the Flora of Italy, with the preparation of two checklists (native and alien vascular plants) which have been published in the first months of 2018 in two separate articles, of which I am co-author. Multivariate analysis of the data deriving from the two checklists, with special emphasis on the floristic differences among regions. 2. Collaboration in the creation of an online Portal to the Flora of Italy, which integrates the data of the two checklists and connects them with additional multimedia resources deriving from several databases of the Dryades Project (University of Trieste), and Acta Plantarum. 3. Analysis of data deriving from 3 interconnected databases: 1) The database of the Flora of Italy; 2) The database of the Flora of Friuli Venezia Giulia by L. Poldini; 3) The database of morphological traits of vascular plants of Project Dryades. The two national checklists include a total of 9792 infrageneric taxa (8195 native and 1597 alien), a number which is much higher than those of previous checklists and floras, with data on nomenclature, taxonomy, synonymy, and regional distribution (presence / absence, doubtful presence, presence not confirmed in recent times, no longer present / extinct, adventitious, naturalized, invasive, etc.). The multivariate analysis of the data related to autochthonous and alien plants at regional level was carried out separately for the two groups taxa, highlighting significant floristic differences among the different regions of Italy, which are correlated to different characteristics of the respective territories (surface area, environmental heterogeneity, altitudinal range, types and number of protected areas, climatic variables, land use, etc.). The Portal of the Flora of Italy allows to carry out queries on the flora of Italy and of the individual Italian regions. For each infrageneric taxon, a "taxon page" is provided, including all the resources deriving from the national checklists, and those deriving from different databases of the Dryades Project: scientific binomial, systematic position (following APG IV) and relative cladograms, regional distribution (with auto-generating maps) and current status, Italian vernacular names and digital images (often showing distinctive characters for identification), plus links to resources from ActaPlantarum. The third and last part of this thesis was devoted to the exploration of the potentialities offered by the achieved interoperability between the database of the Flora of Italy, the resources of Project Dryades, and those of the Database of the Flora of Friuli-Venezia Giulia by Poldini. We show different examples of datasets which is now possible to rapidly obtain from three main databases, including the ecological characterization of vegetation releves by ecological indices, the definition of “virtual habitats” defines by lists of species with a similar ecology, the analysis of complex matrices of ecological data and morphological traits. Such tests will be useful to enrich the future versions of the Portal to the flora of Italy with further data and further functionalities. The creation of the Portal of the Flora of Italy and its interoperability with the resources of the Dryades Project and those of Acta Plantarum are not only a point of arrival, but above all a starting point for future interesting developments. The data made public on the Portal of the Flora of Italy are only a small part of those that can be made searchable in future versions. Some of them are already present in the Flora of Italy Database for more than 80% of the species (biological forms, Ellenberg ecological indexes, flowering periods, altitudinal distribution), others may derive from local databases that can be integrated into the system (e.g. regional floristic databases, the Wikiplantbase Project, etc.), while a wealth of data concerning the morphological features of the species can already be found in the Dryades Project database. With this thesis I have contributed to laying the foundations for a future distributed database on the flora and vegetation of Italy, where data from the Portal to the Flora of Italy can act as a nucleus of crystallization for the aggregation of several other databases that at the moment are not public and/or are not able to communicate with each other.
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Chromosome numbers and karyotypes of seven specific taxa of the Chilean endemic genus Placea were determined. Chromosome numbers of P. lutea, P. ornata, P. grandiflora, P. germainii and P. aff. davidii are described for the first time. All taxa are diploid with 2n=2x=16 and karyotypes are composed of four metacentric (4 m), ten submetacentric (10 sm), and two subtelocentric (2 st) chromosomes. The most symmetrical karyotype was observed in P. lutea (AI: 6.84) while the most asymmetrical karyotype was shown by P. arzae (AI: 9.72). The constancy in karyotype formulae and high similarity in asymmetry indexes suggest that some orthoselection mechanism might be involved in Placea's chromosomal evolution. In spite that no significant karyotypic differences were observed, the species may be differentiated by their chromosomal sizes. Moreover, the tribal position of Placea and its likely relationships with other hippeastroid genera are discussed.
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The results of a survey on the endemic vascular flora of Iglesiente (SW Sardinia) are presented. This flora consists of 140 taxa, including 115 species, 18 subspecies, 4 varieties and 3 hybrids; 92 genera and 34 families are represented. The most represented genera are: Asteraceae (20 taxa), Caryophyllaceae (13) and the Orchidaceae (11). The most represented genera are: Ophrys (9 taxa), Genista (7), Silene and Dianthus (5). The analysis of biologic and chorologic data highlighted the environmental peculiarities of Iglesiente, given by the local evolution of a rich floristic contingent. Eighteen taxa exclusive endemics, together with an unique blend of geolithologic, geomorphologic, paleogeographic, bioclimatic and vegetation features, let to attribute the rank of biogeographic sector to the Sulcis-Iglesiente territory, whose northern part, with 9 exclusive endemics, can be seen as the Iglesiente subsector. From the surveyed taxa, 32 are included in the IUCN Red List and 5 in the EU 92/43 Habitat Directive. Moreover, 11 species (including all the Orchidaceae) are protected by the CITES (Convention on International Trade in Endangered Species of wild flora and fauna) and 1 species by the Bern Conventions.
Article
Dallo studio morfologico e cariologico di alcune specie di Urginea e di Scilla appartenenti alla flora spontanea dei dintorni di Cagliari (Sardegna meridionale) è apparso chiaro che esistono in Sardegna varietà e forme finora non note, e ne sono stati stabiliti i cartogrammi corrispondenti: a) Urginea undulata Steinh., a 2n = 20.b) Urginea fugax Steinh. con due biotipi: uno a 2n = 21 e corrispondente probabilmente alla varietà typica, l'altro a 2n = 24 corrispondente probabilmente alla varietà major Litardier et Maire, quest'ultima finora nota soltanto per l'Africa boreale;c) Urginea maritima Bak., a 2n = 40, corrispondente ad un biotipo tetraploide;d) Scilla obtusifolia Poir. con parecchie varietà: la typica, la var. intermedia (Guss.) Baker con la forma strida n. f., la forma lata n. f., la forma brevis n. f. e infine la var. glauca Gat. et Weiller con la forma nana n. f.: tutte queste entità presentano 2n = 8 cromosomi, che è il cariogramma numericamente più basso nel genere Scilla, di cui viene stabilito il nuovo numero base x = 4.e) Scilla autunnalis L., a 2n—28.Tutte le specie esaminate dei generi Urginea e Scilla presentano cromosomi con « intercalary trabant ». Di tutte le specie viene illustrata la morfologia delle singole coppie cromosomiche.SUMMARYFrom a study of the morphology and caryology of certain species of Urginea and Silla collected near Cagliari (Southern Sardinia), it appears that there exist in Sardinia forms not previously described, with the following chromosome numbers: a) Urginea undulata Steinh,.—2n = 20.b) Urginea fugax Steinh.,—two biotypes: one 2n = 21, probably correponding to var. typica; the other 2n = 24, probably corresponding to var. major Litardière et Maire; this latter so far reported only from North Africa.c) Urginea maritima Bak.,—2n = 40, a tetraploid biotype.d) Scilla obtusifolia Poir.,—various varieties: typica; var. intermedia (Guss.) Bak., with the forms stricta n. f., lata n. f., and brevis n. f.; var.glauca Gat. et Weiller, with the form nana n. f. All these present 2n = 8, numerically the one lowest in the genus Scilla, establishing the new basic number x = 4.e) Scilla autumnalis L.,—2n=28.All the above species of Urginea and Scilla show chromosomes with intercalary trabants The morphology of the single chromosome pairs is given in each case.
Article
A historical survey is given showing the changes of what belonged to the genera Scilla L., Hyancinthus L. and Ornithogalum L. during the centuries. many efforts have been made to find a natural system and to enlighten the phylogeny. According to karyological and morphological characters the species of the extensive genus Scilla L. form related groups all of which have been confirmed by DNA sequencing data. Therefore it is very reasonable to address these groups as genera. The Scilla of the ancient Greeks and Romans and some relative genera form the subfamily Urgineoideae SPETA subfam. nova. The genera Urginea STEINH., Drimia JACQ., Thurantus WRIGHT, Charybdis SPETA nom. novum, Rhadamanthus SALISB., Fusifilum RAF., Rhadamanthopsis (OBERM.) SPETA stat. novus, Igidia SPETA gen. novum, Ebertia SPETA gen. novum, Urginavia SPETA gen. novum, Tenicroa RAF., Urnothigaloideae SPETA subfam. nova and the Hyacinthoideae LINK. Most species of Scilla s. l. belong to the Hyacintheae DUMORT. Scilla L. s. str. (= S. bifolia-retionship and Chionodoxa BOISS.), Schnarfia SPETA gen. novum (= S. messeniaca-rel.), Zagrosia SPETA gen. novum (= S. persica-rel.), Nectaroscilla PARL. (= S. hyacinthoides-rel.), Chouardia SPETA gen. novum (= S. litardierrei-rel.), Fessia SPETA gen. novum (= S. haemorrhoidalis-rel.), Barnardia LINDLEY (= S. scilloides-rel.), while Ledebouria ROTH (= S. hyacinthoides-rel.), Schizocarphus MERWE (= S. nervosa-rel.), Resnova MERWE, Avonsera SPETA gen. novum, Merwilla SPETA gen. novum and Pseudoprospero SPETA gen. novum belong to the Massonieae BAKER. The South American genus Oziroë RAF. forms the subfamily Oziroëoideae SPETA subfam. nova. The North American genus Camassia LINDLEY belongs to the Chlorogaloideae SPETA subfam. nova. Bcause of the new genera 128 new combinations for specific names were necessary.