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Siculosciadium, A New Monotypic Genus of Apiaceae from
Sicily
Author(s): Cristian Brullo , Salvatore Brullo , Stephen R. Downie , Clark A.
Danderson and Gianpietro Giusso del Galdo
Source: Annals of the Missouri Botanical Garden, 99(1):1-18. 2013.
Published By: Missouri Botanical Garden
DOI: http://dx.doi.org/10.3417/2011009
URL: http://www.bioone.org/doi/full/10.3417/2011009
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Volume 99 Annals
Number 1 of the
2013 Missouri
Botanical
Garden
SICULOSCIADIUM, A NEW Cristian Brullo,
2
Salvatore Brullo,
2
MONOTYPIC GENUS OF Stephen R. Downie,
3
Clark A. Danderson,
3
and
1
Gianpietro Giusso del Galdo
APIACEAE FROM SICILY
2
ABSTRACT
The results of a morphological and molecular systematic investigation of the poorly known Sicilian endemic species
Peucedanum nebrodense (Guss.) Nyman (Apiaceae subfam. Apioideae) are presented in order to better characterize the species
and clarify its phylogenetic position. Morphological, karyological, anatomical, and ontogenetic studies indicate that this species
is taxonomically distinct. Phylogenetic analysis of combined morphological and nuclear ribosomal DNA (nrDNA) ITS sequence
data suggests an affinity with Dichoropetalum Fenzl, whereas an unweighted pair group method with arithmetic mean (UPGMA)
analysis, based on 36 morphological characters, suggests that P. nebrodense is similar to Ormosolenia Tausch. Given its unusual
life cycle; distinct suffruticose, evergreen, and dwarf habit; and peculiar fruit anatomy, we recognize P. nebrodense as distinct
from Ormosolenia and Dichoropetalum, and we treat it as the new monotypic genus Siculosciadium C. Brullo, Brullo, S. R.
Downie & Giusso based on S. nebrodense (Guss.) C. Brullo, Brullo, S. R. Downie & Giusso [[Pteroselinum nebrodense Guss.],
which is lectotypified herein.
Key words: Apiaceae, IUCN Red List, Peucedanum, Sicily, Siculosciadium.
The genus Peucedanum L., as traditionally defined family Apiaceae (Winter et al., 2008: 347). The
within the Apiaceae, contains more than 100 species results of molecular systematic investigations have
of Eurasian and African distribution (Pimenov & confirmed the polyphyletic nature of Peucedanum
Leonov, 1993). However, the genus has long been s.l., and, as a consequence, the genus is now reduced
regarded as an unnatural group, and the need for its to only a few species allied to the type species, P.
reduction into smaller, monophyletic units has been officinale L., and several segregates of Peucedanum
proposed by many (Pimenov, 1987, 1992; Burtt, are now recognized as distinct genera (Downie et al.,
1991; Pimenov & Leonov, 1993; Ostroumova & 1998, 2000; Spalik et al., 2004; Winter et al., 2008;
Pimenov, 1997; Reduron et al., 1997; Shneyer et al., Magee et al., 2009). The results of these investiga-
2003; Pimenov et al., 2007). Indeed, the subdivision
of the large polymorphic genus Peucedanum into tions also placed Peucedanum s. str. and other
natural groups is ‘‘among the most urgent challenges’’ segregates in the revised Apiaceae tribe Selineae
facing the realization of a modern classification of the (Spalik et al., 2004; Downie et al., 2010).
1
The authors thank the curators of the herbaria cited in the text for the loan of herbarium materials. We are also very
grateful to M. C. Terrasi for help with the karyological investigations and to G. Venora for help with the fruit anatomy.
2
Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Universita
`di Catania, Via A. Longo, 19, I – 95125,
Catania, Italy. salvo.brullo@gmail.com.
3
Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, Illinois 61801, U.S.A.
salvo.brullo@gmail.com
doi: 10.3417/2011009
ANN.MISSOURI BOT.GARD. 99: 1–18. PUBLISHED ON 19 JULY 2013.
2 Annals of the
Missouri Botanical Garden
One species of Peucedanum whose biology and understand its ontogeny and phenology, fruits,
phylogenetic placement have yet to be investigated is seedlings, and mature plants were cultivated at the
P. nebrodense (Guss.) Nyman. This species was first Botanical Garden of Catania, Italy, for several years.
referred to Imperatoria chabraei Spreng. by Gussone Additional characters considered were those of leaves
(1827), based on specimens from the Madonie Massif and other vegetative organs, inflorescences, flowers,
(northern Sicily), which still represents its sole locality. and fruits. Specimens of P. nebrodense were
Gussone (1844) later attributed the population from examined from herbaria B, CAT, FI, G, M, NAP,
Madonie to a new species, namely Pteroselinum PAL, RO, W, and WU. Specimens from Dichorope-
nebrodense Guss., and highlighted its morphological talum Fenzl, Johrenia DC., Ormosolenia,Peuceda-
differences from Pteroselinum chabraei (Spreng.) num s. str., and Zeravschania Korovin were also
Rchb. Nyman (1855) later ascribed this species to included in this comparative examination of general
the genus Peucedanum, a transfer supported by both morphology, with emphasis placed on P. nebrodense
Cesati et al. (1884) and Strobl (1886). Subsequently, and Ormosolenia. Recently, Pimenov et al. (2007)
Paoletti (1900) and Fiori (1925) considered this taxon considered Holandrea,Johreniopsis Pimenov, and
a variety of Peucedanum carvifolia (L.) Vill. Pignatti some species of Johrenia and Peucedanum as
(1982), however, stressed the unclear and still belonging to the genus Dichoropetalum.
unsolved taxonomic position of P. nebrodense,and
Banfi et al. (2005) attributed this species to the genus KARYOLOGY
Holandrea Reduron, Charpin & Pimenov, although Mitotic metaphase plates were obtained from
without substantiating evidence. It is likely that the squashed root tips of cultivated plants of Peuceda-
classification proposed by Banfi et al. (2005) was num nebrodense, pretreated with 0.3%colchicine
based on the transfer of Peucedanum carvifolia into the water solution for three hours and fixed in ethanol-
genus Holandrea by Reduron et al. (1997). acetic acid (3:1) for six hours, then hydrolyzed in 1N
Because of their rarity and extreme localized HCl for seven minutes and stained, according to the
distribution, specimens of Peucedanum nebrodense Feulgen method. Metaphase observations and chro-
are not commonly found in major European herbaria, mosome measurements were made using the image
and the species is usually not treated in European analysis systems IKAROS 4.6 (Metasystem, Altlus-
floras. The plant does have a very unusual habit, sheim, Germany) and Zeiss (Oberkochen, Germany)
however, when compared with the other known AxioVision 5.1. Karyotyping was done with the
species of Peucedanum, which are commonly hemi- software Cromolab 1.1 (Brullo, 2002–2003), which
cryptophytes. Peucedanum nebrodense is character- was used to recognize homologues, to order chromo-
ized by a dwarf shrubby habit with many woody somes by size, and to classify them according to
branches and persistent leaves. As such, the species morphology and centromere position (Levan et al.,
appears taxonomically well isolated, showing a 1964). Ten mitotic plates from five individuals were
morphological affinity only with P. alpinum (Sieb. used to determine the karyotype.
ex Schult.) B. L. Burtt & P. H. Davis from Crete, a
taxon recently attributed by Pimenov (1992) to the ANATOMY
restored genus Ormosolenia Tausch. The morpholog- Fruit and leaf petiole anatomy was examined from
ical similarities between these species include their living material collected from the type locality, fixed
branched rootstocks, well-developed caudices, and in Karpechenko solution, and then embedded in
polycarpic life form. To better characterize the poorly paraffin. Transverse sections were double stained
known species P. nebrodense and to address its with ruthenium red and light green. Comparisons in
taxonomic position, we conducted morphological, leaf petiole anatomy were made between Peucedanum
karyological, and anatomical studies, and present nebrodense and a putative ally, Ormosolenia alpina
herein the results of a phylogenetic analysis of this (Sieber ex Schultes) Pimenov.
species and its closest allies, using morphological
and molecular evidence.
PHENETIC ANALYSIS OF MORPHOLOGICAL CHARACTERS
MATERIALS AND METHODS We supplemented and slightly modified the
morphological data matrix provided by Pimenov et
GENERAL MORPHOLOGY al. (2007) in their comparative multivariate analyses of
Morphological investigations of Peucedanum ne- Holandrea,Johrenia,Johreniopsis,Peucedanum s.l.,
brodense were based on an examination of living and Zeravschania by including the species Ormosole-
material collected from the type locality. In order to nia alpina,O. pisidica Boiss. & Heldr., and P.
Volume 99, Number 1 Brullo et al. 3
2013 Siculosciadium (Apiaceae) from Sicily
nebrodense and two additional characters (i.e., leaf avschania Korovin and Pimpinella L. (both in tribe
persistence and rootstock habit). In total, the data Pimpinelleae) were used to root the trees. These
matrix comprised 36 morphological characters (Ap- sequence data were analyzed using maximum
pendix 1) and 41 species. The importance of these parsimony as implemented by PAUP* version
characters in distinguishing among these putatively 4.0b.10 (Swofford, 2003), and the resultant strict
related taxa has been highlighted by Pimenov et al. consensus tree revealed that Ormosolenia and P.
(2007). No data cells in the matrix were scored as nebrodense comprised a well-supported clade along
missing. Our rationale for including a phenetic with representatives of Holandrea,Johrenia, and
analysis in this paper was to comprehensively compare Johreniopsis (many of which have been treated as
those diagnostic characters across all included taxa to species of Dichoropetalum, according to Pimenov et
ascertain what species might be most similar morpho- al., 2007). Sister group to this clade was Aethusa
logically to P. nebrodense. The resultant data matrix is cynapium L. Subsequently, another data matrix was
presented in Appendix 2. Cluster (unweighted pair produced to include all species of Holandrea,
group method with arithmetic mean [UPGMA]) Johrenia, and Johreniopsis for which ITS data were
analysis of these data was implemented by using the available in GenBank. Thirteen species comprised
NTSYSpc package (Rohlf, 2000). this matrix, including Ormosolenia and P. nebro-
dense, and A. cynapium was used to root the resulting
DNA SEQUENCING phylogenetic trees. GenBank accession numbers and
voucher information for these species are presented
Total genomic DNAs of Ormosolenia alpina,O. in Table 1. This reduced matrix was analyzed by
pisidica, and Peucedanum nebrodense were extracted using heuristic maximum parsimony searches and
from approximately 20 mg of dried leaf tissue using a 1000 replicate analyses, random stepwise addition of
DNeasy Plant Mini Kit (Qiagen, Valencia, California, taxa, and tree bisection-reconnection (TBR) branch
U.S.A.). The nuclear ribosomal DNA (nrDNA) ITS swapping. Bootstrap values were calculated from
region in each of these species was polymerase chain 1000 replicate analyses, using TBR branch swapping
reaction (PCR)-amplified by using the protocol and simple stepwise addition of taxa. In the
described in Downie and Katz-Downie (1996). For phylogenetic analysis of morphological data, 17
template purification, the QIAquick PCR Purification parsimony-informative morphological characters were
Kit (Qiagen) was used. Sequencing reactions were considered, representing both vegetative and repro-
carried out using a BigDye Terminator Cycle ductive structures (Appendix 3). Eleven characters
Sequencing Kit (version 3.1; Applied Biosystems, were binary, the remaining multistate; all characters
Foster City, California, U.S.A.) and sequenced with were unordered and equally weighted. The taxa
an ABI 3730xl sequencer (Applied Biosystems). The included in this portion of the study corresponded to
simultaneous consideration of both DNA strands those included in the 13-species ITS matrix, and the
across the entire ITS region permitted unambiguous data matrix of morphological characters is presented
base determination. ITS sequence heterogeneity, as in Appendix 4. These ITS and morphological data
evidenced by polymorphisms at single bases in were analyzed separately and combined, with only
otherwise perfect chromatograms, was not detected the results of the combined analysis presented
for any accession. GenBank accession numbers and herein. Separate maximum parsimony analysis of
specimen voucher information for O. alpina,O. each data set resulted in highly consistent yet
pisidica, and P. nebrodense are presented in Table 1. generally poorly resolved and weakly supported trees,
whereas analysis of the combined matrix yielded
PHYLOGENETIC ANALYSES OF MOLECULAR AND trees of greatest resolution and highest branch
MORPHOLOGICAL DATA support overall. The data matrix of combined
To ascertain the phylogenetic positions of Ormo- morphological and ITS sequence data, as well as
solenia alpina,O. pisidica, and Peucedanum nebro- the resulting strict consensus tree, is available from
dense, their ITS sequences were first analyzed with TreeBase (,http://purl.org/phylo/treebase/phylows/
40 additional ITS sequences from other members of study/TB2:S12922.).
the Apiaceae, tribe Selineae. These sequences were
obtained from GenBank and represented most major RESULTS AND DISCUSSION
clades of the tribe. Included here were representa- KARYOLOGY
tives of Eurasian Peucedanum and its segregates,
including P. officinale (the type for Peucedanum), According to Solov’eva et al. (1985) and Pimenov
Imperatoria L., and Pteroselinum Rchb., and Zer- et al. (2003), species of Peucedanum s.l. are usually
diploid (2n¼2x¼22), a feature common in the Karyological data are currently unavailable for other
subfamily Apioideae. Peucedanum nebrodense has a species of Dichoropetalum and Johrenia, as well as
diploid chromosome count 2n¼22 (Fig. 1A, part 1), for Ormosolenia and many other Apiaceae. Never-
as previously reported by Duro et al. (2003), and has theless, the karyotype of P. nebrodense appears to be
a karyotype characterized by five metacentric pairs of unique, bolstering its possible isolated taxonomic
chromosomes, by five pairs of chromosomes approx- position within the Apiaceae.
imate to the submetacentric type, and by one
subtelocentric pair (Fig. 1A, part 2). Two chromo- LEAF MORPHOLOGY AND PETIOL E ANATOMY
some pairs have microsatellites. This karyotype Peucedanum nebrodense has 2-pinnate leaf blades,
differs from those of other examined Peucedanum occasionally 1-pinnate in the cauline leaves, with
species, including those species recently transferred sessile or subsessile leaflets (Fig. 1B, part 1). The
to Dichoropetalum (i.e., D. carvifolia and D. pscha- basal leaves are always long-petiolate, although in the
wicum), and from those of genera segregated from cauline ones the petiole tends to reduce progressively
Peucedanum s.l. (i.e., Imperatoria and Thysselinum upward until disappearing; in such cases, the leaf
Adans.; Solov’eva et al., 1985). The karyotype of blades are inserted directly on the sheaths. The
Peucedanum s.l. is characterized by at least seven petioles in cross section are falcate, with an adaxial
pairs of metacentric chromosomes, while telocentric groove, and have five collenchymatous ribs in the
chromosomes have never been detected. Similarly, in abaxial face and two in the adaxial face. Vascular
Peucedanum s.l., microsatellites are absent or, if bundles are five, all arranged in the peripheral part of
present, occur in one pair of chromosomes only. the parenchyma, and interpose with four secretory
Table 1. Accessions of Apiaceae examined for nuclear ribosomal DNA ITS sequence variation, with specimen voucher
information and GenBank accession numbers. Two GenBank accession numbers for a taxon refer to ITS-1 and ITS-2 data only, with
no data available for the intervening 5.8S region; a single GenBank accession number indicates that 5.8S data were available and
used in the phylogenetic analysis. With the exception of data for the two Ormosolenia species and Siculosciadium nebrodense, all ITS
data were previously published by others and obtained from GenBank. Nomenclature follows Pimenov et al. (2007).
GenBank accession
Taxon Voucher information number(s)
Aethusa cynapium L. Germany. Cult. at UIUC from seeds, Univ. Oldenburg GQ862376
Bot. Gard., Germany, Downie 127 (ILL)
Dichoropetalum achaicum Greece. N. Pelo
´ponnisos, Vouraikos Gorge, Southam s.n. AF164832, AF164857
(Hala
´csy) Pimenov & Kljuykov (E), cult. Royal Bot. Gard. Edinburgh (no. 19912669)
D. aromaticum (Rech. f.) Pimenov Iran. Ajani s.n. (Herb. Hossein Akhani, Univ. Tehran) EU169288
& Kljuykov
D. carvifolia (Vill.) Pimenov & France. Ise
`re, mont Bovinant, Chartreuse, 22 Aug. 1989, AF495828, AF495829
Kljuykov Reduron s.n. (WA)
D. golestanicum (Rech. f.) Iran. Ajani 2053 (TEH) EU169289
Pimenov & Kljuykov
D. paucijugum (DC.) Pimenov & Iran. Ajani 852 (Herb. Hossein Akhani, Univ. Tehran) EU169290
Kljuykov
D. pschawicum (Boiss.) Pimenov & Russia. S. Ossetia, Bad, Pimenov et al. 112 (MW), cult. AF008619, AF009098
Kljuykov Moscow State Univ. Bot. Gard.
D. schottii (Besser ex DC.) France. Alpes-Maritimes, col de Brouis, 30 July 1981, AF495830, AF495831
Pimenov & Kljuykov Reduron s.n. (WA)
D. scoparium (Boiss.) Pimenov & Iran. Arak Prov., Zalion Pass betw. Arak & Borujet, AY941274, AY941302
Kljuykov 338530N, 498000E, 17 June 2001, Pimenov et al. 406
(MW)
D. seseloides (C. A. Mey.) Pimenov Iran. Tehran Prov., Ajani 2057 (TEH) EU169291
& Kljuykov
Ormosolenia alpina (Sieber ex Greece. Crete, Lefka Ori, Mt. Pachnes, 1 Sep. 2006, HQ269391
Schult.) Pimenov Brullo & Giusso del Galdo 2061 (CAT)
O. pisidica Boiss. & Heldr. Turkey. Davraz Dag, 20 June 1998, Brullo & Pavone s.n. HQ269392
(CAT)
Siculosciadium nebrodense (Guss.) Italy. Sicily, Madonie, Pizzo Carbonara, Fosse di S. HQ269390
C. Brullo, Brullo, S. R. Downie Gandolfo, 24 July 2002, Brullo & Giusso del Galdo s.n.
& Giusso (CAT)
4 Annals of the
Missouri Botanical Garden
Figure 1. —A. Mitotic chromosome number of Siculosciadium nebrodense. —Part 1. Somatic metaphase plate. —Part 2.
Karyotype (2n¼22). Root tip preparations from the 24 July 2002 collection, S. Brullo,G. Giusso del Galdo & S. Sciandrello s.n.
(CAT). —B. Leaf variability of S. nebrodense. —Part 1. Living material from Madonie (Sicily). —Part 2. Petiole cross section of
S. nebrodense. —Part 3. Petiole cross section of Ormosolenia alpina. —C. Vegetative stages of S. nebrodense (all living material
taken from Madonie, Sicily). —Part 1. Seedlings. —Part 2. Juvenile plant. —Part 3. Adult sterile plant.
Volume 99, Number 1 Brullo et al. 5
2013 Siculosciadium (Apiaceae) from Sicily
6 Annals of the
Missouri Botanical Garden
ducts (Fig. 1B, part 2). The petiole anatomy of cycle quite different from that known for other
Ormosolenia alpina is rather different from that of P. species of Peucedanum and Ormosolenia.The
nebrodense. The petiole cross section of O. alpina mericarps of P. nebrodense reach maturity between
shows that its collenchymatous ribs are not prominent early September and October, when they gradually
(Fig. 1B, part 3). Interposed among the collenchyma- become loose and fall. These mericarps remain for
tous ribs and vascular bundles are five secretory ducts about four months (December to early April) under
that unlike in P. nebrodense are directly in contact with the snow bed and only after the snow melts do they
the collenchyma. The leaf morphology of P. nebrodense start to germinate (late April to May). The seedlings
suggests an affinity to some species of Dichoropetalum are characterized by well-developed roots, two
(Pimenov et al., 2007), whereas the petiole anatomy of oblanceolate to ovate-oblanceolate cotyledons that
P. nebrodense is unusual within the group. abruptly narrow into a 3-nerved petiole (0.5–0.8 mm),
and a null hypocotyl (Fig. 1C, part 1). The blade is 8–
FRUIT MORPHOLOGY AND ANATOMY 13 31.5–4 mm, 3-nerved, with secondary nervation
at maturity. The primordial leaf is 1- to 2-pinnate,
The mericarps of Peucedanum nebrodense are long-petiolate, and subequal or longer than the
subrounded to elliptical, dorsally compressed, with cotyledons. Afterward, the seedling produces other
distinctive spongy wings. The stylopodium is conical- leaves and, after this stage, it develops rhizomes
flattened and the curved stylodium is much longer bearing new leaves at its apex (Fig. 1C, part 2).
than the stylopodium (Fig. 2A, parts 1, 2). The During the second year, the rhizomes grow and
exocarp is single-layered, interrupted near the ventral branch, producing a well-developed rootstock (Fig.
portion of the wings, and followed by an irregularly 1C, part 3). From June to early September, the
stratified parenchyma interspersed with thin-walled caudices produce flowering stems from the center of
cells in the outer mesocarp and thickened in the ribs; the leaf rosette. Therefore, P. nebrodense is a
the vascular bundles and secretory ducts are sunken perennial, polycarpic, and evergreen plant because
in this parenchyma (Figs. 2B, 2C). The inner layer of its leaves persist through the winter (under the snow
the mesocarp consists of one row of tangentially bed), a feature we observed on both wild and
elongated prosenchymatous cells with slightly ligni- cultivated plants. Peucedanum s. str., Dichoropeta-
fied walls. The endocarp consists of one row of lum,Johrenia, and Zeravschania are characterized by
collenchymatous cells. Five of 11 oil ducts are mono- or polycarpic plants with well-developed
associated with as many vascular bundles, four are taproots, and basal leaves arranged in deciduous
vallecular and two are commissural, but do not reach rosettes, usually with the remains of petioles at the
the mericarp base. According to Pimenov (1992) and stem base, while Ormosolenia is polycarpic with
our own observations, the aforementioned features are branched rootstocks, only basal leaves that are entire
quite different from those of Ormosolenia alpina and or 1-pinnate, with blades reniform to orbicular.
O. pisidica (Figs. 2A, parts 3–6; 2C, parts 2, 3). The Peucedanum nebrodense possesses an unusual life
latter two species are characterized by a more cycle and its suffruticose, evergreen, and dwarf habit
flattened stylopodium, a stylodium shorter than the is distinctive within the group.
stylopodium, three to six vittae per vallecula, and four
to eight commissural vittae (secretory ducts associ- PHENETIC ANALYSIS OF MORPHOLOGICA L CHARACTERS AND
ated with vascular bundles are not always present). COMPARATIVE MORPHOLOGY
The inner layer of the mesocarp of both species Peucedanum nebrodense has been considered
consists of three to five rows of elongated prosen- morphologically similar to P. carvifolia (Paoletti,
chymatous cells and the single-layered endocarp also 1900; Fiori, 1925; Banfi et al., 2005), the latter
has tangentially elongated cells. The mericarp recently included by Pimenov et al. (2007) in
anatomy of Dichoropetalum,Johrenia, and Zerav- Dichoropetalum sect. Holandrea. In contrast, the
schania is rather variable, but is always well unweighted pair group method with arithmetic mean
differentiated from that of P. nebrodense (Pimenov (UPGMA) phenogram (Fig. 3A) revealed that P.
et al., 2007). In summary, the peculiar fruit nebrodense clustered with Ormosolenia alpina and O.
morphology and anatomy of P. nebrodense do not pisidica and was separate from any species attribut-
suggest any close taxonomic ally. able to Dichoropetalum (including Holandrea),
Johrenia,orZeravschania. There are, however,
LIFE CYCLE remarkable morphological differences between Or-
Both field and greenhouse observations reveal that mosolenia and P. nebrodense, such that the inclusion
Peucedanum nebrodense is characterized by a life of the latter species into Ormosolenia is difficult to
accept upon the basis of these characters, several of basal leaves that are deciduous, reniform to ovate in
which were not included in the phenetic analysis. outline, entire to 1-pinnate, and with up to one pair
These differences chiefly concern their ontogenetic of leaflets. Its cauline leaves are usually absent. The
cycle, life form, leaf shape, and petiole and fruit petiole has five secretory ducts between the
anatomy (Table 2). The genus Ormosolenia is collenchyma and vascular bundles, the petals are
characterized by a nonsuffruticose rootstock and yellowish green, the mericarps have three to six
Figure 2. —A. Mericarp variability in Siculosciadium and Ormosolenia. —Part 1. Mericarp, dorsal view, S. nebrodense (30
Sep. 2009, Brullo et al. s.n., CAT). —Part 2. Mericarp, commissural view, S. nebrodense (30 Sep. 2009, Brullo et al. s.n., CAT).
—Part 3. Mericarp, dorsal view, O. alpina (1 Sep. 2006, Brullo & Giusso 2061 CAT). —Part 4. Mericarp, commissural view (1
Sep. 2006, Brullo & Giusso 2061, CAT). —Part 5. Mericarp, dorsal view O. pisidica (20 June 1998, Brullo & Pavone s.n., CAT).
—Part 6. Mericarp, commissural view (20 June 1998, Brullo & Pavone s.n., CAT). —B. Mericarp cross sections of S. nebrodense
(all living material from Madonie, Sicily). —Part 1. Mericarp, lateral view. —Part 2. Mericarp, dorsal central view. —Part 3.
Mericarp, commissural central view. —C. Mericarp cross sections of Siculosciadium and Ormosolenia. —Part 1. Mericarp, S.
nebrodense (30 Sep. 2009, Brullo et al. s.n., CAT). —Part 2. Mericarp, O. alpina (1 Sep. 2006, Brullo & Giusso 2061, CAT).
—Part 3. Mericarp, O. pisidica (20 June 1998, Brullo & Pavone s.n., CAT).
Volume 99, Number 1 Brullo et al. 7
2013 Siculosciadium (Apiaceae) from Sicily
Figure 3. —A. UPGMA phenogram showing the morphological relationships among the genera Ormosolenia,
Dichoropetalum,Johrenia,Zeravschania, and Siculosciadium. —B. Strict consensus tree of ten 129-step trees resulting from
maximum parsimony analysis of 17 parsimony informative morphological characters and DNA sequence data from the nrDNA
ITS region. The numbers above the branches are bootstrap values.
8 Annals of the
Missouri Botanical Garden
.Zeravschaniaand,Johrenia,Dichoropetalum,Peucedanum,Ormosolenia,SiculosciadiumofcharactersmorphologicaldistinguishingtheofschemeComparative2.Table
Character Siculosciadium Ormosolenia Peucedanum s. str. Dichoropetalum Johrenia Zeravschania
Life form polycarpic polycarpic polycarpic or polycarpic or monocarpic polycarpic
monocarpic monocarpic
Taproot no no yes yes yes yes
Leaf persistence evergreen deciduous deciduous deciduous deciduous deciduous
Rootstock suffruticose not suffruticose not suffruticose not suffruticose not suffruticose not suffruticose
Caudices well developed well developed absent absent absent absent
Fibrous collar absent absent present present present present
Basal leaves, insertion apex of caudices apex of caudices apex of taproot apex of taproot apex of taproot apex of taproot
Basal leaf blade, in outline oblong to oblong-ovate reniform to ovate ovate to semicircular triangular to lanceolate triangular to ovate triangular to ovate
Basal leaf shape 2-pinnate entire to 1-pinnate ternate to multisect 1- to 3-pinnate 2-pinnate 1- to 3-pinnate
Pairs of leaflets, number 3 to 4 absent to 1 3 to 5 1 to 6 1 to 6 1 to 3
Leaflet shape ovate to deltoid reniform to ovate linear linear to ovate linear to lanceolate ovate to ovate-oblong
Cauline leaves present usually absent present present present or strongly present
reduced
Umbel rays, number 4 to 8 3 to 6 10 to 45 3 to 20 3 to 20 3 to 12
Bracts absent absent absent to 3 absent absent (1)3 to 8
Bracteoles, number 1 to 5 absent to 2 2 to 10 1 to 5 1 to 5 2 to 8
Bracteoles, texture herbaceous setaceous herbaceous herbaceous herbaceous herbaceous
Bracteoles, relative to umbellulae subequal to shorter shorter shorter usually shorter shorter subequal to shorter
Calyx teeth absent inconspicuous developed inconspicuous inconspicuous inconspicuous to
triangular
Petal color white yellowish green yellow white, yellow, pink yellow white (rarely yellowish
white?)
Stylopodium conical-flattened conical-flattened conical flat to conical flat to shortly conical shortly conical
Mericarp wings developed short developed short to well developed absent short
Dorsal ribs keeled indistinguishable filiform filiform to inflated indistinguishable filiform
Vittae, per vallecula 1 3 to 6 1 to 3 absent to 3 absent to 3 1
Commissural vittae 2 4 to 8 2 absent to 4 absent to 4 2
Secretory ducts associated with 5 usually absent absent absent to 5 absent to 5 absent
vascular bundles
Volume 99, Number 1 Brullo et al. 9
2013 Siculosciadium (Apiaceae) from Sicily
10 Annals of the
Missouri Botanical Garden
vittae per vallecula and four to eight commissural trees were restricted to the relative placements of
vittae, the secretory ducts associated with vascular several Dichoropetalum species. The strict consensus
bundles are not always present, and the stylodium is of these 10 trees is presented in Figure 3B. The
shorter than the stylopodium. Peucedanum nebro- results of this analysis support the recognition of
dense, in contrast, has a suffruticose rootstock, basal Dichoropetalum as a monophyletic group (albeit the
leaves that are evergreen, oblong to oblong-ovate in clade is weakly supported, with a 60%bootstrap
outline, and 2-pinnate with three to four pairs of value). The two species of Ormosolenia unite as a
leaflets. Its cauline leaves are present. The petiole strongly supported clade (100%bootstrap value).
has four secretory ducts interposed among the Peucedanum nebrodense is sister to the Dichoropeta-
vascular bundles, the petals are white, the mericarps lum clade, although this relationship is also weakly
have only a single vitta per vallecula and two supported (54%bootstrap value), with only four or
commissural vittae, five secretory ducts are associ- five molecular characters supporting this relation-
ated with its vascular bundles, and the stylodium is ship, depending upon the tree. Constraining both
much longer than the stylopodium. Remarkable species of Ormosolenia and P. nebrodense to
morphological differences are also apparent between monophyly and repeating the maximum parsimony
P. nebrodense and other genera included in our analysis resulted in trees just one step longer than
comparative study, namely Dichoropetalum (includ- those without the constraint invoked. These results
ing Holandrea), Johrenia, and Zeravschania. These are inconclusive in establishing whether P. nebro-
differences are presented in Table 2, but are also dense is more closely related to Dichoropetalum or to
reflected in the groupings of the UPGMA tree (Fig. Ormosolenia; indeed, P. nebrodense may actually
3A). comprise an isolated lineage separate from these two
Ormosolenia pisidica is considered by Boissier taxa upon the collapse of a single node. Further
(1849) as distinct from O. alpina by its better moleculardatawillberequiredtoelucidate
developed stems, nonthickened umbel rays, and unequivocally the phylogenetic position of P. nebro-
yellow flowers. Subsequently, Boissier (1872) trans- dense.
ferred both species to the genus Peucedanum, while
more recently Chamberlain (1972), Hartvig (1986), CONCLUSIONS
and Pimenov (1992) treated O. pisidica as a synonym The results of this study, incorporating data from
of O. alpina. In this study, both morphological and morphology, karyology, fruit and petiole anatomy, and
anatomical characters support the separation of O. ontogeny, as well as results of both phenetic analysis
pisidica from O. alpina. In particular, O. pisidica is of morphological data and phylogenetic analysis of
characterized by stems 10–20( 30) cm tall, leaves 2– combined morphological and molecular data, strongly
4 cm, umbels 3- to 4-rayed, umbellules 3- to 6- support the recognition of Peucedanum nebrodense as
flowered, petals yellow, fructiferous pedicels thin and a distinct taxon, closely allied to Dichoropetalum and
subequal to the fruit, and mericarps with filiform ribs Ormosolenia. An affinity to Peucedanum s. str. is not
and four(six) commissural vittae. Ormosolenia alpina,supported, nor is a close relationship to segregate
on the other hand, has stems 3–10( 15) cm tall, taxa such as Imperatoria and Pteroselinum. Instead,
leaves 2–10 cm, umbels 4- to 7-rayed, umbellules up maximum parsimony analysis of combined data
to 10-flowered, petals greenish white (often violet in suggests an affinity with Dichoropetalum, whereas
bud), fructiferous pedicels thickened and shorter than the UPGMA tree, based largely upon characters
the fruit, as well as mericarps with slightly inflated selected by Pimenov et al. (2007) to discern among
ribs and six to eight commissural vittae. species of Dichoropetalum,Johrenia, and Zeravscha-
nia, suggests that P. nebrodense is more similar
PHYLOGENETIC ANALYSES OF MOLECULAR AND morphologically to Ormosolenia. Of these two anal-
MORPHOLOGICAL DATA yses, we favor the maximum parsimony trees as
The data matrix of combined ITS sequence and indicating true relationship, but we acknowledge that
morphological data for 13 species comprised 544 branch support is weak and that further molecular
invariant, 43 autapomorphic, and 41 informative studies incorporating chloroplast DNA will be
positions, the latter including 17 parsimony-informa- necessary to confirm phylogenetic relationships
tive morphological characters. Maximum parsimony among these taxa. Nevertheless, Peucedanum nebro-
analysis of these combined data resulted in ten 129- dense can clearly be distinguished from Ormosolenia
step trees (consistency indices ¼0.7442 and 0.6071, and Dichoropetalum upon a variety of characters, with
including and excluding uninformative characters; many of these characters supporting its isolated
retention index ¼0.6452). Differences among these position within the group. Furthermore, P. nebrodense
Volume 99, Number 1 Brullo et al. 11
2013 Siculosciadium (Apiaceae) from Sicily
presents an isolated geographic distribution, known Genus novum Ormosoleniae Tausch affine, a qua caudice
only from one population exclusively localized in a suffruticoso sempervirenti, foliis basalibus bipinnatis seg-
high mountain doline of the Madonie Massif. No other mentis lateralibus ambitu oblongis vel oblongo-ovatis seg-
mento terminali laterales subaequenti vel quam eis minore,
species examined herein occur in Sicily or in foliis caulinis praesentibus, bracteolis umbellularum 1 ad 5
southern Italy. Therefore, we recognize P. nebrodense herbaceis margine hyalinis, dentibus calycinis nullis, petalis
as comprising the new genus Siculosciadium. In order albis colore roseo-purpurascenti suffusis apice truncatis,
to highlight the morphological differences between strato interno mesocarpii ex cellularum prosenchymatarum
this new genus and its putatively allied genera, the serie una et cellularum collenchymatarum serie una constanti,
following analytical key is provided. quaque vallecula vitta valleculari solitaria et vittis commissu-
laribus duabus praedita, canaliculis jugalibus solitariis atque
KEY TO THE CLOSELY RELATED GENERA OF SICULOSCIADIUM stylodio in fructu bene evoluto reflexo quam stylopodio per
anthesin multo longiore differt.
1a. Rootstock not well developed with numerous long
caudices, without fibrous collar; basal leaves Evergreen polycarpic dwarf shrub; rootstock
inserted at apex of caudices. branched, covered by a thin rhytidome, without
2a. Basal leaves 2-pinnate, with 3 to 4 pairs of
ovate to deltoid leaflets; cauline leaves fibrous collar; stems numerous, leafy, erect to
present; mericarps with 1 vitta per vallecula ascending, usually with umbels at the nodes. Basal
and 2 commissural vittae, with secretory leaves petiolate, inserted in the terminal part of the
ducts associated with vascular bundles caudices; sheath linear, whitish; blade 2-pinnate,
................................ Siculosciadium with 3 to 4 pairs of leaflets, terminal leaflet subequal
2b. Basal leaves entire or 1-pinnate, with up to 1
pair of reniform to ovate leaflets; cauline or smaller than the lateral ones; lobes oblong to ovate,
leaves usually absent; mericarps with 3 to 6 rounded at apex; cauline leaves becoming increas-
vittae per vallecula and 4 to 8 commissural ingly smaller upward, petiolate to sessile; blade 1- or
vittae, without secretory ducts associated with 2-pinnate; lobes linear to linear-lanceolate. Inflores-
vascular bundles ................... Ormosolenia cence composed of long pedunculate compound
1b. Rootstock robust without caudices, covered at the
top by a fibrous collar; basal leaves inserted umbels, unequally rayed; bracts absent; umbellule
directly at apex of the rootstock. with 1 to 5 bracteoles, herbaceous with hyaline
3a. Calyx teeth developed; leaf petiole round in margin, subequal to or shorter than rays. Flowers
cross section; leaf blade ternate-multisect, hermaphrodite or masculine, pedicellate; calyx teeth
tridimensional, with primary segments long
petiolate and ultimate segments entire, 2–9 absent; petals white, tinged with pink purplish
cm .......................... Peucedanum s. str. outside, inflexed, thick, truncate at apex; stylopodium
3b. Calyx teeth inconspicuous or very short; leaf conical-flattened. Mericarps lenticular, shallowly
petiole falcate in cross section with a ventral compressed dorsally with well-distinct spongy wings,
groove; leaf blade once to 3-pinnate, plane, 5-ribbed, 3 of which are dorsal and 2 marginal;
with primary segments sessile or short
petiolate and ultimate segments dentate or exocarp 1-layered; mesocarp 2-layered, its outer layer
lobate (rarely entire), 0.5–2 cm. consisting of parenchymatous cells; inner layer of
4a. Bracts (1 to)3 to 8; mericarps dorsally mesocarp consisting of 1 row of prosenchymatous
convex without occurrence of secretory cells and 1 row of collenchymatous cells; vascular
ducts associated with vascular bundles bundles 5; five secretory ducts associated with
.............................. Zeravschania
4b. Bracts absent; mericarps dorsally com- vascular bundles, 4 more are vallecular and 2
pressed usually with secretory ducts commissural not reaching the base. Stylodium in
associated with vascular bundles. fruit well developed, reflexed, and much longer than
5a. Leaf sheath very long; mericarps with the stylopodium in flower.
dorsal ribs indistinguishable and
wings absent .................. Johrenia 1. Siculosciadium nebrodense (Guss.) C. Brullo,
5b. Leaf sheath rather short; mericarps
with dorsal ribs filiform to inflated Brullo, S. R. Downie & Giusso, comb. nov.
(keeled) and wings more or less Basionym: Pteroselinum nebrodense Guss., Fl.
developed ............. Dichoropetalum Sicul. Syn. 1: 336. 1844. Imperatoria chabraei
Guss., Fl. Sic. Prodr. 1: 368.1827, nom. illeg., non
Imperatoria chabraei Spreng., Sp. Umbell.: 64.
TAXONOMIC TREATMENT 1818. Peucedanum nebrodense (Guss.) Nyman,
I. Siculosciadium C. Brullo, Brullo, S. R. Downie & Syll. Fl. Eur.: 153. 1855. Peucedanum nebrodense
Giusso, gen. nov. TYPE: Siculosciadium nebro- (Guss.) Strobl, Flora 69(36): 567. 1886. Peuceda-
dense (Guss.) C. Brullo, Brullo, S. R. Downie & num carvifolia (L.) Vill. var. nebrodense (Guss.)
Giusso Paol. in Fiori & Paol.,Fl. Anal. Ital. 2: 179. 1900.
12 Annals of the
Missouri Botanical Garden
Holandrea nebrodensis (Guss.) Banfi, Galasso & bundles 5 (3 small and dorsal and 2 big and at the
Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. wing base); 5 secretory ducts associated with the
Nat. Milano 146(2): 233. 2005. TYPE: [Italy, vascular bundles, 4 more are vallecular and 2
Sicily. Palermo:] Sub Pteroselinum nebrodense,commissural not reaching the base; endocarp 1-
Madonie, alle Fosse di S. Gandolfo, julio [manu layered; endosperm ventrally flat. Stylodium in fruit
Gussone], (lectotype, designated here, NAP- elongating to ca. 2 mm, slender, reflexed, curved, and
GUSS!; isolectotype, FI!). Figure 4. much longer than the stylopodium.
Dwarf glabrous perennial, polycarpic, suffruticose, Distribution and ecology. This Italian species is
rhizomatous, evergreen, with a well-developed creep- exclusively found in the Madonie Massif on the island
ing and branched rootstock, 4–12 cm, 2–5 mm diam., of Sicily, where it grows on the southern slopes of a
covered by a thin, pale brown rhytidome; fibrous dolina near Pizzo Carbonara, locally known as Fosse
collar absent; stems numerous, leafy, erect to di S. Gandolfo. It grows at an altitude of about 1870
ascending, 15–30 cm tall, usually with umbels at m.s.m. on Mesozoic carbonatic substrates, within a
the nodes. Basal leaves green, glaucous, 5–15 cm, dwarf shrubby plant community characterized by
inserted in the terminal part of the caudices; sheath several endemic chamaephytes and hemicryptophytes
long linear, whitish, 1–2.5 cm 33–5 mm; petiole (Brullo et al., 2005), e.g., Astragalus nebrodensis
slightly striate, 1.5–7 cm; blade oblong to oblong- (Guss.) Strobl (Fabaceae), Erysimum bonannianum C.
ovate in outline, flat, 1.5–6 31–3 cm, 2-pinnate, with Presl (Brassicaceae), Acinos alpinus (L.) Moench
3 to 4 pairs of leaflets, ovate to deltoid, 6incised, subsp. nebrodensis (Kerner & Strobl) C. Brullo &
terminal leaflet subequal or smaller than the lateral Brullo (Lamiaceae), Minuartia verna (L.) Hiem subsp.
ones; lobes oblong to ovate, rounded at apex with grandiflora (C. Presl) Hayek (Caryophyllaceae),
hyaline mucro; primary lobes 6–17 mm and second- Petrorhagia saxifraga (L.) Link subsp. gasparrini
ary lobes 3–11 mm; cauline leaves becoming (Guss.) Greuter & Burdet (Caryophyllaceae), Centau-
increasingly smaller upward with petiole 0–25 mm; rea parlatoris Heldr. (Asteraceae), Cerastium tomen-
sheath 2–5.5 cm 32–4 mm; blade lanceolate to
triangular in outline, 1- or 2-pinnate; lobes linear to tosum L. (Caryophyllaceae), Herniaria microcarpa C.
linear-lanceolate, primary ones 5–22 mm and Presl (Illecebraceae), and Silene sicula Ucria (Car-
secondary ones 5–13 mm, usually acute at apex. yophyllaceae).
Inflorescence composed of long pedunculate com- Phenology.Siculosciadium nebrodense has been
pound umbels, unequally 4- to 8-rayed, rays 0.5–1.5 observed in flower from July to August, and in fruit
cm; bracts absent; flexuous when flowering, rigid from August to October.
when fruiting; umbellule with 1 to 5 bracteoles,
subulate, herbaceous with hyaline margin, subequal Etymology.Siculosciadium comes from the
to or shorter than rays. Flowers hermaphrodite or Greek words ‘‘siculo,’’ referring to ‘‘Sicilian,’’ and
masculine, 4 to 8 per umbellule, 2–2.2 mm diam., ‘‘sciadion,’’ referring to ‘‘umbel.’’
pedicellate; calyx teeth absent; petals ovate, 1.5 mm,
white tinged with pink purplish outside, inflexed, IUCN Red List category.Siculosciadium nebro-
thick, truncate at apex; stamens long exserted, dense is represented by few individuals, usually
inrolled, wholly white; staminal filament 1.7 mm; forming a thick layer and occupying a surface area of
anther 0.5 mm; stylopodium conical-flattened; stylo- about 1000 m
2
. For this reason, we propose to include
dium ca. 1 mm long, cylindrical, erect; ovary this species in the Red List of Threatened Species as
subcylindrical to obovoid, 1–1.2 mm. Mericarps Critically Endangered (CR). Based on the criteria
subrounded to elliptical, lenticular, 6–8.5 34–5 adopted by IUCN (2001, 2003, 2005), we herein
mm, shallowly compressed dorsally with well-distinct proposed its inclusion in the following categories: CR
spongy wings, 0.5–0.8 mm wide, 5-ribbed, 3 of which B2ab(ii, iii, iv, v), C1.
are dorsal and 2 marginal; exocarp 1-layered,
interrupted near the ventral part of the wings; Additional specimens examined. ITALY, SICILY. Pa-
mesocarp 2-layered, its outer layer consisting of lermo: Madonie, Fosse di S. Gandolfo, s.d., manu Gussone,
parenchymatous cells with thin walls; in the distal sub Pteroselinum nebrodense (NAP-GUSS); in pascuis
part of the wings these cells have slightly lignified montosis Madonie, July, Citarda 1450 (FI, PAL), 1 Sep.
1881, Lojacono 223 (FI); Madonie, Fosse di S. Gandolfo e
walls with chinked pores; the inner layer of mesocarp Vallone Reale, Tineo (FI); Madonie, Pizzo Carbonara, Fosse
consisting of 1 row of tangentially elongated prosen- di S. Gandolfo, 9 Sep. 2001, S. Brullo, G. Giusso del Galdo
chymatous cells with slightly lignified walls; endocarp & S. Sciandrello s.n. (CAT); Madonie, Pizzo Carbonara,
consisting of 1 row of collenchymatous cells; vascular Fosse di S. Gandolfo, 24 July 2002, S. Brullo, G. Giusso del
Figure 4. Siculosciadium nebrodense. —A. Flowering plant habit. —B. Inflorescence. —C. Fructified umbel. —D. Male
flower. —E. Hermaphrodite flower. —F. Two petals, internal view. —G. Pistil. Living material taken from Madonie, Sicily, and
drawn from the collection, 9 Sep. 2001, S. Brullo,G. Giusso & S. Sciandrello s.n. (CAT).
Volume 99, Number 1 Brullo et al. 13
2013 Siculosciadium (Apiaceae) from Sicily
14 Annals of the
Missouri Botanical Garden
Galdo & S. Sciandrello s.n. (CAT), 30 Sep. 2009, C. Brullo, Downie, S. R., S. Ramanath, D. S. Katz-Downie & E.
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family Apioideae: Phylogenetic analyses of nuclear
Specimens examined for Ormosolenia alpina. GREECE. ribosomal DNA internal transcribed spacer and plastid
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hautes montagnes de Sphakia, June 1845, Raulin 485 (G- Apioideae (Apiaceae): Relationships among some mem-
BOISS); Creta, Distr. Sphakia, Levka Ori, in glareosis calc. bers of tribe Peucedaneae sensu lato, the placement of
montis Pachnes, 16 June 1942, Rechinger 13848 (G); Crete, several island-endemic species, and resolution within the
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Volume 99, Number 1 Brullo et al. 15
2013 Siculosciadium (Apiaceae) from Sicily
Pimenov, M. G. & M. V. Leonov. 1993. The Genera of the 6. Leaf shape: 0, linear to lanceolate; 0.5, subrounded to
Umbelliferae. A Nomenclator. Royal Botanic Gardens, reniform; 0.75, oblong to oblong-ovate; 1, ovate to
Kew. triangular.
Pimenov, M. G., M. G. Vasil’eva, M. V. Leonov & J. V. 7. Leaf sheath relative length: 0, long; 1, short.
Daushkevich. 2003. Karyotaxonomical Analysis in the 8. Leaf sheath shape: 0, linear; 1, triangular.
Umbelliferae. Science Publishers, Enfield, New Hamp- 9. Leaf basal primary segments: 0, sessile; 0.5, with petiole
shire. less than 5 mm; 1, with petiole exceeding 5 mm.
Pimenov, M. G., E. Kljuykov & T. Ostroumova. 2007. 10. Shape of terminal leaf lobes: 0, filiform or linear; 0.5,
Critical taxonomic analysis of Dichoropetalum,Johrenia,lanceolate; 1, ovate.
Zeravschania and related genera of Umbelliferae– 11. Terminal leaf lobes: 0, clustered; 1, not clustered.
Apioideae–Peucedaneae. Willdenowia 37: 465–502. 12. Upper cauline leaves: 0, absent; 1, present.
Reduron,J.-P.,A.Charpin&M.Pimenov.1997. 13. Bracts: 0, present; 1, absent.
Contribution a
`la nomenclature ge
´ne
´rique des Apiaceae 14. Mininum number of rays per umbel: 0, fewer than 5; 1,
(Ombellife
`res). J. Bot. Soc. Bot. Fr. 1: 91–104. equal to or more than 5.
Rohlf, F. J. 2000. NTSYSpc. Numerical Taxonomy and 15. Maximum number of rays per umbel: 0, ,10; 1, .10.
Multivariate Analysis System, Version 2.1. Exeter 16. Umbel rays, relative proportions: 0, very unequal; 0.5,
Software, New York. slightly unequal; 1, almost equal.
Shneyer, V. S., N. G. Kutyavina & M. G. Pimenov. 2003. 17. Bractlet texture: 0, completely herbaceous, 1, with narrow
Systematic relationships within and between Peuceda- white margins.
num and Angelica (Umbelliferae–Peucedaneae) inferred 18. Bractlet shape: 0, subulate to linear; 0.5, linear-
lanceolate; 1, lanceolate to broadly lanceolate.
from immunological studies of seed proteins. Pl. Syst. 19. Bractlets/umbellules, relative proportions: 0, bractlets
Evol. 236: 175–194. longer than umbellules, 1, shorter than umbellules.
Solov’eva,N.M.,M.Pimenov,G.Vasil’eva,N.N. 20. Calyx teeth: 0, present; 1, absent.
Mz
ˇigareva & V. D. Turkov. 1985. Karyotaxonomic study 21. Petals color: 0, white; 1, yellow.
of some species of Peucedanum (Umbelliferae). Pl. Syst. 22. Stylopodium shape: 0, flat; 0.5, short-conical; 1, conical.
Ecol. 151: 89–101. 23. Mericarps shape: 0, dorsally convex; 1, dorsally com-
Spalik, K., J.-P. Reduron & S. R. Downie. 2004. The pressed.
phylogenetic position of Peucedanum sensu lato and 24. Dorsal mericarp ribs: 0, indistinguishable; 0.5, filiform;
allied genera and their placement in tribe Selineae 0.75, keeled or slightly inflated; 1, strongly spongy
(Apiaceae, subfamily Apioideae). Pl. Syst. Evol. 243: inflated.
189–210. 25. Marginal mericarp ribs: 0, not developed; 1, 6
Strobl, G. 1886. Flora der Nebroden. Flora 44(36): 564– developed.
574. 26. Mericarp parenchyma (commissural side): 0, with
Swofford, D. L. 2003. PAUP*: Phylogenetic Analysis Using destroyed parenchyma near carpophore; 1, commissural
Parsimony (*and other methods), Version 4.0b10. parenchyma not destroyed.
Sinauer Associates Inc., Sunderland, Massachusetts. 27. Parenchyma cells with lignified pitted walls on dorsal
Winter, P. J. D., A. R. Magee, N. Phephu, P. M. Tilney, S. mericarp side: 0, developed; 1, absent or sometimes only
R. Downie & B.-E. van Wyk. 2008. A new generic under vascular bundles.
classification for African peucedanoid species (Apia- 28. Marginal ribs inflated: 0, composed of large mesocarp
ceae). Taxon 57: 347–364. cells with lignified pitted walls; 1, without lignified pitted
walls.
29. Inner lignified layer of mesocarp (hypendocarp): 0,
absent; 0.5, faintly marked; 1, strongly expressed.
APPENDIX 1. List of 36 morphological characters and 30. Secretory ducts (vittae) in mericarp furrows, presence: 0,
character states included in the morphological matrix used in absent or inconsistent, not visible in each furrow; 1,
the phenetic analysis. always developed.
31. Secretory ducts (vittae) in mericarp furrows, organization:
1. Habit: 0, monocarpic with unbranched rootstock; 1, 0, solitary; 1, several in each furrow.
polycarpic with branching rootstock. 32. Commissural secretory ducts (vittae): 0, absent; 0.5, two
2. Leaf petioles at stem base: 0, present (at least as dense in each mericarp; 1, more than two per mericarp.
remnants); 1, absent. 33. Rib secretory ducts: 0, absent; 1, present.
3. Stem height: 0, 7–40 cm; 0.5, 41–100 cm; 1, exceeding 34. Mericarp vascular bundle organization and number: 0,
100 cm. compact; 1, consisting of several groups of vascular
4. Stem ribs: 0, without prominent ribs; 1, with 6prominent elements.
ribs. 35. Leaf persistence: 0, deciduous; 1, evergreen.
5. Stem pubescence: 0, glabrous; 1, pubescent. 36. Rootstock habit: 0, not suffruticose; 1, suffruticose.
1.AppendixinthosetocorrespondstatescharactercodingandCharacteranalysis.phenetictheforusedcharactersmorphological36theofmatrixData2.Appendix
Character states for characters 1–36
Taxa 1 2 3 4 5 6 7 8 9 10 1112131415 16 17 18 192021 22 23 24 25262728 29 3031 32 3334353
Dichoropetalum achaicum (Hala
´csy) 100 001100 1 101110 00 1111 10.5 11101 101 100
Pimenov & Kljuykov
Dichoropetalum alpinum Fenzl 100 001100 1 111000 00 1110.510.75 11101 100.5100
Dichoropetalum aromaticum (Rech. 000.5100110 1 111100 00 1110.511 11000.5000 100
f.) Pimenov & Kljuykov
Dichoropetalum aureum (Boiss. & 100 001100 0.5011000 00 1110.510.75 11101 100 100
Balansa) Pimenov & Kljuykov
Dichoropetalum carvifolia (Vill.) 100.5001100 0.5101110 00 1100.510.5 11101 111 100
Pimenov & Kljuykov
Dichoropetalum chryseum (Boiss. & 000.5100100 0.5111110 00 1110.510.5 11100.5100.5100
Heldr.) Pimenov & Kljuykov
Dichoropetalum depauperatum 100.5001100.51 111000 00 1110.510.75 11101 100.5100
(Boiss. & Balansa) Pimenov &
Kljuykov
Dichoropetalum golestanicum (Rech. 000.5000100 1 101000 00 1111 10.5 11001 100 100
f.) Pimenov & Kljuykov
Dichoropetalum graminifolium 000.5001100 0.5101110 00 1100.510.75 11101 100.5100
(Boiss.) Pimenov & Kljuykov
Dichoropetalum isauricum (Parolly & 000.5001100 1 101000 10 1100.510.75 11101 100.5100
Nordt) Pimenov & Kljuykov
Dichoropetalum junceum (Boiss.) 101 001100.51 111000 00 1110.510.5 11101 100.5000
Pimenov & Kljuykov
Dichoropetalum lavrentiadis (Strid & 100 101100 0.5111100.500 1100.510.75 11100.5100.5100
Papan.) Pimenov & Kljuykov
Dichoropetalum minutifolium (Janka) 000.5010100 1 111110 00 1110 10.5 11101 111 100
Pimenov & Kljuykov
Dichoropetalum munbyi (Boiss.) 101 001100 0.5101000 00 1100.510.5 11101 100 100
Pimenov & Kljuykov
Dichoropetalum oligophyllum 000 001100 0.5101100 00 1100.510.5 11101 110.5100
(Griseb.) Pimenov & Kljuykov
Dichoropetalum palimbioides (Boiss.) 000.5010100 1 111110.500.51110.510.75 11100.5111 100
Pimenov & Kljuykov
Dichoropetalum paucijugum (DC.) 000 100100 1 111110 01 1110 11 11000.5000 100
Pimenov & Kljuykov
Dichoropetalum platycarpum (Boiss.) 100.5010100 0.5111000 00 1111 11 11000 000 100
Pimenov & Kljuykov
Dichoropetalum pschawicum (Boiss.) 100.5001100 0.5101111 00 0100.510.5 11101 111 100
Pimenov & Kljuykov
Dichoropetalum ramosissimum 001 000100 1 111000 00 1110.510.5 11001 000 000
(Mozaff.) Pimenov & Kljuykov
6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
16 Annals of the
Missouri Botanical Garden
Continued.2.Appendix
Character states for characters 1–36
Taxa 1 2 3 4 5 6 7 8 9 10 1112131415 16 1718192021 22 23 24 25262728 29 3031 32 3334353
Dichoropetalum schottii (Bess) 100.5001 100 0.5100110 001100.510.5 11101 111 100
Pimenov & Kljuykov
Dichoropetalum scoparium (Boiss.) 101 010 100 0.5101000 001111 10.5 11100.5110 100
Pimenov & Kljuykov
Dichoropetalum seseloides (C. A. 000.5000 100 0.5111110 001111 10.5 11101 100.5100
Mey.) Pimenov & Kljuykov
Dichoropetalum stridii (Hartvig) 1 1 0 1 0 1 1 0 0 0.5 1 1 1 1 0 0.5 0 0 1 1 0 0.5 1 0.75 1 1 1 0 0.5 0 0 0 1 0 0 0
Pimenov & Kljuykov
Dichoropetalum vittijugum (Boiss.) 001 000 100 1 111110 001110 10.5 11101 100.5100
Pimenov & Kljuykov
Johrenia dichotoma DC. 011 001 001 1 111000 001110.500 00001 000 100
Johrenia distans (Griseb.) Hala
´csy 010.5001 000 0.5111000 001110.500 00001 011 100
Johrenia polyscias Bornm. 010.5001 000.50.5111100 001110.510 00001 000 100
Johrenia selinoides Boiss. & Balansa 011 001 001 1 111110 001110 00 00001 000 100
ex Boiss.
Johrenia tortuosa (Fisch. & C. A. 011 001 000.50.5111100 001110.500 00001 000 100
Mey.) Chamberlain
Ormosolenia alpina (Sieber ex 110 100.5 000 1 111000 001000.510 11100.5111 100
Schult.) Pimenov
Ormosolenia pisidica Boiss. & Heldr. 110 100.5 000 1 111000 001010.510.5 11100.5111 100
Peucedanum nebrodense (Guss.) 1 1 0 1 0 0.75 0 0 0 1 1 0 1 0 0 0 1 0 1 1 0 0.5 1 0.75 1 1 1 0 0.5 1 0 0.5 1 0 1 1
Nyman
Zeravschania aucheri (Boiss.) 100.5001 111 1 110111 111100.500.5 11111 100.5000
Pimenov
Zeravschania ferulifolia (Gilli) 100.5001 111 1 110001 111001 00.5 11111 100.5000
Pimenov
Zeravschania knappii (Bornm.) 101 001 101 0 110001 001110.500.5 11111 100.5100
Pimenov & Kljuykov
Zeravschania membranacea (Boiss.) 101 001 111 0 110001 111110.500.5 11111 100.5000
Pimenov
Zeravschania pauciradiata 101 001 111 1 110001 111110.500.5 11111 100.5000
(Tamamsch.) Pimenov
Zeravschania regeliana Korovin 100.5001 111 1 110001 111100.500.5 11111 100.5100
Zeravschania scabrifolia Pimenov 100 001 111 1 110001 111101 00.5 11111 100.5000
Zeravschania stricticaulis (Rech. f.) 100.5011 100 0.5110001 111010.500.5 11111 100.5000
Pimenov & Kljuykov
6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Volume 99, Number 1 Brullo et al. 17
2013 Siculosciadium (Apiaceae) from Sicily
Appendix 4. Data matrix of the 17 morphological characters used for the phylogenetic analysis. Characters and the coding of
character states correspond to those in Appendix 3.
Taxa
Character states for characters 1–17
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Aethusa cynapium 01213102001301110
Dichoropetalum achaicum 10003102012111120
D. aromaticum 00110111011300001
D. golestanicum 00100111012110100
D. paucijugum 00010101010300001
D. pschawicum 10103100001111120
D. schottii 10103100001111120
D. scoparium 10200110012111100
D. seseloides 00100100012111111
D. carvifolia 10103100001111120
Ormosolenia alpina 11011002101011120
O. pisidica 11011002111111120
Peucedanum nebrodense 11012002001211110
18 Annals of the
Missouri Botanical Garden
APPENDIX 3. List of 17 morphological characters and 9. Calyx teeth: 0, absent; 1, present.
associated character states used in the phylogenetic analysis 10. Petal color: 0, more white; 1, more yellow.
11. Stylopodium shape: 0, flat; 1, short-conical; 2, conical.
1. Habit: 0, monocarpic with unbranched rootstock; 1, 12. Dorsal mericarp ribs: 0, indistinguishable; 1, filiform; 2,
polycarpic with branching rootstock. keeled or slightly inflated; 3, strongly spongy inflated.
2. Stem base: 0, densely covered by remains of leaf petioles; 13. Marginal mericarp ribs: 0, not developed; 1, 6
1, without remains of petioles. developed.
3. Stem height (in cm): 0, 7–40; 1, 41–100; 2, more than 14. Parenchyma cells with lignified pitted walls on dorsal
100. mericarp side: 0, developed; 1, absent or sometimes only
4. Stem ribs: 0, without prominent ribs; 1, with 6prominent under vascular bundles.
ribs. 15. Secretory ducts (vittae) in mericarp furrows: 0, absent or
5. Leaf shape: 0, linear to lanceolate; 1, subrounded to inconsistent, not visible in each furrow; 1, always
reniform; 2, oblong to oblong-ovate; 3, ovate to triangular.
6. Leaf sheath length: 0, long; 1, short. developed.
7. Leaf sheath shape: 0, linear; 1, triangular. 16. Commissural secretory ducts (vittae): 0, absent; 1, two in
8. Shape of terminal leaf lobes: 0, filiform to linear; 1, each mericarp; 2, more than two per mericarp.
lanceolate; 2, ovate. 17. Rib secretory ducts: 0, absent; 1, present.
