A phylogenetic analysis of genus Lomelosia Rafin. (Dipsacaceae) and allied taxa

Abstract

The species of genus Lomelosia (Dipsacaceae), as well as an appropriate set of outgroups, were investigated to elucidate their phylogenetic relationships based on 19 multistate morphological, palynological and karyological characters. The resulting phylogenetic trees suggest that evolution in Lomelosia proceeded from perennial to annual species, from species with prismatic epicalyx tubes to species with campanulate ones, from species with entire outer flower margins to species with flabellate ones. The majority of the characters show rampant homoplasy in the whole tree. Within the outgroup, the species of genus Scabiosa, which collapse at the base of the phylogenetic tree, are interpreted as similar to the last common ancestor of the inclusive group in study.

Full text

Delpinoa, n.s. 41: 29-45. 1999
A phylogenetic analysis of genus Lomelosia Rafin.
(Dipsacaceae) and allied taxa
OLGA DE CASTRO, PAOLO CAPUTO
Dipartimento di Biologia vegetale, Università degli Studi di Napoli Federico
II, Via Foria 223, I-80139 Napoli, Italy
Riassunto. Analisi filogenetica del genere Lomelosia Rafin.
(Dipsacaceae) e dei taxa affini
È stata effettuata un’indagine filogenetica delle specie del genere
Lomelosia (Dipsacaceae), insieme con appropriati outgroup. Sono stati
impiegati 19 caratteri multistato relativi alla morfologia, alla
palinologia e alla cariologia. Gli alberi filogenetici risultanti
suggeriscono che nell’evoluzione del genere ci sia stato un passaggio
da specie perenni a specie annuali, da specie con tubi dell’epicalice
prismatici a specie con tubi campanulati, da specie con i margini dei
fiori esterni interi a specie con margini flabellati. La maggior parte dei
caratteri mostra omoplasia elevata. Nell’outgroup le specie di Scabiosa
che si trovano, non risolte, alla base dell’albero, sono probabilmente
simili all’ultimo antenato comune di tutto il gruppo in studio.
Key Words: Dipsacaceae, Lomelosia, Phylogeny.
INTRODUCTION
Within Dipsacaceae, tribe Scabioseae includes the greatest
diversity in terms of evolutionary novelties and morphology of
single taxa. This tribe, as presently circumscribed (MAYER &
EHRENDORFER, 1999) includes the genera Lomelosia Rafin.
(=Scabiosa sect. Trochocephalus Mert. & Koch), Pterocephalus
(Vaill.) Adans. pro parte, Pycnocomon Hoffmans. & Link,
Scabiosa L. sensu stricto (=Scabiosa sect. Scabiosa), Sixalix
Rafin. (=Scabiosa sect. Cyrtostemma Mert. & Koch). The other
taxa which were formerly included in the tribe, i.e.,

Pseudoscabiosa Devesa (=Scabiosa sect. Asterothrix Font Quer),
the Himalayan species of genus Pterocephalus (now
Pterocephalodes V. Mayer & Ehrend.), Pterocephalidium G.
López, Succisa Necker, and Succisella Beck have been excluded
from the tribe on the basis of the anatomy of the diaspores
(MAYER & EHRENDORFER, 1999; 2000). Tribe Scabioseae, as
presently circumscribed, is composed of taxa whose epicalyx (a
rigid structure, synapomorphic for the family, which encases the
ovary) is radially symmetrical and roughly prismatic. This
structure is differentiated in a tube which ends in the majority of
the taxa with a membranous expansion called corona. Almost all
taxa show a diaphragma which contributes to keep the ovary, and
later the achene, in place. The area of the epicalyx tube above the
diaphragma, until the interface with the corona, is differentiated
in a region which varies in dependence with the various taxa and
is named epidiaphragma (MAYER & EHRENDORFER, 1999).
Relationships among the genera of Dipsacaceae have been the
object of various past studies (VERLÁQUE, 1977a,b; 1984a,b;
1985a,b; 1986a,b; CAPUTO & COZZOLINO, 1994). These
contributions indicated slightly different hypotheses of descent
for the taxa which belong to the “Scabiosa” morphotype (i.e.,
taxa whose involucel shows a wide corona and five calyx
bristles). In particular, according to VERLÁQUE (1986b),
Scabiosa s.s. was closely related to Pterocephalus, whereas the
other taxa (Lomelosia, Sixalix and Pycnocomon) formed a closely
knit unit. CAPUTO & COZZOLINO (1994) suggested that the whole
group of genera made a monophyletic unit, with Scabiosa s.s. at
the base, as a sister group to a clade composed of Lomelosia,
Sixalix and Pycnocomon (the latter two in a sister group
relationship).
However, recently MAYER & EHRENDORFER (1999) provided
another hypothesis, indicating a sister group relationship between
Lomelosia and Pycnocomon, as well as between Scabiosa and
Sixalix. However, all the above mentioned contributions dealt

with phylogeny at genus level and no attempt has been made to
resolve relationships below that level in the genera of Scabioseae,
regardless of the fact that some genera show evolutionary
tendencies which have been variously treated for the past as
parallelisms or synapomorphies (e.g., production of pits at the
distal end of the epicalyx tube, shape of the diaphragma and
epidiaphragma, heterocarpy).
This paper deals with a cladistic analysis of genus Lomelosia
based on morphological, karyological and palynological
characters. Lomelosia, which can be distinguished from the other
related genera for the presence of eight pits on the epicalyx tube,
includes over 50 perennial and annual species, diffused around
the Mediterranean sea, with the greatest diversity concentrated in
the eastern Mediterranean and Middle East, and outliers reaching
China and Japan.
MATERIAL AND METHODS
Taxa examined
In order to reduce the complexity of the investigation, the taxa
of genus Lomelosia have been fused in various cases in informal
groups (several of them corresponding to those of VERLÁQUE,
1986b) which share at least an exclusive synapomorphy.
A list of the investigated species and species groups follows:
L. argentea group [including L. argentea (L.) Greuter & Burdet, L.
cosmoides (Boiss.) Greuter & Burdet and L. hispidula (Boiss.)
Greuter & Burdet]
L. bicolor (Boiss.) Greuter & Burdet
L. brachiata (Sibth. & Smith) Greuter & Burdet
L. calocephala (Boiss.) Greuter & Burdet

L. camelorum group [including L. camelorum (Coss. & Dur.)
Greuter & Burdet, L. kurdica (Post) Greuter & Burdet, L. cyprica
(Post) Greuter & Burdet and L. paucidentata (Hub.-Mor.) Greuter
& Burdet]
L. candollei (Wall. Ex DC.) Soják
L. caucasica group [including L. caucasica (M. Bieb.) Greuter &
Burdet, L. sulphurea (Boiss. & Huet) Greuter & Burdet, L. balianii
(Diratz.) Greuter & Burdet, L. alpestris (Kar. & Kir.) Soják, L.
soongarica (Schrenk) Soják, L. speciosa (Royle) Soják].
L. crenata group [including L. crenata (Cyr.) Greuter & Burdet, L.
oberti-manetti (Pamp.) Greuter & Burdet, L. pulsatilloides (Boiss.)
Greuter & Burdet , L. robertii (Barr.) Greuter & Burdet, L. isetensis
(L.) Soják]
L. cretica group [including L. albocincta (Greuter) Greuter &
Burdet, L. cretica (L.) Greuter & Burdet, L. hymettia (Boiss. &
Spruner) Greuter & Burdet, L. minoana (P.H. Davis) Greuter &
Burdet, L. variifolia (Boiss.) Greuter & Burdet]
Scabiosa deserticola Rech. f. [No name is available under
Lomelosia as yet (P. Caputo, in prep.); it is informally referred to as
L. deserticola in the cladogram of Fig. 1].
L. flavida (Boiss. & Hausskn.) Soják
L. graminifolia group [including L. graminifolia (L). Greuter &
Burdet, L. epirota (Halácsy & Bald.) Greuter & Burdet, L.
hololeuca (Bornm.) Greuter & Burdet, L. pseudograminifolia (Hub.
Mor.) Greuter & Burdet, L. rhodopensis (Stoj. & Stefanov) Greuter
& Burdet]
L. leucactis (Patzak) Soják
L. micrantha group [including L. divaricata (Jacq.) Greuter &
Burdet, L. micrantha (Desf.) Greuter & Burdet]
L. olivieri (Coult.) Soják
L. polykratis (Rech. f.) Greuter & Burdet
L. prolifera (L.) Greuter & Burdet
L. reuteriana (Boiss.) Greuter & Burdet
L. rhodantha (Kar. & Kir.) Soják

L. rotata group [including L. aucheri (Boiss.) Greuter & Burdet, L.
palaestina (L.) Rafin., L. persica (Boiss.) Greuter & Burdet, L.
porphyroneura (Blakelock) Greuter & Burdet, L. rotata (M.B.)
Greuter & Burdet]
L. rufescens (Freyn & Sint.) Greuter & Burdet
Scabiosa schimperiana Boiss. & Buhse [No name is available under
Lomelosia as yet (P. Caputo, in prep.); it is informally referred to as
L. schimperiana in the cladogram of Fig. 1].
L. sphaciotica (Roem. & Schult.) Greuter & Burdet
L. stellata group [including L. stellata (L.) Rafin., L. simplex (Desf.)
Rafin.]
Scabiosa transcapica Rech. f. [No name is available under
Lomelosia as yet (P. Caputo, in prep.); it is informally referred to as
L. transcapica in the cladogram of Fig. 1].
Outgroups
As outgroups, all species of genera Pycnocomon and Sixalix,
as well as a selection of species from genus Scabiosa have been
chosen. The reason of selecting a subsample of species in the
latter genus is mainly related to the fact that over thirty names
available for segregates of S. columbaria L. have been regarded
as a single species.
A list of the outgroups follows:
Pycnocomon intermedium (Lag.) Greuter & Burdet
P. rutifolium (Vahl) Hoffmans. & Link
Scabiosa africana L.
S. columbaria L.
S. japonica Miq.
S. parviflora Desf.
S. silenifolia Waldst. & Kit.
S. tenuis Boiss.

Sixalix arenaria (Forssk.) Greuter & Burdet
S. atropurpurea (L.) Greuter & Burdet
S. cartenniana (Pons & Quézel) Greuter & Burdet
S. farinosa (Coss.) Greuter & Burdet
S. lybica (Alavi) Greuter & Burdet
S. parielii (Maire) Greuter & Burdet
S. semipapposa (DC.) Greuter & Burdet
S. thysdrusiana (Le Houérou) Greuter & Burdet
Investigated characters
The employed characters have been assessed on at least one
flowering and one fruiting specimen for each of the species listed
above. In this context, an extended loan from B and W is
gratefully acknowledged. Characters have been double checked
with the available literature sources, such as VERLÁQUE (1977b;
1984a; 1985a; 1986a,b), DEVESA (1984), HILGER & HOPPE
(1984), CAPUTO & COZZOLINO (1994), RECHINGER (1991),
MAYER & EHRENDORFER (1999).
A list of the characters follows:
1. Life form. This is a self-explicative character. A detailed
choice of states was preferred over a simple dual choice (i.e.,
perennial vs. annual) because of the fact that in various
genera, and notably within Lomelosia, a correlation may be
found between derivativeness and terophytic life style,
whereas the most archaic forms are usually fruticous. (0) =
Scapous hemicryptophyte. (1) = Suffruticous
chamaephyte. (2) = Fruticous chamaephyte. (3) = Biennial
hemicryptophyte. (4) = Scapous terophyte.
2. Radiant capitula. In the majority of the taxa in study,
capitula have zygomorphic external flowers, so that the

inflorescence has a radiant appearance. On the contrary,
capitula are haemispheric or, however, do not have a radiant
appearance in few plesiomorphic species of Lomelosia,
Scabiosa and Sixalix. Also some quite derived members of
genus Lomelosia (e.g., L. olivieri) do not have this character.
(0) = Capitula globose. (1) = Capitula radiant.
3. Number of flowers in the capitula. In the inclusive group in
study, capitula are multiflowered (i.e., with over 30 flowers
each). Only in few members of genus Lomelosia (i.e., L.
olivieri and related annual species of sect. Olivierianae) the
number of flowers in the capitula is reduced to less than
fifteen. (0) = multiflowered. (1) = pauciflowered.
4. Involucral bract connation. Involucral bracts are free in the
majority of Scabioseae. Only in the genus Pycnocomon they
are usually connate (VERLÁQUE, 1977b; 1986a). In P.
rutifolium, they are connate in their basal half and in P.
intermedium are connate only at their very base. (0) =
Absent. (1) = Basal.
5. Involucral bract length. In the inclusive group in study,
involucral bracts (i.e., the bracts which surround the
capitulum in the appearance of a calyx) are either shorter than
the radius of the head or, at maximum, equal. They are
distinctly longer only in few species of Lomelosia (i.e., in the
group of L. micrantha). This character has to be observed in
fully expanded flowering capitula only, and, for this reason, it
is scored as unknown for various taxa which have been
observed as herbarium specimens only. (0) = Shorter than
head. (1) = Longer than head.
6. External flower margin. In the taxa in which capitula are
radiant, the corolla lobes of the outer flowers are different in

shape as compared to the others. The outer margins of these
corolla lobes (which contribute the most to the overall shape
of the capitulum) may be entire or subentire (e.g., in the
majority of the outgroup taxa, in L. argentea and L.
graminifolia) or, in various taxa, crenulated (typically in the
group of L. crenata, but also in L. bicolor and in L. prolifera).
Rarely, but however in various species of Lomelosia, this
margin may be visibly flabellate (in L. calocephala, L.
leucactis, L. schimperiana). (0) = (Sub)entire. (1) = Crenate.
(2) = Flabellate.
7. Epicalyx tube pits. The epicalyx grooves are usually smooth.
The top of the epicalyx tube shows a minute pit in each
groove in Pycnocomon. In Lomelosia, the epicalyx tube is
terminated by eight deep cavities below the attachment to the
corona. (0) = Absent. (1) = Shallow depressions at the top
of the tube. (2) = Deep round or elliptical cavities at the
top of the tube.
8. Epicalyx sclerenchyma. Cross sections of the dipsacaceous
epicalyx show that the sclerenchyma is either diffuse or
organized in bundles or rings surrounding the vascular
bundles. In the investigated species, it is either present in the
form of a single ring or as a double, concentric ring (in
Lomelosia and Pycnocomon, MAYER & EHRENDORFER, 1999)
(0) = Single ring . (1) = Double ring
9. Involucel tube length. In the majority of the taxa in study,
the involucel tube is vertically developed, i.e., the length is
greater than the width. This is the plesiomorphic condition for
the whole family and, certainly, also for the taxa taken into
consideration here. However, in various taxa of Lomelosia,
the involucel tube is roundish in appearance, because width is
similar to length. This is the case, for example, of L.

brachiata, L. micrantha, L. olivieri. The epicalices of several
annual species of genus Sixalix (i.e., S. arenaria and S.
lybica) also have a roundish appearance but, in that case, the
involucel tube prolongs into the erected epidiaphragma region
and, taking into consideration the whole of the tube, the
involucel is longer than large. (0) = Longer than large. (1) =
as long as large.
10. Sulcus between pits. As already stated, genus Lomelosia
shows a definite pitting in the distal part of the epicalyx tube.
The eight resulting foveoles are separated by thin strands of
tissue. These strands may show a central furrow (e.g., in
various perennial taxa, as L. crenata, L. cretica, L. caucasica
and related species) or not, in the latter case being smooth
(e.g., in many annuals, as L. olivieri, L. rhodantha, as well as
in the group of L. argentea). The character is not applicable in
Scabiosa and Sixalix. (0) = Present. (1) = Absent.
11. Epicalyx corona. In all taxa taken into account, i.e.,
Lomelosia, Pycnocomon, Scabiosa and Sixalix, the corona is
formed by a membranaceous limb (MAYER & EHRENDORFER,
1999); this interpretation is different from that presented in
VERLÁQUE (1985a,b) and CAPUTO & COZZOLINO (1994).
Those authors, in fact, suggested that the corona in
Pycnocomon and Sixalix was woody and fenestrated;
regardless, MAYER & EHRENDORFER (1999) clearly
demonstrated that what the previous author misconstrued as a
corona was indeed part of the involucel tube, and that the
corona in Pycnocomon and Sixalix has the same nature as in
the other mentioned taxa. However, the corona is expanded in
Lomelosia and Scabiosa, but very diminutive in Pycnocomon
and Sixalix. (0) = Corona wide. (1) = Corona narrow.

12. Corona veins. The corona limb is lined by rigid veins, which
allow it to be kept expanded. These veins may merely reach
the rim of the membranous expansion (as in Scabiosa, Sixalix,
and various species of Lomelosia, notably the groups of L.
cretica, L. crenata, L. rotata) or protrude from it in the
fashion of an umbrella (as in the group of L. argentea and in
L. brachiata). (0) = Veins not excurrent. (1) = Veins
excurrent.
13. Epidiaphragma position. The studies by MAYER &
EHRENDORFER (1999) showed the relevance of the
epidiaphragma for the understanding of the reproductive
biology and systematics of Scabioseae. Such epidiaphragma
(which is present also in some taxa not taken into account in
the present study) is flat (i.e., horizontal) in Lomelosia as well
as in the majority of the species of Scabiosa, and shows a
roughly vertical disposition in Sixalix, Pycnocomon and
Scabiosa parviflora. (0) = Horizontal. (1) = Vertical.
14. Epidiaphragma length. The epidiaphragma is short in
Scabiosa, Pycnocomon and Sixalix farinosa, and long,
regardless of its position (see char. 13), in both Lomelosia and
the other species of Sixalix (MAYER & EHRENDORFER, 1999).
(0) = Short. (1) = Long.
15. Bristle position. In various of the taxa in study, calyx bristles
are expanded, so as to give a flat appearance to the calyx; in
few of them, and mainly in some species of Lomelosia,
bristles are erect. (0) = Erect. (1) = Widening.
16. Bristle size. In various taxa in study, calyx bristles greatly
protrude from the corona limb; in various species, however,
bristles are diminutive and so entirely hidden by the
membranous corona. (0) = Hidden. (1) = Evident.

Table 1 - Data matrix for the cladistic analysis. Bracketed states are
polymorphic. Question marks indicate unknown states. Dashes indicate
inapplicability. Characters are listed as in the text.
Lomelosia_candollei 110000210000011111?
L._argentea_group [03]100[01]02101010111111
L._bicolor 4[01]0001210[01]01011111?
L._brachiata 41?0?02110010111112
L._calocephala 410002210100011111?
L._camelorum 10010-210000010011?
L._caucasica_group 01000021000[01]0110111
L._crenata_group [01]100012100000111111
L._cretica_group 2100002100000100111
L._deserticola 40100-211100011111?
L._flavida 411000211100011111?
L._graminifolia_group 1100002100000100111
L._leucactis 410002210101011111?
L._micrantha_group 40001-2110010111111
L._olivieri 40100-2111000111111
L._polykratis 1100[01]0210101011111?
L._prolifera 41000121?0000101111
L._reuteriana 4100[01]1210001011111?
L._rhodantha 401000211100011111?
L._rotata_group 4[01]00[01]?21100001[01]111[012]
L._rufescens 40000-21110?011111?
L._schimperiana 310002210?00011111?
L._sphaciotica 11?00?2101010111111
L._stellata_group 4[01]00[01]?21000001[01]111[012]
L._transcapica 411000211100011111?
Pycnocomon_intermedium 01010011[012]-111010111
Pycnocomon_rutifolium 01010011[012]-111010111
Scabiosa_africana 0100?0000-001010000
Sc._columbaria_group 0100?0000-000011000
Sc._japonica 0100?0000-00000000?
Sc._parviflora 40000-000-001000100
Sc._silenifolia 0100??000-000010000
Sc._tenuis 4100??000-000011000
Sixalix_arenaria 410000001-10111100?
Si._atropurpurea [34]10000000-101111000
Si._cartenniana 1100?0000-10111100?
Si._farinosa 10000-000-001000000
Si._lybica 410000001-10111100?
Si._parielii 1000?-000-101110000
Si._semipapposa 410000000-101111000
Si._thysdrusiana 0100?0000-10111100?

17. Pollen type. Within Scabioseae, the genus Sixalix, as well as
the great majority of the species of Scabiosa has tricolpate
pollen; Lomelosia, Pycnocomon and Scabiosa parviflora have
triporate pollen (VERLÁQUE, 1985a; 1986a,b). (0) =
Tricolpate. (1) = Triporate.
18. Aperture ornamentations. Sixalix and Scabiosa parviflora
have apertural membranes (VERLÁQUE, 1985a; CAPUTO &
COZZOLINO, 1994; MAYER & EHRENDORFER, 1999).
Lomelosia, Pycnocomon, as well as the other species of genus
Scabiosa have opercula. (0) = Membrane. (1) = Operculum.
19. Chromosome haploid number. The chromosome haploid
number in Pycnocomon and Lomelosia p.p. is n=9, in
Lomelosia brachiata is n=7, and in Scabiosa and Sixalix is
n=8 (VERLÁQUE, 1977a, 1985a, 1986a,b). Some
polymorphism occurs in Lomelosia because of dysploid
series. (0) = Nine. (1) = Eight. (2) = Seven.
The resulting matrix (41 terminals, 19 characters) (Tab.1) was
analyzed by using the cladistic software environment Winclada
(NIXON, 1999), running Nona (GOLOBOFF, 1993-99) as a
daughter process, with the following parameters: hold 100000;
hold/100; mult*100; max*, and treating all multistate characters
as unordered. The resulting cladograms were investigated with
Winclada.
RESULTS
After the cladistic analysis, 27 maximum parsimony
cladograms were obtained (length = 55, C.I. = 0.47, R.I. = 0.83),
the strict consensus of which is shown in Fig. 1. The tree has
been rooted by using Scabiosa and Sixalix, leaving Pycnocomon

in the

Fig. 1 - Consensus tree for the taxa in study out of 27 maximum parsimony
cladograms (length = 55, C.I. = 0.47, R.I. = 0.83). Black dots indicate
synapomorphy; white dots indicate homoplasy. Numbers above each dot are
the character numbers as indicated in the text and in the matrix. Numbers
below dots represent the state at the internode.

ingroup. The topology of the ingroup shows Pycnocomon as
sister group to Lomelosia. Pycnocomon is monophyletic in
having connate involucral bracts (char. 4), as well as characters
pertaining to the corona and epidiaphragma (chars. 11-13). The
latter three characters, however, behave as synapomorphies only
locally. Lomelosia is monophyletic in having a wide
epidiaphragma (char. 14). Such character is, however, only
locally synapomorphic, because it develops in the derived
members of genus Sixalix in a parallel fashion. Topology in
genus Lomelosia is not entirely resolved (Fig. 1). However,
several clades are well defined. The genus has a basal clade made
of the group of L. caucasica, then a clade composed of L.
camelorum, and the species of the L. cretica and L. graminifolia
groups. All members of this clade show a reduction of the calyx
bristles (char. 15). The more internal clade has a basal, local
synapomorphy, also related to calyx (char. 16, widening
bristles). This clade shows L. candollei basally and then L.
crenata, the latter sister being group to an inner clade composed
(primarily) of annual species (char. 1). The character, however,
reverts once in some apomorphic species. The remaining species
and species groups, which are of primarily Eastern distribution,
are mainly characterized by excurrent corona veins (char. 12,
reverting later). In this clade, besides two basal collapses of two
central and Eastern Mediterranean species each, two clades are
visible, one including the group of L. argentea, L. polykratis, L.
sphaciotica, L. leucactis, L. calocephala and L. schimperiana
and the other with L. brachiata (the only species of the genus
with ten calyx bristles), the group of L. micrantha, the group of L.
rotata as sister group to the annual Eastern sect. Olivierianae
(Rech. f.) V. Mayer & Ehrendorfer. This section is composed by
small, pauciflowered (char. 3) annuals.
In the outgroup, no synapomorphy keeps genus Scabiosa
together. Genus Sixalix, on the contrary, is characterized by a
vertical epidiaphragma (char. 13). It is to be noted that Scabiosa

parviflora nestes together with one plesiomorphic member of
genus Sixalix (S. farinosa).
DISCUSSION
The phylogenetic analysis of genus Lomelosia allows
recognition of various evolutionary trends, some of which had
already been identified, although on an intuitive basis, in previous
literature. The most plesiomorphic species of the genus, for
example, are scapous hemicryptophytes (i.e., the group of L.
caucasica). Several species of the genus then evolved to
(sub)fruticous chamaephytes. A definite shrubby habit is present
only in the group of L. cretica; suffruticous habit, however is
rather diffuse in the genus (e.g., L. camelorum, L. crenata, L.
graminifolia, L. polykratis, L. sphaciotica), and probably
developed in an independent fashion at least twice. The great
majority of the most apomorphic taxa, however, is annual.
Many species, both in genus Lomelosia and in the outgroups,
have radiant capitula. The occurrence of radiant capitula, which is
apomorphic in the family (CAPUTO & COZZOLINO, 1994), is
however plesiomorphic in the inclusive group in study.
Regardless, globose capitula are present in several taxa and, in
particular, in several species of the outgroup (Scabiosa
parviflora, Sixalix farinosa, Sixalix parielii) as well as in the
plesiomorphic L. camelorum. For these taxa the globose capitula
seem to have been apomorphically acquired; however, it is
difficult to state whether the condition is truly apomorphic or not
(it may depend upon our outgroup choice). Globose capitula,
however, are also present (in one case the character is
polymorphic) in several derived species of Lomelosia (the group
of L. micrantha, the group of L. rotata, as well as in L. rufescens,
L. deserticola, L. olivieri, L. rhodantha). The change in the
overall shape of the capitulum may be related to pollinator shifts,

in particular for the last mentioned group of species, in several of
which a reduction in the number of flowers in the capitulum
occurs.
Outer florelets margins modify their shape throughout the
genus. The most plesiomorphic species have entire flower
margins, whereas several independently evolve crenate margins
(e.g., the group of L. crenata or L. reuteriana) or even visibly
flabellate margins (L. leucactis, L. calocephala, L. schimperiana).
Another character which can be followed throughout the genus is
related to the overall proportions of the epicalyx tube, which is
roughly cylindrical in most species, but becomes campanulate
(i.e., as long as large) in L. brachiata, L. micrantha, L. olivieri
and few other species.
As far as the outgroups are concerned, we would like to point
out that the lack of synapomorphies within genus Scabiosa s.s. is,
in our opinion, not an artifact of our analysis. In fact, this genus
developed an expanded, membranous corona which, however, is
also present in all other genera taken into account (as well as in
others which are not present in this investigation). Scabiosa
probably represents a successful body plan, which has been
further exploited in Lomelosia and Sixalix. From some ancestor
which had to be similar to present-day plesiomorphic species of
the genus (e.g., Scabiosa japonica), Lomelosia, Sixalix,
Pycnocomon, as well as the most derived species of Scabiosa
have probably originated.
A clear artifact of our analysis, on the contrary, is the sister
group relationship between Scabiosa parviflora, an annual and
aberrant species from Sicily, Italy, and Sixalix farinosa. The
presence of S. parviflora within genus Sixalix may derive from
the fact that some of its autapomorphies are misconstrued as
synapomorphies; most likely, however, a deeper investigation
would bring also Sixalix farinosa to collapse, together with the
species of Scabiosa, at the base of the tree. Sixalix farinosa, in
fact, is a very plesiomorphic species, and may represent the

ancestor of all the other species of Sixalix (MAYER &
EHRENDORFER, 1999). Eventually, Pycnocomon, a genus which
was regarded in the past as closely related to Sixalix (VERLÁQUE,
1986a; CAPUTO & COZZOLINO, 1994), appears from this analysis
as sister taxon of Lomelosia. This depends on the reassessment of
the homology of the epicalyx parts carried out by MAYER &
EHRENDORFER (1999), who showed the close resemblance
between the two genera.
The results shown here, in terms of genus-level topology,
coincide, to the extent of the taxa in common, with preliminary
results based on molecular characters (Caputo et al., in prep.).
In conclusion, genus Lomelosia, as well as the related
Scabiosa, Sixalix, and Pycnocomon, are taxa beset with
parallelisms (as shown by the very low C.I. of the cladograms),
which may often obscure true synapomorphy. Various of the
tendencies which are observed in one of the genera, develop also
in the others in a parallel fashion. This, up to recent times, has
prevented a clear understanding of the phylogenetic relationships.
Ongoing studies, based on molecular characters, will allow
further insights in the relationships of some of the taxa (mainly,
Scabiosa parviflora and Sixalix farinosa, but also all the other
species of Scabiosa) which have not been elucidated here.
Abstract. The species of genus Lomelosia (Dipsacaceae), as well as an
appropriate set of outgroups, were investigated to elucidate their
phylogenetic relationships based on 19 multistate morphological,
palynological and karyological characters. The resulting phylogenetic
trees suggest that evolution in Lomelosia proceeded from perennial to
annual species, from species with prismatic epicalyx tubes to species
with campanulate ones, from species with entire outer flower margins
to species with flabellate ones. The majority of the characters show
rampant homoplasy in the whole tree. Within the outgroup, the species
of genus Scabiosa, which collapse at the base of the phylogenetic tree,
are interpreted as similar to the last common ancestor of the inclusive
group in study.

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Finito di stampare nel Febbraio 2001