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Systematics of the Complex Pulmonaria saccharata-P. vallarsae and Related Species (Boraginaceae)

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Abstract and Figures

The complex Pulmonaria saccharata-P. vallarsae has been studied by means of morphological, biometric and biosystematic analysis of dried and living specimens, in natural environments and under cultivation. Hydroponic cultures have been used to distinguish the effects of environmental conditions and of phenological phases in determining phenotypic plasticity. An electrophoretic study of the seed proteins has been done. Inter- and infra-specific crossability has been experimentally tested. Chromosome counts on populations of all italian species have been made.
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~ Webbia 51(1): 1-20. 1996
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~ Systematics of the Complex Pulmonaria saccharata-P. vallarsae and
Related Species (Boraginaceae) ,',
GIOVANNAPUPPI and GIOVANNICRISTOFOLINI
Dipartimento di Biologia Evo!. Sper., Università di Bologna
Via Irnerio 42, 1-40126 Bologna.
Accettato per la stampa il
1
aprile 1996
Sistematica del complesso Pulmonaria saccharata-P. vallarsae e delle specie affini
(Boraginaceae). - Pulmonaria saccharata Miller e specie affini sono state studiate
per mezzo di analisi morfologiche e biometriche. Sono stati analizzati sia campioni
d'erbario che campioni in coltivazione; la coltivazione è stata condotta sia in terra
che in idrocoltura, ond~ distinguere la variabilità genetica da quella direttamente
indotta dall'ambiente. E stata studiata pure la variabilità nella composizione delle
proteine di riserva dei semi. Prove di infertilità infra- ed inter-specifica sono state
fatte in condizioni controllate. Conte cromosomiche sono state fatte su popolazioni
di tutte le specie presenti in Italia. La ricerca ha portato a riconoscere le seguenti
specie:
- P. saccharata Miller, da escludersi dalla flora italiana; si tratta di una specie
presente soltanto nell'Europa settentrionale (Belgio e, dubitativamente, Olanda).
- P. affinis Jordan, strettamente correlata a P. saccharata, rappresenta questo
complesso nella Francia ad ovest del Rodano ed in Spagna, nei Pirenei.
- P. picta Rouy
(=
P. saccharata Auct. Fl. Ita!.), distribuita dalla Francia Sud-
orientale all'Italia, prevalentemente sul versante tirrenico, dalle Alpi Marittime al
Lazio.
- P. vallarsae Kerner, endemica delle Prealpi Trentine e Venete.
- P. apennina sp. nov.
(=
P. vallarsae Auct. Fl. Ital., p. p.), endemica
dell'Italia, diffusa dall'Appennino settentrionale alla Calabria, gravitante soprat-
tutto sul versante adriatico.
Le specie sono parzialmente interfertili; tuttavia la specializzazione ecologica e
la segregazione geografica determinano un consistente isolamento genetico. L'iden-
tificazione, sempre problematica, può essere fatta con certezza solo sull'indumento
delle foglie basali estive.
Key words: biosystematics, Boraginaceae, chorology, flora, Italy, protein
polymorphism, Pulmonaria, taxonomy.
The species of Pulmonaria
L.
are highly polymorphic, and intergrade the
one into the other along complex patterns of variation. Hence, De Candolle is
quoted to have remarked that «de Pulmonariis tot capita tot sensus»
(KERNER,
1878), and Lacaita to have stated that «le Pulmonarie fanno disperare» (PIG-
NATTI,
1982).
,', Research financed
by
MURST (40%, project: "Biosistemarica e corologia delle piante") and CNR
(project: "Flora mediterranea: sistematica e corologia di generi critici").
)
/
2
G. PUPPI and G. CRISTOFOLINI
After KERNER'S (1878) exhaustive monograph, hardIy any of the folIowing
treatments gained a generaI consensus. TARNAVSCHI (1935) first used the
caryological analysis to interpret the apparently chaotic variation. FolIowing his
example, MERXMULLER & GRAU (1969), MERXMULLER & SAUER (1972), SAUER
(1975, 1979), BOLLIGER (1982) also gave a remarkable weight to the caryotype
in order to define systematic patterns. Nevertheless, not alI conclusions
reached so far are convincing - partly due to the high infraspecific variation.
The relationships of
P. saccharata-complex
within the other species is contro-
versial. SAUER (1975, 1979) assumed that
P. saccharata
is related to the western
species
P. longifolia, P. affinis
and
P. montana,
whereas
P. vallarsae
would be
derived from the eastern complex
P. stiriaca-P. carnica-P. rubra.
At divergence with SAUER'S views, the group of
P.saccharata
is intended
here in a broad sense, as to include not only
P. saccharata s.l.,
but
P. vallarsae s.l.
as well. We agree indeed with MERXMULLER & GRAU (1969) and MERXMULLER
& SAUER (1972) in observing that
P. vallarsae
is related to the cycle of
P.
saccharata
by a remarkable morphological affinity, and by intermedia te forms.
The
P. saccharata-complex
seems the result of a recent radiation. Distin-
guishing the species by reliable diagnostic characters is often difficult (LA-
CAlTA, 1921, PUPPI & CRISTOFOLINI, 1991). AlI species of this complex have the
lamina of the basaI leaves more or less abruptly narrowed into petiole, never
cordate, more or less distinctly spotted, more or less densely covered by long
setae,
short
puberes
and glandular hairs. All species occur in mesic broad-Ieaved
forests
(Fagetalia, Quercetalia robori-petraeae, Quercetalia pubescentis);
their geo-
graphic range is from w-(sub)mediterranean to subatlantic. The generaI distri-
bution ranges from Southern Italy to North-eastern France and Belgium,
through the Apennines, the Maritime Alps and CentraI France; such a distribu-
tion pattern seems to match with the pattern of the quaternary migration from
the refuge-area in Southern ltaly toward Northern Europe. The range is
interrupted at some pIace, e.g. by the Po plain: such interruption is possibly
due to the deforestation during the late Middle Age.
Most chromosome counts gave a soma tic number 2n
=
22, that has been
interpreted by TARNAVSCHI(1935) as an hypertriploid on a base number
x=
7.
An exception is given by some populations of the Apennines, with
2n
=
28,27,26, possibly a tetraploid level, with derived aneuploidy. No specific
morphological traits are correlated with such cytotypes, whose origin is rather
puzzling.
The following taxa were studied in the present research: 1)
P. saccharata
Miller
sensu stricto
(see PUPPI & CRISTOFOLINI, 1991); 2)
P. affinis
Jordan; 3)
p.
picta
Rouy (=
P. saccharata
Auct.fUtal. non Miller); 4)
P. vallarsae
Kerner
sensu
stricto,
an endemie of Italian Eastern Pre-Alps; 5) The apenninic race of
P.
vallarsae sensu lato,
that is here described as the new species
P. apennina.
S
METHODS
a)
Morpholo
object of morpl
herbaria were s
SIENA,TSB, \
To detect e
P. vallarsae
and
in the Garden,
solution (Lewa
following locali
Baccio (Moden~
(Trento); Vallar
(Cuneo); BologI
Cervati (Salernc
Several indi
detect somatic \
Macroscopic
(especially colol
exsiccata. The
(tab. 1).
Four trichorr
mm), usualIy sI
stipitate multict
stipitate, unicell
summer leaves,
unit areas of 0.7
per leaf.
b)
Protein ani
picta,
of
P. valla
environments, a
(open ground, 9:
Brasimone (wooc
1150 m, Trento.
Casalecchio (woc
Prati di Mugnanl
Nutlets prodl
together. Pericatj
with quartz sand;
buffer pH 8.0, ~
1efollowing
st used the
~ollowinghis
972),
SWER
he caryotype
conclusions
fic variation.
.es is contro-
I
the western
ae
would be
~i.sintended
'. vallarsae
S.l.
I1ERxMuLLER
~cycle
of
P.
diate forms.
tion. Distin-
!ifficult (LA-
llex have the
=tiole, never
~red by long
)road-leaved
IS);
their geo-
eneral distri-
nd Belgium,
:h a distribu-
gration from
"he range is
n is possibly
at has been
1mber x=
7.
nines, with
. No specific
gin is rather
P.
saccharata
rordan; 3)
P.
Kerner
sensu
c race of
P.
~nnina.
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
3
METHODS
a)
Morphology and biometry -
Herbarium specimens of alI species were the
object of morphological and biometrical analysis. Exsiccata from the following
herbaria were studied: BM, BOLO, BR, FI,
K,
LY, NAP, OXF, PAVIA, RO,
SIENA,TSB, VER, WU, ZT.
To detect environment-controlled variability, living specimens of
P. picta,
P. vallarsae
and
P. apennina
were coIIected in natural stands and cultivated: 1)
in the Garden, in pots in the open, under uniform conditions; 2) in hydroponic
solution (Lewatit-Bayer) in the greenhouse. Collections were made at the
following localities: a)
P. picta -
Vergato (Bologna); Riola (Bologna); Lago
Baccio (Modena); Monti Cimini (Viterbo); b)
P. vallarsae -
Pian delle Fugazze
(Trento); Vallarsa (Trento); Valle Agordina (Belluno); c)
P. apennina -
Priero
(Cuneo); Bologna; Corniolo (Forlì); Senarica (Teramo); Palena (Chieti); Monte
Cervati (Salerno).
Several individuals were vegetatively developed from single plants, to
detect somatic variation due to environmental conditions.
Macroscopic and microscopic characters were recorded on fresh plants
(especiaIIy colour of corolla and colour of the spots on the leaves) and on
exsiccata. The leaves characters were recorded on leaves of the same age
(tab. 1).
Four trichome types were separately recorded:
bristles
(=
setae):
long
(>
0.5
mm), usuaIIy stiff hairs;
puberes:
short hairs
«
0.5 mm);
glandular hairs:
stipitate multicellular glands, about as long as setae;
microglands:
shortly
stipitate, unicelluiar glands. Trichomes were counted on the upper surface of
summer leaves, using a stereomicroscope (60 to 90 magnif.); on each leaf 20
unit areas of 0.7 mm
2
were recorded. The length was measured on 50 trichomes
per leaf.
b)
Protein analysis -
Mature nutlets were collected from wild plants of
P.
picta,
of
P. vallarsae,
and of
P. apennina.
Collections were made in different
environments, at the foIIowing localities: a)
P. picta -
Serra del Zanchetto
(open ground, 920 m, Bologna), Granaglione (open ground, 600 m, Bologna),
Brasimone (wood, 860 m, Bologna); b)
P. vallarsae -
Vallarsa (open ground,
1150 m, Trento); c)
P.apennina -
Pieve del Pino (wood, 350 m, Bologna),
Casalecchio (wood, 90 m, Bologna), San Lazzaro (clearing, 60 m, Bologna),
Prati di Mugnano (wood, 290 m, Bologna).
Nutlets produced by different plants of the same population were pooled
together. Pericarps were manuaIIy removed. The seeds were ground in a mortar
with quartz sand; the flour was suspended in
lO
parts (v/w) 0.2M Tris-Glycine
buffer pH 8.0, and centrifuged (15' at 15,000 rpm). The liquid phase was
4
G. PUPPI
and
G. CRISTOFOLINI
Length of lamina (mm)
Width of lamina (mm)
WL/LL
Shape of the base of lamina (angle, in degrees)
Distance of maximum width of lamina from the base (mm)
DW/LL
Length of petiole (mm)
LP/LL
Width of petiole at its middle, incl. wings (mm)
Width of petiole excl. wings (mm)
WP/LL
Thickness (dorsiventral) of petiole (mm)
WPP/TP
Width of petiole wings (mm)
Colour of petiole (green to violet, in four classes)
Presence and evidence of bright spots (from absent to very sharp, in four classes)
Dimension and confluence of spots (in five classes)
BIade margin (from flat to undulate, in four classes)
Density of glandular hairs (no. per mm')
Density of microglands (no. per mm')
Density of
setae
(no. per mm')
Density of
puberes
(no. per mm')
Middle length of indumentum (mm)
A sarr
different
pots. Th
insects fl
flowers. '
alI throu
flowerin
t
counted.
As
eal
produced
interferti
of the re~
The
SI
under la!:
ate gradu
TABLE1 - Macroscopic and microscopic characters recorded on basaI leaves. The characters
marked with an asterisk (,,) present significant differences between species.
LL "
WL
NWL'"
SE
DW
NDW
LP
NLP
WP"
WPP
NWP
TP ,',
NP
WW'"
CP "
EM"
DM'"
OND"
DGH'"
DMI ,',
DS "
DP "
LI ,',
d)
Ch
Radical
colchicin
were squ
.
.
lmmerslO
emulsified with Freon (15', ratio 4:1) to separate lipids. The lipophylic phase
was removed with a micropipette, the hydrophylic phase was re-centrifuged as
above and filtered through a column (cm 0.9 x 25) of Sephadex G-25, at a flow
rate 0.8 mI/min. The eluate was monitored at 280 nm. The protein containing
fraction was salted out (50% saturation (NH4
)2
S04). After centrifuging (15'
at 10,000 rpm) the pellet was resuspended in distilled H20 and desalted by gel
filtration as above. The samples to be analysed by electrophoresis were added
with 1% Sodium Dodecylsulphate and 1% ~-mercaptoethanole. Polyacryla-
mide gel electrophoresis was done under different conditions (homogeneous
gels 12.5%, and gradient gel 8-25% and 10-15%) using a Phast-Gel Pharmacia
apparatus.
The phenetic similarity between patterns was computed as the ratio
c/(a
+
b
+
c),
where
a
and bare the number of protein bands present in only one
pattern, and c is the number of bands common to both patterns. Clustering was
done by UPGMA.
e)
Ph,
were stue
basallem
(MARCEL
f)
5yr.
taxon w(
behaviou
sociologi<
VV.AA.,
RESULTS
a) Mc
the flow
corolla a
P. picta (
be easily
c)
Crossing experiments -
AlI species of
Pulmonaria
are distylous self-
incompatible. Cross pollination is mainly due to bees, but also to ants.
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
5
The characters A sample of 48 plants of
P. picta, P. vallarsae, P. apennina
were collected at
different stands in the Northern Apennines and the Alps, and cultivated in
pots. The infIorescences were protected by a thin tissue bag, that prevented
insects from enter but allowed air to move through and light to reach the
fIowers. The fIowers were hand poIlinated using a thin brush, every other day
alI through the fIowering time. At the end
oE
the reproductive time the
fIowering scapes were collected, and the number of nutlets produced was
caunted.
As each fIower can produce up to four nutlets, the ratio (number
oE
nutlets
produced)j(max. theoretical number of nutlets) was assumed as a measure of
interfertility. Only fully developed nutlets were counted. The variance analysis
of the results was done.
The seed germinability was not tested, as germination
oE Pulmonaria
seeds
under laboratory conditions is extremely difficult to obtain, and seeds germin-
ate gradually, during several years (see also BOLLIGER, 1982).
lur
classes)
d)
Chromosome counts -
All counts were made on somatic metaphases.
Radical tips developed from rhyzomes in hydroculture were treated with
colchicine (Fluka) and stained in leucobasic fuchsin (Merck); the radical tips
were squashed on microscope slides, mounted with Euparal, and observed in
immersion at 1500x.
phylic phase
:ntrifuged as
25, at a fIow
11
containing
"ifuging (15'
;alted by gel
were added
Polyacryla-
)mogeneous
l Pharmacia
e)
Phenology -
The phases of vegetative and reproductive development
were studied on plants cultivated in the Botanical Garden. The growth rate of
basalleaves was recorded. The phases
oE
reproductive processes were observed
(MARCELLO, 1935, PUPPI BRANZI, 1989).
s the ratio
in only one
Jstering was
E) Synecology and chorology -
The geographic distribution data of each
taxon were derived mainly from herbarium records. The phytosociological
behaviour was defined from field observations and from a screening
oE
phyto-
sociological literature (FEoLI
&
LAGONEGRO, 1982, UBALDI et al., 1987,
VV.AA., 1995).
RESULTS
;tylous self-
nts.
a)
Morphology and biometry -
The morphology
oE
the fIoraI scape and
oE
the fIowers are
oE
little use to discriminate between species. The colour of
corolla and its internaI hairiness are differential, e.g. between
P. saccharata
and
P. picta
(PUPPI
&
CRISTOFOLINI, 1991). Unfortunately, these characters cannot
be easily evaluated in dry specimens.
6
G. PUPPI
and
G. CRISTOFOLINI
The basaI leaves present a remarkable variability. The macroscopic morpho-
logy (shape, dimension, presence of white spots) varies in relation with the
phenological phase and the environmental conditions (tab. 2). On the contrary,
the indumentum of fully developed basaI leaves ("summer leaves") is confirmed
as the most reliable among all morphological traits. The number of hairs and
the number of glands per uni t area, the aver age length of hairs and their
distribution in classes of length, are characters which proved constant, inde-
pendent of growth conditions, and are significantIy different from species to
species (tab. 3).
The average length and the density of hairs are negatively correlated with
each other. The indumentum of
P. vallarsae
and of
P. apennina
is short and
dense, whilst in
P.
picta
it is long and sparse.
P.
saccharata
and
P.
affinis
are
intermediate under this aspect.
T
ABLE
2 - Variance and covariance of the morphological characters in relation to phenological
phase and edaphic conditions. Only characters whose values are significantly different (P
=
0.05)
between species are listed.
Column
1:
characters (abbreviations as in tab. 1); columns 2-4: mean value and standard deviation
in the three italian species; column 5: significance of the difference between leaves of the same plant
developed in different seasons (n.s,
=
not significant; ,',
=
significant for p
=
0.05; ,',,',
=
significant
for P
=
001); column 6: significance of the difference between plants of the same clone grown under
different edaphic conditions (symbols as above).
TABLE
3 -
p, apennina.
A
(
l
]'I
V
T
~
C
El
Di
01
D~
DJ
m
DI
LI
b)
Protein
electrophoret
P. vallarsae, P
Protein pattel
the same spec
lower similari
terns of differ
single polype]
species. The (
species are of
but the apenn
each other th~
1
2
3
4
5
6
Characters Ppicta P vallarsae Psaccharata
LL
118.3 +/- 31.7 133.5 +/- 38.6 107.4 +/- 30.2 ns
**
NWL 0.40 +/- 0.06 0.50 +/- 0.11 0.48 +/- 0.08
**
IlS
WP 5.6+/-2.7 6.5 +/- 3.1 4.5 +/- 1.5
IlS
*
TP
2.2 +/- 0.5 2.5 +/- 0.7 2.1 +/- 0.6
IlS
**
WW 1.7 +/-
l.
l
I.
9 +/- 1.3 1.2 +/- 0.6
IlS
*
CP 1.7 +/- 0.9
l.3
+/- 0.6 2.0 +/- 1.0
** **
EM 2.7+/- 1.3 1.7 +/- 0.8 2.0 +/- 1.0
***
DM 2.2 +/- 1.6 0.7 +/- 0.7
1.3
+/-
l.3
IlS
*
OND 2.0 +/- 0.9 2.5 +/- 0.6 2.2 +/- 0.6
IlS
**
DGH 1.32 +/- 0.54 1.46 +/- 0.38 0.62 +/- 0.30
IlS IlS
DMI 0.68 +/- 0.19 0.47 +/- 0.11 0.94 +/- 0.28
IlS
llS
DS 2.85 +/- 0.97 1.74 +/- 0.44 1.84 +/- 0.43
IlS IlS
DP 13.20 +/- 0.77 31.64 +/- 6.11 32.85 +/- 4.83
IlS
**
LI
0.24 +/- 0.02 0.13 +/- 0.03 0.16 +/- 0.02
** *
c)
Crossing
to produce fu
is generally hi
species (see a
terspecific fe
cantly lower
t
cies. Most ren
produced by
c
nina
is signif:
produced by
f
Alps crossed
I
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
7
pie morpho-
)il
with the
ne con trary,
.s confirmed
)f hairs and
sand their
stant, inde-
1
species to
TABLE
3 - Macroscopic and microscopic characters: pairwise comparison of
p,
picta, P. vallarsae,
P. apennina. Abbreviations as in tab.
1.
""""~~~~~~,
Characters
Ppicta
vs .
Ppicta
vs.
P vallarsae
vs.
P vallarsae Papennina Papennina
LL
ns ns
**
NWL
** **
ns
WP
ns ns
**
TP
*
ns
*
WW
ns ns
**
CP
ns ns
**
EM
** *
ns
DM
** *
ns
OND
**
ns ns
DGH
ns
** **
DMI
* ** **
DS
** **
ns
DP
** **
ns
LI
** ** *
'elated with
s short and
) affinis
are
o phenological
rent (P=005)
dard deviation
the same pIan t
=
significan t
~ grown under
b)
Protein analysis -
The variability of
electrophoretic patterns (fig. 1) of
P. picta,
P. vallarsae, P. apennina
has been analysed.
Protein patterns of different accessions of
the same species are similar. By contrast, a
lower similarity is observed between pat-
terns of different species, although no one
single polypeptide is differential of any
species. The distances between the three
species are of the same magnitude (fig. 2)
but the apenninic taxa are more similar to
each other than to the alpine
P. vallarsae.
+
6
lS
**
,*
ns
*
**
*c)
Crossing experiments -
The ability
to produce fully developed fruits (tab. 4)
is generally high, both within and between
species (see also BOLLIGER,1982). The in-
terspecific fertility is, however, signifi-
cantly lower than the fertility within spe-
cies. Most remarkably, the amount of seed
produced by crossing
P. picta
with
P. apen-
nina
is significantly higher that the seed
produced by plants of
P. vallarsae
from the
Alps crossed with plants of
P. apennina.
*
**
**
S
*
IS
**
IS
ns
IS
ns
IS
ns
ap
va pl
*
*
Fig. 1 - A sample of the electro-
phoretic patterns of the seed globulins of
p.
apennina (ap),
p
vallarsae (va), p, picta
(pi),
.s
**
8G. PUPPI and G. CRISTOFOLINI
Th
1ll
J
-
>-
....,
.....
0.5
m
E
(J)
• P.
VALLARSAE
~ P. APENNINA
~ P.
PICTA
spe
rytl
apr
var
ane
SlgI
nUTI
fIo\!
nu
22,
or
o
I
Fig. 2 - Dendrogram of phenetic similarity among 8 accessions of the three ltalian species, based on the
electrophoretic pattern of seed globulins, at a
on I
rem
lea\
TABLE
4 - Results of crossing experiments among
p,
vallarsae,
P.
picta
and
p,
apennina.
In each
box: CentraI figure
=
Number of nutlets produced per 100 flowers; upper right
=
absolute number of
nutlets produced; lower left
=
number of plants; lower right
=
number of flowers pollinated.
~
P.va//arsae P.apennina Ppicta
Acceptor
353 164 280
P.va//arsae
69
48
55
12 511 13 345 10 510
178 203 73
P.apennina
39 63
49
7460 3324 6149
53 154 90
P.picta
51 45
58
5104 8345 5156
~,
j
alI
S
3, 4
alor
mOl
Eas
(Fag
stag
bria
Ap(
ased on the
~a.In each
number of
ted.
fa
280
510
73
149
90
156
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
9
The geographic segregation determined by the Po plain has obviously resulted
in a significant reduction of interfertility.
d)
Chromosome counts -
Our counts were consistent with the previously
known data (BoLLIGER,1982, CAPINERI,1986, MERXMULLER
&
GRAU, 1969,
SAUER,1975). AlI populations of
P. vallarsae
and
P. apennina
have a somatic
number
2n
=
22, whilst
P. picta
is composed of several caryological races
(2n
=
22, 26, 28), whose distribution does not seem to correlate with any geographic
or ecological pattern.
e)
Phenology -
The observations were limited to the Italian taxa. Inter-
specific differences were detected both in the anthesis and in the vegetative
rythm. Anthesis in
Pulmonaria
usually begins in february-march and lasts until
apriI or even early may. The number of flowers and the duration of anthesis
vary greatly from pIan t to plant within a species, and depend on environmental
and growing conditions. In spite of such variability,
P. picta
resulted to have a
significantly shorter flowering time than the other species, and to yield a lower
number of flowers. Under our experimental conditions
P. apennina
started
fIowering up to twenty days before
P. picta
and
P. vallarsae.
As regards phenology of vegetative characters, the basalleaves are produced
at a rate of 1 to 3 leaves per month. The rate varies during the year, depending
on the environmental conditions and on the development of the floral scape. A
remarkable difference was observed between
P. apennina
and
P. vallarsae,
the
leaves being produced at a faster rate in the former species than in the latter.
f)
Synecology and chorology -
The geographic and ecological distribution of
alI species was defined, with particular attention to the Italian distribution (fig.
3, 4), both on the basis of literature and of originaI observations.
AlI species grow mainly in mesic to submesic woods, often in clearings,
along footpaths, roads and streams.
P. picta
grows from the Ligurian Alps to the Northern and CentraI Apen-
nines (incl. Alpi Apuane), mainly on the Thyrrenic side. lts habitat are the
mesic mixed broad-Ieaved forests
(Laburno-Ostryon
UBALDI 1980) and the
montane beech forests
(Fagion),
up to the upper limit of the woods.
P. vallarsae
is widespread from the Adige Valley around Rovereto,
Eastwards to the area of Agordo (Province Belluno), mainly in beechwoods
(Fagion),
but it can be also found in spruce plantations in the same altitudinal
stage.
P. apennina
is widespread throughout the peninsula from Liguria to Cala-
bria, and is more thermophylic than
P. picta
and
P. vallarsae.
In the Northern
Apennines it grows in mixed broad-Ieaved formations
(Laburno-Ostryon),
while
lO
G. PUPPI
and
G. CRISTOFOLINI
Pulmonaria picta
5,
I
r
l'''
27l(
41ò.
V
54lc: \
6~ ~:
78/ ~ _
9l~~/.. te. _
102~ 104~ _ -J-J-
~ 111 ~ 1-1-
119
11 -I-
~
,2
248
1
AJr\I\
L
:c:+ti'( \
L
'-
Fig. 3 - Geographic distribution of
p, piela.
more southwards it can be found at higher elevations, in turkey oak and beech
woods
(Doronico-Fagion
Corbetta &Ubaldi in UBALDIet al., 1987).
DISCUSSION
AlI
macro-morphological characters are subject to a remarkable variability,
and can hardly help to draw interspecific boundaries. The phenotypic plasticity
of al:
game
Nevt
bet\),
shor
spec
.I~--~~----~~==~------------~
j
beech
lbility,
lsticity
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
• Pulmonaria apennlna
~ Pulmonaria vallarsae
1r-
-
V1
2
r-.
J
'\.
5r
Il,
~~
~->
15
1,5
f'-,
/
IV ~~~ ~
_\1"- ~~ ~
lr-.\
41 ~
V
J:'
~
,.:
·I~
••
\
9U~
r..
f
~
1021--- 104I-
••
• •
'-----'--
111 \
D.
119 • •
\
126~d
"1.,
• •
135I~~
.",
142
;'\
1'\
149
\'.
• •
I~~
15
8j1e.
166 ~
0.:
170
'\
N
179
l'-.
v
l'"\'
183
lJt"
,......."
192 "'\ ) 196
.
h ~
1-
213 }, J
205 20~
216 220
\
2231'-J
224 228 J '--
232 236
-Y
a
Ir""
24
1)
/'"
239
I~
~~ 244n
.)
248
A
~
/J-"~
L
2561~ ...
(
~
265
'"'
Ì"\
l
....
271I~r-- ~
275
.D
27
54
66
78
11
of all species causes an amount of variation within biological units (populations,
gamodemes etc.), which is comparable to the difference among different units.
Nevertheless, a set of characters, mainly microscopic, allows to discriminate
between species (tab. 5). In particular, the density and the measure of bristles,
short hairs, microglands and glandular hairs on the summer leaves are species-
specifico
Fig. 4 - Geographic distribution of p, apennina and P. vallanae.
I
r
12
G. PUPPI
and
G. CRISTOFOLINI
TABLE 5 - Synapsis of the main diagnostic characters of the rosette summer leaves. Absalute
numbers indicate appraximate density of indumentum (trichomes per mm'); fractional numbers
indicate the approximate of
setae
and ratia of
puberes
on the whale indumentum.
ba~
Total Se/ae Puberes Gland, Micro Surface Margin
hairs hairs glands Base Peliole Spo/s 1.
2.
P.saccharata
8-25
115
415
0,5
2.5 rough flat cuneate +/- wh~e,
broadly confluent
winged
2.'
3.
Paffinis
5-25
1/3
2J3
0.1 2
rough Hat abrufily narrowly whttish,
contracted winged nei:
conftuent
3."
P.picta
2-25
3/4
1/4
1-10
saft
+/-
ftat
+/- +/-
wh~e,
cuneate broadly conftuent
winged
4.
4.'"
P.vallarsae
30-70 1/10 9/10
1.5
0.5
soft, undulate abruptly broadly obscure,
sticky contracted winged not
confluent
or
unspctted
1.
P.apennina
25-70
115
415
0.5
1.5 saft, undulate abruptly narrowly whitish,
sticky contracted winged not
confluent
or
unspctted
Ph)
Ma
Ma
BM
223
The results of experimental crossing, and the patterns of variation, indicate
that some interspecific gene fIow is present, although it is not quantified.
However, reproductive isolation is certainly present to some extent, due to the
joint effect of ecological specialisation, of phenological differences, and of
reduced interfertility. Evidence of isolation is provided by the different pro-
tein patterns as welI as by the morphological characters, that are associated
with the geographic distribution and with ecological attitudes.
Therefore, it seems appropriate to give taxonomic recognition to a number
of elementary units, even if they do not represent biological species. Such a
praxis is in accordane e with the main trend in Pulmonaria taxonomy, that has
been followed from KERNER'S monograph onwards, and seems to be the only
practical way of affording the problem. On this basis, the Pulmonaria saccharata
aggr. is here resolved into five elementary species.
lO
fla
O.:
mI
ba
bit
al.
XI
Sp
[I
leaves.Absolute
:tional numbers
PeNale Spats
+/-
wMe,
broadly conftuent
Ninged
larrowly wMish,
winged nd:
conftuent
+/-
wh~e,
broadly conftuent
winged
broadly obscure,
winged no!
confluen!
or
unspctted
larrowly wh~ish,
winged no!
confluen!
or
unspctted
ion, indicate
t
quantified.
t, due to the
lCes, and of
ifferent pro-
:e associated
to a number
~cies. Such a
my,
that has
be
the only
riasaccharata
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
13
KEY TOTHE SPECIES
NOTE : ali characters of the indumentum should be observed on fully developed summer
basalleaves.
1.
Lamina of summer leaves covered with conspicuous white or bright green spots; surface not
sticky; hairs rare
«
25 per mm'); short hairs
(puberes)
less than 4/5 of the indumentum
2. Summer leaves surface soft; long hairs
(setae)
form 1/2 to 3/4 of the indumentum; glandular
hairs ca. 1 per mm' P. picta
2." Summer leaves surface rough; long hairs from 1/5 to 1/3 of the indumentum; leaf margin flat
3. Basalleaves lanceolate, cuneate at the base; bIade covered with large, confluent white spots;
puberes
<
0.2 mm long; petiole
±
broadly winged; corolla blue; tube glabrous below the
ring of hairs P. saccharata
3." BasaI leaves ovate, abrupt!y narrowed into petiole; bIade with scattered, whitish spots;
puberes 0.2-0.3 mm long; petiole narrowly winged; corolla pink to violet; tube hairy belo w
the ring of hairs P. affinis
1."
Lamina of summer leaves obscurely spotted or unspotted, abrupt!y contracted into petiole;
leaf margin undulate; surface sticky; hairs dense (25-70 per mm2); short hairs
(puberes)
from
4/5 to 9/10 of the indumentum
4. Glandular hairs ca. 1.5 per mm', microglands scattered (ca. 0.5 per mm'); petiole green,
usually broadly winged (wings ca. 2 mm wide) P. vallarsae
4.'" Glandular hairs rare (0.5-0.7 / mm'); microglands ca. 1.5 per mm'; petiole greenish-violet,
narrowly winged (wings
<
1 mm wide) P. apennina
1. Pulmonaria saccharata Miller, Gard.Dict., 8 Ed., n. 3 (1768).
SYN.:
Symphytum maculosum sive Pulmonaria maxima foliis quasi saccharo incrustatis
Plukenet,
Phytogr. T.227 (1691);
Pulmonaria Batavica maxima foliis longioribus maculis majoribus conspersis
Morison, Pl. Hist. Universo Oxon., 3: 444 (1699).
Pulmonaria grandiflora
DC., Cat. Hort.
Monsp.: 135 (1813).
Typus -
«Pulmonaria maxima foliis quasi saccharo incrustatis».
(Royal Society Collection,
BM). (designated by PUPPI
&
CRISTOFOLINI,1991).
IcONOGRAPHY- PWKENET (1691),
T.
227, fig. 4, reprinted by PUPPI
&
CRISTOFOLlNI(1991:
223)
DESCRIPTION - Lamina of summer basalleaves ovate-Ianceolate, 9-25 x 3-
lO
cm, covered by numerous, large, often confluent, whitish spots; margins
flat; base cuneate; hairs sparse (8 to 25 per mm
2),
mostly short
puberes
(0.08-
0.2 mm); rare microglands (ca. 2 per mm2)and glandular hairs (ca. 0.5 per
mm
2);
petiole more or less broadly winged; caulinar leaves spear-shaped, the
base haH embracing the stalk; corolla tube longer than the empalement; corolla
blue; tube glabrous below the ring of hairs. Flowers from ApriI to May.
CHROMOSOME NUMBER - not known.
GENERAL DISTRIBUTION - Belgium,The Netherlands? (see DUVIGNEAUD et
al., 1976; LAwALRÉE, 1949; LE]EUNE, 1813, 1824, 1828; MORISON, 1699)
SPECIMENSSEEN. Belgium: colI.
Dr. Lejeune
(PAVIA); Belgien, XII.1833,
Lejeune,
(K); Spa,
XII.1816,
Lejeune misit
(K); Environs de Veruieres?, XII.1816,
Lejeune misit
(K); environs de
Spaa, s.d.,
Lejeune
(K);
e flora Ha llensi ,
Weche, s.d.,
Lejeune
(BR); Belgien, s.d.,
Dr. Lejeune
14
G. PUPPI
and
G. CRISTOFOLINI SYST
(BR);
e fl. belg.,
s,d" M.
Martens,
(BR);
e fl. belg.,
s.d.,
M. Martens,
(BR); n. 306 P
saccharata
Miller, s.d"
Du Mortier?,
(BR); Herb. H. Brunner, s.d.,
Lawalree,
(BR); Siemers, en montant de
Corneilla au Vernet, s.d.,
scollo
(BR); Lebend 1861 von Spaa enthalten, IV.1865,
Schliekum
(BR)
The Netherlands:
(?)
Herb. Morison:
Pulmonaria Batavica maxima foliis 10ngi01ibus, maculis
majoribus conspersis
(OXF).
OBSERVATION- l
Flore de France, as "
centrale"
with
"feuille!
en pétiole étmit ailé, ..
courts et inégaux, .. " '
this group. Unfortuna
that may be considere
in any other pIace. Re
Typus - We des
Lupitana, 6.V.1951,}
IcONOGRAPHY-
I
2. Pulmonaria affinis Jordan, in Schultz, Arch. Fl. Fr. Allem., ser.I: 321
(1854).
SYN.:
Pulmonaria affinis
Jordan, Cat. Jard. Bot. Dijon :13 (1848),
nomen nudum; P confusa
Rouy, Fl. Fr. :296 (1908);
p saccharata sensu
GRENIERet GODRON,Fl. Fr., II: 527 (1850)
et sensu
COLMEIRO,Enum, Rev. Plant. Penino Hisp. IV: 121 (1888);
p,
ovalis
(Bast.) Boreau, Flore
Centre France: 458 (1857);
P. alpestris
Lamotte, Prodr.
FL
Plat. Centro Fr.: 535 (1881),
Typus - «"L'Arbresle" (Dept, Rhone),
Jordan,
1848» (G) (designated by BOLLIGER,1982).
IcONOGRAPIIY- KERNER(1878), Taf. VIII
&
XIII/V.
OBSERVATION-
A
race growing in the Swiss Jura, strict!y related to
P. affinis,
has rosette
leaves obscurely spotted. It has been described by BOLLIGER(1982) as an independent species
under the name
Pulmonaria helvetica.
We could not examine any specimen of this taxon. For the
diacritic characters of
P. helvetica
reference is made to BOLLIGER(cit.).
DESCRIPTION -
late, 10-25
X
5-~
gradually narrowir
whitish spots; ind
late; hairs 2 - 25 I
tum, microglands
mm
2);
petiole mor,
cm long; caulinar
florescence spread
tube hairy inside,
pink at first, later
CHROMOSOME
Sorgente del Tev
(SAUER, 1971). 2
Cimini (Viterbo)
Monte Rondinaio
DESCRIPTION - Lamina of summer basalleaves ovate to ovate-Ianceolate,
7-
20 x4-10 cm, abruptly narrowed into petiole; autumn leaves eIIiptic-lanceolate,
narrower than the summer leaves; lamina covered by numerous, large, often
confluent, whitish spots; indumentum rough; hairs 0.06 to 1.5 mm long, sparse
(ca. 5-25 per mm
2),
with a prevalence (70%) of short
puberes
(0.06 - 0.20 mm),
rare microglands (ca. 2per mm
2)
and glandular hairs (ca 0.1 per mm
2);
petioles
narrowly winged, as long as 1/2 to 3/4 the lamina; caulinar leaves lanceolate,
sessile, not embracing the stalk, spotted; inflorescence lax; calyx laciniae
exceeding the coroIIa tube; tube pubescent below the ring of hairs; corolla at
first reddish, then blue-violet to violet. Flowers in ApriI.
CHROMOSOME NUMBER - 2n
=
22: Isère, Basses Alpes, Tende (SAUER,
1975).
G ENERAL DISTF
SYN :
Pulmonaria saccharata
Auct. Fl. Ital.,
non
Miller;
Pulmonaria affinis
Auer.,
non
Jordan.
SPECIMENSSEEN
(BOLO); Taglieto (Va
di La Spezia, su calcar
Emilia-Romagna:
(BOLO); Lago Sante
11.X.1989, E.
Tibilet
(BOLO); Pievepelago
(Bologna), 460 m" 24
18.IX.1983, D.
Ubala
(BOLO); Lago Brasirr
Madonna dell' Acero
I
(Bologna), 960 m., 19
Toscana: Barberir
Tombaccio, Alpi Apu
Marcello Pistoiese,
2:
cuum inter loca S, Min
GENERAL DISTRIBUTION - France (only West of the Rhone), Spain [Pyren-
ees. (BoLLIGER, 1982)].
SPECIMENS SEEN - France: L'Arbresle, Lyon, s.d.,
s,collo
(NAP); Lyon
à
l'Arbresle,
28.IV .1870,
Jordan
(BR);
ibidem,
IV .1873,
JOldan
(BR); L'Arbresle pres Lyon, 1852, Jordan
(K);
Marcigny sur Loire, 25.IV et 5.VI.1896,
Dr. Gillot
(Herb Rouy, LY); Bois albi Tarn, III. 1994,
Sudre
(RO)
3. Pulmonaria picta Rouy, Fl. Fr., 2: 7 (1908).
6
P.
saccharata
en montant de
:65,
5chliekum
oribus, maculis
, ser.I:
321
~m; P. confusa
(1850)
et sensu
Boreau, Flore
1881).
LLIGER,1982).
!is,
has rosette
~ndent species
taxon. For the
nceolate, 7-
:-lanceolate,
large, often
long, sparse
- 0.20 mm),
TI
2);
petioles
; lanceolate,
lyx laciniae
5;
corolla at
de (SAUER,
lain [Pyren-
à
l'Arbresle,
2, Jordan (K);
arn,
III.1994,
, non
Jordan.
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
15
OBSERVATION- Rouy described this species almost incidentally in an
"Observation"
in his
Flore de France, as
"P. picta
Rouy
(P. saccharata
A. Kern,
non
Mill.) ,
plant de l'Italie surtout
centrale"
with
"feuilles abondamment maculées, ... feuilles l'ad. estivales lancéolées, ... subcontractées
en pétiole étroit ailé, ... page sup, munie de poils peu raides et d'une villosité composé de poils mous,
courts et inégaux, ... ".
This is out of doubt the first name validly published for the italian race of
this group. Unfortunately the protologue does not mention any collection or any other element
that may be considered for typification. As far as we know, Rouy did not return on this species
in any other pIace. Research for originaI material in FI, G, LY, P was unsuccessful.
Typus - We designate here the following neotype: Cerbaie (Toscana), nel bosco di Valle
Lupitana, 6.V.1951, A.
Contardo
(FI).
ICONOGRAPHY- KERNER(1878). Taf.
VII
&
XIII/III,
sub P. saccharata.
DESCRIPTION - Lamina of summer basalleaves lanceolate to ovate-Ianceo-
late, 10-25
X
5-9 cm, fIat or slightly undulate at the margin, cuneate and
graduaIIy narrowing into petiole, covered by numerous, large, often confIuent,
whitish spots; indumentum smooth, not sticky; autumn leaves elliptic-lanceo-
late; hairs 2 - 25 per mm
2,
short hairs forming 25% to 50% of the indumen-
tum, microglands scatteted (1 to lO per mm2), glandular hairs rare (ca. 1per
mm
2);
petiole more or less narrowly winged, often more or less violet, lO to 20
cm long; caulinar leaves lanceolate to oblong-spatulate, sessile, spotted; in-
fIorescence spread, not sticky; calyx laciniae exceeding the corolla tube; corolla
tube hairy inside, bu.t sometimes glabrescent below the ring of hairs; fIowers
pink at first, later blue-violet. Flowers from March to ApriI.
CHROMOSOME NUMBER - 2n =
22:
Lago Baccio (Modena)(!); Vallombrosa,
Sorgente del Tevere (MERXMULLER
&
GRAU, 1969); Val Varaita, Limone
(SAUER, 1971). 2n=26: Vergato (Bologna) (!), Riola (Bologna) (!), Monti
Cimini (Viterbo) (!). 2n =26 +b: Appennino Ligure (SAUER, 1975). 2n =28:
Monte Rondinaio (SAUER, 1975), Arezzo, Sarteano (Siena) (CAPINERI, 1982).
GENERAL DISTRIBUTION- France (SE), ltaly (Piemonte to Lazio). (fig. 3)
SPECIMENS SEEN -
Liguria:
Scogna (Sesta Godano), 600 m., 21.VI.1990,
S. Ballelli
(BOLO); Taglieto (Varese Ligure), 500 m., 21.VI.1990,
5,
Ballelli
(BOLO); a Romito, dintorni
di La Spezia, su calcare, 30.III.1984, D.
Marchetti
(SIENA).
Emilia-Romagna: Lago Baccio (Modena), 1560 m., 17.VI.1986, G.
Cristofolini
&
G.
Puppi
(BOLO); Lago Santo modenese, 7.VII.1937,
Lunardi
(FI); Merizzana (Modena), 1000 m.,
11.X.1989, E.
Tibiletti
(BOLO); Valle delle Tagliole (Modena) 1500 m., 1986,
M. Tomaselli
(BOLO); Pievepelago (Modena) 1290 m., 11.X.1989, E.
Tibiletti
(BOLO); Riola di Vergato
(Bologna), 460 m., 24.III.1985, G.
Cristofolini
(BOLO); Molino del Pallone (Bologna), 500 m.,
18.IX.1983, D.
Ubaldi,
(BOLO); Porretta (Bologna) margine di strada, 17 .IX.1989, E.
Tibiletti
(BOLO); Lago Brasimone (Bologna), 845 m., V.1977, D.
Ubaldi
(BOLO); Valle del Dardagna,
Madonna dell' Acero (Bologna), 850 m., 20.IX.1988, D.
Ubaldi
(BOLO); Passo della Raticosa
(Bologna), 960 m., 1990, G.
Cristofolini
(BOLO).
Toscana: Barberino del Mugello (Firenze), 450 m., 20 IV 1985,
A.L. Zanotti
(BOLO); Passo
Tombaccio, Alpi Apuane, 1300 m., 22.VI.1987,
M. Tomaselli
(BOLO); Maresca: comune di S.
Marcello Pistoiese, 22.VI.1941, R.
COI'I'adi
(FI); Vallombrosa,
in sylvaticis castanearum et quer-
cuum inter loca
5.
Miniato in Alpe et Lago dicta,
800-900m,
folia
16.IX.1909,
fIor.
21.IV .1910,
A.
16
G. PUPPI
and
G. CRISTOFOLINI
Fiori
(FI); Firenze in Gamberaia, 10.IV.1948,
A. Contardo
(FI); Montecuccoli presso Firenze,
5.IV.1871 (FI); M, Pejano presso Bigoli? 20.II.1864,
Beccari
(FI); lungo il canale di Arsina in
Vallebuia, 3.III.1861,
Beccm'Ì
(FI); Solco degli Streghi a Tempignano in val di Serchio,
8.IV.1861 e M. Pisano a Pozzuolo, 19.III.1861,
Beccm'Ì
(FI); Giogara (Casentino), 10-
20.V,1871,
Siemoni
(FI); Comune di Pontassieve, S. Brigida da cima di Monte Rotondo a
Messeri, m 772 fino a 700 ca., 27.V.1954,
C.
Gaito
(FI); Toscana alla Consuma (Firenze),
13VII.1942,
R.
Con'adi
(FI); Vallombrosa (Firenze), 29.VI.1939,
R.
Conadi
(FI); Monte Senario
(Firenze), 17.VI.1946,
R.
Con'adi
(FI); Florence, III.1866,
Chiarugi,
(FI); S. Romolo (Signa),
12.VII.1933,
R. Corti
e
R.
Con'adi
(FI); Lardarello e Serrazzano, nel Cerreto (Pisa), m.500 ca.,
29,VI.1953,
Chiarugi et al.
(FI); Grotta all'Onda-Casoli (M. Matanna), 6.IV.1938,
Chiarugi
(FI);
Cerbaie: parte inferiore di Valle Maggiore, 6.V.1951,
A.
Contardo
(FI); S.Casciano di Val di
Pesa, 20.VIII.1955, P
Paradossi
(FI); Monte Cetona (Siena), 5.VII.1935,
R.
Corti et
A.
Messeri
(FI);
leg, in nemoribus prope Carthusiam Florentinam,
19.IV.1857,
Calandrini
(RO); Consuma,
29.IV, 1902, G.
Sala
(RO); Querceto-cerreto tra S. Galgano e bivio per Casette, 24.X.1984,
M,
Mariotti et al.
(SIENA), tra Orgia e Brenna nei bordi boscosi dei campi, 19.V.1987, F
Semplici
(SIENA); Poggio Ermicciolo (M. Amiata) 1100 m. faggeta, 22.IV.1992, F
Selvi
(FI)
Marche: boschi prossimi ad Ascoli, s.d.,
Orsini?
(RO); Calmencio (Pesaro), 9.III.1974,
Ubaldi
(BOLO).
Lazio: Monti Cimini (Viterbo), 800 m., 30.VII.1986, G.
Cristofolini
(BOLO); Soriano nel
Cimino (Lazio), VIII. 1957,
Anzalone
(RO); Colli Albani (Roma) 10.IV.1881,
Baccarini
(SIENA).
4, Pulmonaria vallarsae A. Kerner, Monogr. Pulm.: 33 (1878).
Typus -
«Ti1'Olia australis, Vallarsa, ca, 1870»
(Herb. Kerner, WU), designated here.
IcONOGRAPHY- KERNER(1878), Taf.
XI
&
XIII/VI.
DESCRIPTION- Lamina of summer basalleaves 8-27
X
5-12 cm, ovate to
ovate-Ianceolate, more or less undulate at the margin, rounded at the base,
more or less abruptIy narrowed into the petiole; autumn leaves elliptic-Ianceo-
late; narrowing gradually into petiole, undulate; colour usualIy uniformly green,
bright-green spots sometimes present, but never whitish nor confluent; surface
soft, more or less sticky; hairs dense (30 to 70 per mm
2),
short (average length
0.15 mm); glandular hairs ca. 1.5 per mm2,microglands ca. 0.5 per mm2;
petiole broadly winged, green, as long as 1/2 to 3/4 the bIade: caulinar leaves
ovate to lanceolate, sessile, sometimes bright-greenish spotted; inflorescence
dense; calyx laciniae exceeding the corolla tube; corolla tube hairy or glabres-
cent inside; flowers pink at first, then blue-violet. Flowers from March to
April.
CHROMOSOMENUMBER- 2n
=
22: Pian delle Fugazze (Tren to)
(!),
Vallarsa
(MERXMULLER
&
GRAU, 1969).
GENERALDISTRIBUTION- Italy: Trentino-Alto Adige, Veneto (fig. 4).
SPECIMENS SEEN - Trentino: Pian delle Fugazze (Trento), 18.V.1985, G,
Cristofolini
(BOLO); Vallarsa (Trento), 18.V.1985, G.
Cristofolini
(BOLO).
Alto Adige: Fie', Valle di Tires (Bolzano), Faggeta, VII.1990, D,
Ubaldi
(BOLO).
li presso Firenze,
naIe di Arsina in
val di Serchio,
(Casentino), 10-
[onte Rotondo a
nsuma (Firenze),
:);Monte Senario
Romolo (Signa),
Pisa), m.500 ca.,
38, Chiarugi
(FI);
Isciano di Val di
:ortiet A. Messeri
(RO); Consuma,
ò, 24.X.1984,
M,
1987, F
Semplici
)i
(FI).
aro), 9.III.1974,
LO); Soriano nel
!ccarini
(SIENA).
78).
ated here.
!
cm, ovate to
l
at the base,
:lliptic-lanceo-
iformly green,
]uent; surface
lverage length
J.5
per mm
2;
aulinar leaves
infIorescence
iry or glabres-
Dm
March to
)) (!),
Vallarsa
l
(fig. 4).
, G.
Cristofolini
OLO).
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA
17
Veneto: Cogul di Val Agordina (Belluno), IV.1985,
M. Tomaselli
(BOLO); Losen (Belluno),
730 m., 1988,
Lasen
(BOLO); Bellunese, s.d.,
Minio
(FI); Dopo S. Dionisi (Verona), l.IV.1982,
Bianchini
(VER).
5. Pulmonaria apennina
Cristof. et Puppi, sp. nov.
SYN.:
Pulmonaria mollis
Ten., Fl. Nap. Syll.: 84 (1830) non Wulfen ex Hornemann, Hort.
Reg. Bot. Hafn., 1: 179 (1813);
P. vallarsae
Auct. Fl. Ital.
pro parte.
Typus - «Parco Talon, Casalecchio di Reno, 26.IV .1996,
Giovanna Puppi»
(Holotype in FI,
isotype in BOLO).
ICONOGRAPHIA- fig. 5.
DIAGNOSIS- E
Pulmonariae vallarsae
Kernerii affinitate, a qua differt indumento folio rum
basalium, ex densis (circa 1.5 per mm') glandulis minutis, et radioribus (circa 0,5 per mm')
glandis stipitatis composito; petiolo subtiliori, strictis (ca. 1 mm) alis praedito, violaceo colore.
Habitat in nemoribus Italiae peninsularis. Floret Martio et Aprile. Sporophyti chromosomata 22.
Nomen
è
montibus Apenninis ductum, ubi pIanta crescit.
DESCRIPTION- Lamina of summer basalleaves ovate to ovate-Ianceolate,
10-25x5-10 cm, often undulate at the margins, abruptly contracted into petio-
le; autumn basaI leaves elliptic-lanceolate; bIade unspotted, or spread with
bright-green, circular, usually not confluent spots; surface soft and sticky,
indumentum very dense (25 to 70 hairs per mm2); short
puberes
prevailing (up
to 80% of hairs), long
setae
rare, glandular hairs ca. 0.5 per mm2,microglands
ca. 1.5 per mm
2;
petiole narrowly winged, from a haH to as long as the bIade;
caulinar leaves ovate to lanceolate, sessile, sometimes spotted; inflorescences
compact at first, then spreading; calyx laciniae exceeding the corolla tube;
corolla usually hairy inside below the ring of hairs, pink at first, then blue-
violet. Flowers from March to ApriI.
CHROMOSOMENUMBER- 2n
=
22: Bologna (!), Corniolo (Forlì) (!), Palena
(Chieti)
(!),
Senarica (Teramo) (!); Sirente, Foresta Umbra, Pollino (MERXMUL-
LER
&
GRAU,1969); M. Vulturino (Potenza), Roma, Bagni di Lucca (CAPINERI,
1986). 2n
=
22 +2b: M. Procinto (Alpi Apuane) (CAPINERI,1986).
GENERALDISTRIBUTION- Italy (from Liguria to Calabria) (fig. 4).
SPECIMENSSEEN- Liguria: S. Lazzaro Reale (Imperia), 8.VIII.1937,
Corradi
(FI); strada di
Viozene-Ponti di Nava, 8.IX.1948,
Corradi
(FI); Val di Vara su argille a palombini, 125 m., s.d.,
D,
Marchetti
(SIENA); a Pignone tra Monterosso al Mare e la S.S. Aurelia, su calcare, 200 m.,
3.IV.1982, D.
Marchetti
(SIENA).
Piemonte: Priero (Cuneo), 500 m., 22.V.1986, G.
Cristofolini
(BOLO); Montezemolo
(Cuneo), 700 m., 22.V.1986, G.
Cristofolini
(BOLO); Val Bormida (Cuneo), 300 m., 22.V.1986,
G.
Cristofolini
(BOLO); Cuneo, Certosa di Pesio, 4.III.1946, R.
Berenti
(FI); Roccavione,
Cuneo, s.d.,
Biadego
(VER).
Emilia-Romagna: Valle Tresinaro (Reggio Emilia), 300 m., 1984, D.
Ubaldi
(BOLO); S.
Lazzaro di Savena (Bologna), 60 m., 1990, G.
Bugamelli
(BOLO); Parco Talon di Casalecchio di
Reno (Bologna), 90 m., 9.IX.1985, G.
Puppi
(BOLO); Monte Adone (Bologna), 400 m.,
IV.1985,
M. Tomaselli
(BOLO); Rio Croara, Vergato (Bologna), 300 m., V.1987,
M. Tomaselli
18
G. PUPPI
and
G. CRISTOFOLINI
(BOLO
414
m.,
m,
III
Lupo, l
(FIl; Se
Tm
(Lucca)
2.V.18
Marche
M. FaI
COrl'adl
Beccari
Boscoh
S. Stef
19.IV.
Ma
del
MI
(BOLe
presso
Tibilett
Piceno
Un
Padule
La:
Oscura
5.III.,
culI, e
Albane
- Valle
M
m.,
V
2.VU
Pellegr.
Castel
Cesati
M(
Bianch
Pu
Ca
Cervat
leclis,
presso
Bl
Poten:
Madd,
VU9'
C~
(RO);
ACKNC
Tl
Curate
WU,
Cartoe
Fig. 5 -
Pulmonaria apennina
Cristof. et Puppi.
SYSTEMATICS OF THE COMPLEX PULMONARIA SACCHARATA 19
(BOLO); Corniolo (Forlì), 600 m., X1.1986, N.
Agostini
(BOLO); S.Benedetto in Alpe (Forlì),
414 m., III 1987, N.
Agostini
(BOLO); Modigliana (Forlì), 1988 (BOLO); Strigara (Cesena), 500
m., III.1987, N,
Agostini
(BOLO); Valle Baganza (Parma), 6.IV.1920,
Minio
(FI); Passo del
Lupo, 1500 m., 25.V.1955,
Chiarugi
(FI); Colli dell'Osservanza a Bologna, 5.IV.1890,
A. Fiori
(FI); Sestola (App. modenese), 1020 m., VIII. 1962,
s. collo
(RO).
Toscana: Alpi Apuane (Massa) 125 m., 20.IV.1985,
M. TomaseUi
(BOLO); Le Pizzorne
(Lucca), 900 m., 24.III.1985,
M. Tomaselli
(BOLO); Fl. Italica, Etruria,
Florenz in umbrosis,
2,V.1886, E.
Levier
(RO); nei pressi di Bibola sotto Aulla (Massa) Alpi Apuane, 16.IV.1980, D.
Marchetti
(SIENA); presso Redicesi di Massa, Alpi Apuane, 8.IV.1978, D.
Marchetti
(SIENA);
M. Falterona, 700-900 m., S. Godenzio, 1952,
Forasassi
(FI); La Burraia, Campigna, 1952,
Con'adi
(FI); dalla Calla a Poggio Scali per la Giogaia, 1912,
Baccarini
(FI); Montagioli, 1863,
Beccal'i;
Viale dei Colli (Firenze), 1945,
Con'adi
(FI); Cafaggiolo, 7.VII.1940,
Corradi
(FI);
Boscolungo, 1883,
Levier
(FI); Versilia, Stazzema, 1923,
Chiarugi
(FI); Passo di Viamaggio-Pieve
S. Stefano (Arezzo), 19.IV.1976, D.
Ubaldi
(BOLO); Pieve S. Stefano - La Verna (Arezzo),
19.IV.1976, D,
Ubaldi
(BOLO).
Marche: M. Carpegna (Pesaro), 900 m., 31.V.1983, D.
Ubaldi
(BOLO); Monastero - Valle
del Mutino (Pesaro), 1972,
Ubaldi
(BOLO); Sasso Simone (Pesaro), l.V.1976, D.
Ubaldi
(BOLO); S. Michele tra Matelica e Fabriano, 300 m., 26.X.1990, E.
Tibiletti
(BOLO); S. Angelo
presso Matelica, 550 m., 26.X.1990, E.
Tibiletti
(BOLO); Camerino, 350 m., 26.X.1990, E.
Tibiletti
(BOLO); Visso (Macerata) 520 m., 28.X.1990, E.
TibiZetti
(BOLO); Amandola (Ascoli
Piceno) 590 m., 28.X.1990, E.
Tibiletti
(BOLO).
Umbria: Scheggia (presso Gubbio), 680 m., 27.X.1990, E.
Tibiletti
(BOLO); Gubbio -
Padule, 510 m., 27.X.1990, E.
Tibiletti
(BOLO).
Lazio: Monti Albani (Roma), 13. V1.1881,
Pedicino
(VER); nel castagneto presso Valle
Oscura nei colli Albani, G.
Lusina
(RO);
plantae in agro romano lectae,
S. Cesareo nel bosco,
5.III.,
A. Cacciato
(RO);
culto e radic. in mont, prope Albano lectis,
21.11.1920,
C.
Lacaita
(RO);
cult. e radic. in silvis supra Nemi lectis,
21.II.1920,
C.
Lacaita
(RO); Colli Albani tra Capuccini e
Albano,
Biadego
(RO); Riano (Roma), 1980,
Caprolati
(RO); Monti Simbruini (Lazio) zona Trevi
- Valle Pietra, 24.V.1985,
Anzalone
(RO); M. Terminillo, VI.1983,
Anzalone
(RO).
Abruzzo: Senarica (Teramo), 700 m., IX.1986,
AL Zanotti
(BOLO); Palena, Majella, 1000
m., VII.1987, G.
Cristofolini et al.
(BOLO); Parco Nazionale d'Abruzzo, verso Bisegna,
2.VI.1956,
Anzalone
(RO); Tagliacozzo (Aquila), 2.IV.1961, S.
colf.
(RO); Monte Sirente, s.d.,
Pellegrini
(RO); Castel di Sangro,
ex herb.
Gussoniano, s.d.,
Cesati
(RO); Monte di Mezzo prope
Castel di Sangro, 23.V.1837,
Gussone
(NAP); Italien, Castel di Sangro ex herb. Gussonii, s.d.,
Cesati
(Herb. Kerner, WU).
Molise: Flora del Matese, Faggeta alle pendici del M. Miletto, 1459 m., l.VII.1968,
Bianchini
(VER); tra S. Massimo e Campitello, 1200 m., 26.V.1971,
Bianchini
(VER).
Puglia: Foresta Umbra, Gargano, 800 m., 4.V.1982,
Bianchini
(VER).
Campania: Monte Gelbison (Salerno), 1600 m., VI1.1987, D.
Ubaldi
(BOLO); Monte
Cervati (Salerno), 1000 m., VI1.1987, D.
Ubaldi
(BOLO);
cult ... e radic. in montibus supra Amalfi
lectis,
800 m., 21.II.1920,
Lacaita
(RO); selve di .... sopra Maddaloni? e bosco di Querciacupa
presso Valle, 10.IV.1838,
Gussone
(NAP).
Basilicata: Potenza,
in siZvaAria Silvana,
900-1200 m., 26.IV.1936,
Gavioli
(RO); Lucanie,
Potenza, bois de Quercus cerris, sol argileux, altitude 1000 m., 24.IV.1933, O.
Gavioli
(RO); La
Maddalena, bosco di cerro-faggio 1300 m., VIII. 1977, D.
Ubaldi
(BOLO); Villa d'Agri,
VI.1991, G.
Puppi
(BOLO).
Calabria: in Calabria, s.d.,
B. Longo
(RO); in monto di Pecoraro (Calabria), 1877,
Arcangeli
(RO); monti da Giffone a Serra S. Bruno, con Lacaita, 1907,
PasquaZe
(NAP).
ACKNOWLEDGEMENTS
The authors gratefully acknowledge the invaluable help received by the Directors and
Curators of the Herbaria: BM, BR,
FI,
K, LY, NAP, OXF, PAVIA, RO, SIENA, TSB, VER,
WU, ZT. The type of
P. apennina
was skilfully drawn by Cinzia Gasperini (Lucrezia di
Cartoceto, Pesaro).
20
G. PUPPI
and
G. CRISTOFOLINI
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H.
&
SAUERW., 1972. - Pulmonaria L. In: TUTIN T,G. et a!' (eds.), Flom Europaea: 3: 100-
102. Cambridge University Press, Cambridge.
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CRISTOFOLINIG., 1991. - Sul significato del binomio Pulmonaria saccharata Miller. Webbia 45
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Summary
The complex Pulmonaria saccharata-P. vallarsae has been studied by means of morphological,
biometric and biosystematic analysis of dried and living specimens, in natural environments and
under cultivation. Hydroponic cultures have been used to distinguish the effects of environmen-
tal conditions and of phenological phases in determining phenotypic plasticity. An electrophore-
tic study of the seed proteins has been done. Inter- and infra-specific crossability has been
experimentally tested. Chromosome counts on populations of ali italian species have been made.
The following species have been recognised: P. saccharata Miller, confined to Belgium and
perhaps the Netherlands, to be excluded from Italy; P. affinis Jordan, which is strictly related to
P. saccharata, and is distributed throughout France and in Eastern Spain; P. picta Rouy
(=
P.
saccharata Auct. Fl. Ital.), distributed from SE France to CentraI Apennines, mainly on the
Tyrrenic side of the Italian peninsula; P. vallarsae Kerner, endemie to the italian Pre-Alps; P.
apennina Cristof. et Puppi sp. nov.
(=
p,
vallarsae Auct. Fl. Ital. pro parte), endemic to Italy,
widespread from Liguria to Calabria, mainly on the Adriatic side of the peninsula.
... The latter taxon is currently treated as a subspecies of the allopatric P. vallarsae because of striking morphological similarity (Cecchi & Selvi, 2015;Bartolucci & al., 2018). All these taxa are putatively different in features of summer basal leaves, such as shape, maculation, and hair pattern (Kerner, 1878;Bolliger, 1982;Puppi & Cristofolini, 1996). In addition, they also show differences in chromosome number: 2n = 28 in P. hirta, although 2n = 22 and 26 cytotypes have also been reported (Puppi & Cristofolini, 1996); 2n = 22 in P. apennina, although 2n = 26 has also been reported (Astuti & al., 2019); 2n = 22 in P. vallarsae (Puppi & Cristofolini, 1996). ...
... All these taxa are putatively different in features of summer basal leaves, such as shape, maculation, and hair pattern (Kerner, 1878;Bolliger, 1982;Puppi & Cristofolini, 1996). In addition, they also show differences in chromosome number: 2n = 28 in P. hirta, although 2n = 22 and 26 cytotypes have also been reported (Puppi & Cristofolini, 1996); 2n = 22 in P. apennina, although 2n = 26 has also been reported (Astuti & al., 2019); 2n = 22 in P. vallarsae (Puppi & Cristofolini, 1996). Pulmonaria apennina overlaps with P. hirta in the northern portion of its range (Tuscan-Aemilian Apennine), where a high morphological convergence and intermediate chromosome numbers have been observed (Puppi & Cristofolini, 1996;Vosa & Pistolesi, 2004). ...
... All these taxa are putatively different in features of summer basal leaves, such as shape, maculation, and hair pattern (Kerner, 1878;Bolliger, 1982;Puppi & Cristofolini, 1996). In addition, they also show differences in chromosome number: 2n = 28 in P. hirta, although 2n = 22 and 26 cytotypes have also been reported (Puppi & Cristofolini, 1996); 2n = 22 in P. apennina, although 2n = 26 has also been reported (Astuti & al., 2019); 2n = 22 in P. vallarsae (Puppi & Cristofolini, 1996). Pulmonaria apennina overlaps with P. hirta in the northern portion of its range (Tuscan-Aemilian Apennine), where a high morphological convergence and intermediate chromosome numbers have been observed (Puppi & Cristofolini, 1996;Vosa & Pistolesi, 2004). ...
Article
Full-text available
Hybridization and introgression have a significant impact on the taxonomically controversial genus Pulmonaria. Within this genus, the P. hirta complex shows puzzling systematic relationships among P. hirta s.str. (2n = [22, 26]28), P. apennina (2n = 22[26]), and P. vallarsae (2n = 22), showing range overlaps and mixed phenotypes in Southern Europe. We carried out morphometric analyses of basal leaves and flower features along with AFLP characterization of 236 plants belonging to 11 populations within the complex and 1 population of P. officinalis. We also implemented an already available phylogeny with sequences from our target populations and characterized their karyotype. For all the populations within the complex, we found molecular evidence of a hybrid origin involving species belonging to different clades (angustifolia and officinalis clades). However, there is a certain morphological differentiation between some populations (“hirtoid” morph) and others (“vallarsoid” morph), albeit single individuals or entire populations show intermediate features. According to our results, hybridization and/or backcrossing/introgression have occurred, and gene flow is currently taking place among these "taxa". Following the hybridization event(s), we can elaborate three possible evolutionary scenarios: 1) one hybrid "vallarsoid" (2n = 22) species spread across Italian peninsula and from this originated the "hirtoid" morph (2n = 28) through dysploidy; 2) two geographically distinct hybridization events produced both "vallarsoid" and "hirtoid" morphs; 3) one "hirtoid" alloploid hybrid species originated and backcrossed with P. officinalis generating "vallarsoid" plants. Under scenarios 1 and 2, the different morphs met again in C Italy, with massive current gene flow. Under scenario 3, “vallarsoid” plants spread across Italian peninsula, but further backcrossed with "hirtoid" plants in C Italy, leaving pure lineages of "vallarsoid" plants only in the extreme north and south of their range. This latter scenario is supported by populations with 2n = 22, 26 chromosomes, having karyotype asymmetry indices intermediate between those of 2n = 16 and 2n = 28 cytotypes. Irrespective of the evolutionary dynamics, today a single lineage showing three cytotypes occurs throughout the Italian peninsula, supporting the circumscription of a single polymorphic species, namely P. hirta.
... More recently, the name P. picta has been rejected in favour of P. hirta L., the earlier name holding priority (Conti et al. 2007). Puppi & Cristofolini (1996) provided evidence that Alpine and Apennine populations attributed to P. vallarsae are distinct enough to be separated at species level, so that the peninsular populations were described as Pulmonaria apennina Cristof. & Puppi, sharing with P. vallarsae the same chromosome number 2n = 22 (2n = 22 + 2B for specimens from Alpi Apuane: Merxmüller & Grau 1969;Capineri 1986), although 2n = 26 has now been reported for a population of P. apennina On the other hand, most chromosome counts for P. hirta gave 2n = 28, but some reports had 2n = 22 as well as intermediate numbers such as 2n = 26 and 27, thus partially overlapping P. apennina and P. vallarsae (Merxmüller & Grau 1969;Sauer 1975;Capineri 1986;Puppi & Cristofolini 1996;Astuti et al., 2019). ...
... Puppi & Cristofolini (1996) provided evidence that Alpine and Apennine populations attributed to P. vallarsae are distinct enough to be separated at species level, so that the peninsular populations were described as Pulmonaria apennina Cristof. & Puppi, sharing with P. vallarsae the same chromosome number 2n = 22 (2n = 22 + 2B for specimens from Alpi Apuane: Merxmüller & Grau 1969;Capineri 1986), although 2n = 26 has now been reported for a population of P. apennina On the other hand, most chromosome counts for P. hirta gave 2n = 28, but some reports had 2n = 22 as well as intermediate numbers such as 2n = 26 and 27, thus partially overlapping P. apennina and P. vallarsae (Merxmüller & Grau 1969;Sauer 1975;Capineri 1986;Puppi & Cristofolini 1996;Astuti et al., 2019). In a survey of some two hundred populations of Pulmonaria in northern Apennines, Vosa & Pistolesi (2004) found three major karyotypes: 2n = 16 (P. ...
... range, dominating the high course of river Reno, has been an important religious area since Etruscan times. Most of these plants show elongated summer leaves with conspicuous ("mortar-like") to confluent spots, and a strigose indumentum formed by bristles (setae) and a few short hairs (puberes) (Puppi & Cristofolini 1996). All P. hirta plants sampled had 2n = 28 chromosomes. ...
Article
Full-text available
Populations of Pulmonaria on Apennine highlands in the province of Bologna were investigated. Except for a few isolated sites with P. officinalis, most populations are referable to P. apennina on the basis of leaf shape, indumentum and chromosome number (2n = 22). Pulmonaria apennina occurs in two morphs: i) a green morph with few or no leaf spots, common in mountain sites and similar to its Alpine vicariant, P. vallarsae; ii) a variegated morph with diffuse, clear leaf spots occurring in cool locations at low altitude, often not distant from cities. Hilly areas along the Reno valley, however, are colonized by populations of P. hirta with 2n = 28 chromosomes, characterized by lanceolate leaf shape and extensive maculation. Certain similarities between P. hirta and the variegated morph of P. apennina have long caused confusion and misidentifications, but the two species in the area of investigation are distinguishable on the basis of leaf morphology and indumentum.
... Pulmonaria vallarsae subsp. vallarsae is endemic to Trentino-Alto Adige and Veneto (Cecchi and Selvi 2015), and it was described by Kerner (1878) on plants occurring in Vallarsa, the valley of the river Leno situated southeast of Rovereto, which is the locus classicus for this species (Puppi and Cristofolini 1996). Pian delle Fugazze, the northern summit of the valley, is among the localities cited in the protologue. ...
... These plants display intermediate morphological features between the typical P. hirta and the typical P. vallarsae subsp. apennina (Puppi and Cristofolini 1996;Cecchi and Selvi 2015), although showing a closer resemblance to the former species, whose range spreads from SE France to C Italy (Cecchi and Selvi 2015). Four out of six samples in this population were found to have 2n = 28 chromosomes, which is typical for P. hirta Pupillo et al. 2019), whereas the remaining two individuals were found to have 2n = 22 chromosomes. ...
Article
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In this contribution, new chromosome data obtained on material collected in Italy are presented. It includes counts from six populations of three taxa within the genus Pulmonaria , two of which are endemic to Italy (P. vallarsae subsp. apennina and P. vallarsae subsp. vallarsae); the other is the widespread European P. officinalis . In addition, two counts from Potentilla detommasii and Stachys thirkei , two eastern Mediterranean species, are also reported.
... Observations. Puppi and Cristofolini (1996) collected the holotype of the name P. apennina from Parco Talon (Emilia-Romagna). All the 18 plants sampled from this population show the typical chromosome number of P. apennina, i.e. 2n = 22, without Method. ...
... We found 2n = 28 chromosomes, the chromosome number typical of this species (Cecchi and Selvi 2015), and no variation among the 16 individuals sampled in this topotypical popula-tion. A specimen coming from Valle Lupitana (Toscana) was selected by Puppi and Cristofolini (1996) as the neotype for the name Pulmonaria picta Rouy, a heterotypic syonym of P. hirta. Also for the 18 individuals sampled in this population, we found 2n = 28 chromosomes, in agreement with other counts from the same geographical area (Vosa and Pistolesi 2004). ...
Article
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In this contribution, new chromosome data obtained on material collected in Italy are presented. It includes a total of 105 chromosome counts for three populations of Pulmonaria vallarsae A.Kern. subsp. apennina (Cristof. & Puppi) Cecchi & Selvi and for three populations of P. hirta L.
... The collected plants were identified using standard floras (Fiori 1923(Fiori -1929Zangheri 1976;Tutin et al. 1964Tutin et al. -1980Tutin et al. , 1993Pignatti 1982;Pignatti et al. 2017aPignatti et al. , 2017bPignatti et al. , 2018Pignatti et al. , 2019Castroviejo et al. 1986Castroviejo et al. -2019Tison and de Foucault 2014). To identify samples belonging to critical groups, we used specific taxonomic studies (Grau 1970;Tornadore and Garbari 1979;Chrtekjun 1992;Puppi and Cristofolini 1996;Del Carratore et al. 1998;Garbari et al. 2003Garbari et al. , 2007Peruzzi et al. 2007;Brullo et al. 2009;Arnelas and Devesa 2011;Ardenghi et al. 2014Ardenghi et al. , 2015aArdenghi et al. , 2015bArrigoni 2014;Roma-Marzio et al. 2015Conti et al. 2019;Bartolucci et al. 2020). Some herbarium specimens belonging to critical genera were sent to specialists for revision: Alchemilla L. The exsiccata of collected plants are stored in the 'Sandro Ballelli' Herbarium, part of the Herbarium Universitatis Camerinensis (CAME). ...
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This study aims to increase floristic knowledge of Marche by means of a survey in the Montagna di Torricchio State Nature Reserve (central Italy). The Reserve, located in the central Apennines, covers about 3.2 km ² at altitudes ranging from 820 to 1,491 m a.s.l. It has been owned and managed as a strict reserve by the University of Camerino since 1970: all the anthropic activities ceased about 50 years ago, except for a minimal area where mowing and cattle grazing are still allowed. The floristic list consists of 789 specific and subspecific taxa belonging to 81 families and 352 genera. Two species are new for Italy ( Taraxacum calocarpum and T. pulchrifolium ) and 14 for Marche regional flora. Compared to previous floristic studies, we found 127 more taxa but we showed a certain stability in the life-form spectrum, suggesting limited effects of dynamic processes related to climate and land-use changes. The negligible number of alien species (11) is probably related to the limitations to anthropic activities in the Reserve. The occurrence of taxa never recorded for Italy and Marche highlights the floristic value of the Reserve for species conservation in the central Apennines.
... garganicum e Rumex scutatus. ANZALONE, 1983ANZALONE, , 1984ANZALONE, , 1991ANZALONE, BAZZICHELLI, 1960;ANZALONE, CORAZZI, 1998, 1998aANZALONE, LATTANZI, 1988;ANZALONE, VERI, 1975;BIGAZZI, RAFFAELLI, 2000;BOTTEGA, GARBARI, 2003;COLASANTE, ALTAMURA, 1988;CORAZZI, 2003;CORAZZI et al., 2003;CORAZZI, TILIA, 1998;DE ANGELIS, SCACCHI, 1991;DE PERSIS, 1987;DE PERSIIS, DE PERSIIS, 1986;DE PISI et al., 2005;DI PIETRO et al., 1997;FALQUI, 1899;FORTINI, , 1996aFRONDONI, IBERITE, 1998;FURNARI, 1970;GARBARI, 1984;GARBARI et al., 2003;GIGLI, 1996;GRANDE, 1920GRANDE, , 1922GUARRERA, 1996;IBERITE, ANZALONE, 2001;LATTANZI et al., 1983LATTANZI et al., , 2000LATTANZI et al., , 2003LATTANZI, TILIA, 1995, 2000, 2004LEPORATTI et al., 1994;LUCCHESE, 1986LUCCHESE, , 1996LUCCHESE, , 1996aLUCCHESE, , 1996bMARTINOVSKY, MORALDO, 1980;MONTELUCCI, 1956MONTELUCCI, , 1964MONTELUCCI, , 1971MORALDO, 1986;NARDI, 1984;PALOMBI, 1986;PIGNOTTI, 2003;PUPPI, CRISTOFOLINI, 1996;RASETTI, 1980;RAFFAELLI, BALDOIN, 1997;RAFFAELLI, RICCERI, 1987;STADLMANN, 1906;STEFFAN, STEFFAN, 1983TRAVAGLINI et al., 1999;VERI, 1988;VERI, BRUNO, 1978. La flora informatizzata, contenuta nel CD allegato, consente la determinazione delle 1500 entità censite. ...
Article
In this paper the computerised analytical Flora of the Special Protected Area Simbruini-Ernici mountains is presented. The Flora contains 1500 entities identified through floristic field analysis carried out between the year 2000 and the year 2005. The Flora allows the classification of a specimen trough two different approaches: the dichotomous keys of the Flora d'Italia, a research module based on species characters such as sexual and vegetative structures, habitus, habitat and geographical distribution. Both classification processes end with a form containing the descriptive information of each species and one or more photos taken in the field. There are a total of 4.000 photopraphs in the archive. Moreover a complete list of 1812 entities was obtained adding data from scientific literature and specimens from Rome's Herbarium (RO). The study area extends for about 53.000 ha. The lithology is mainly made of limestone formations. The bioclimate belongs to the Mediterranean temperate region. The vegetation consists mainly of beech woodlands and montane pastures. 48 among the 1500 entities are escaped from cultivation or are adventitious naturalized or introduced. The spontaneous 1460 entities are distributed in 1431 species, 568 genera and 112 families. Some species are interesting because very rare (Equisetum hyemale, Minuartia capillacea, Silene vallesia subsp. graminea, Ranunculus magellensis, Descurainia sophia, Aubrieta columnae subsp. columnae, Parnassia palustris, Potentilla supina, Geranium subcaulescens, Cerinthe minor subsp. auriculata, Scutellaria alpina, Scutellaria altissima, Euphrasia illyrica, Pinguicula vulgaris, Campanula foliosa, Campanula latifolia, Leontopodium alpinum subsp. nivale, Doronicum orientale, Cirsium oleraceum, Veratrum album subsp. lobelianum, Allium schoenoprasum subsp. sibiricum, Iris marsica, Carex umbrosa subsp. umbrosa, Orchis militaris, Orchis spitzelii, ecc.) and 3 species have been confirmed (Draba aizoides, Myosotis sylvatica s.l., Alopecurus pratensis). The biological spectrum underlines a predominance of hemicryptophytes (40,8%), in accordance with the geological and climatical features of the area. The chorological spectrum shows a prevalence of Eurasiatic (25,5%) element, followed by Mediterranean element (11,9%), in accordance with the climate.
... Gli exsiccata sono conservati presso gli erbari degli autori. Nel redigere l'elenco ci siamo attenuti alla nomenclatura della Flora d'Italia (PIGNATTI, 1982), salvo alcuni casi in cui abbiamo utilizzato il Prodromo della Flora Romana (ANZALONE, 1994(ANZALONE, , 1996 o recenti revisioni tassonomiche (BARBO, CELA RANZONI, 1998;DIAZ LIFANTE, VALDÈS, 1996;PUPPI, CRISTOFOLINI, 1996;SELVI BIGAZZI, 1998). Nell'elenco, oltre al binomio latino, sono indicati la forma biologica e il tipo corologico desunti da PIGNATTI (l.c.). ...
Article
The paper describes the floristic, structural and chorological features of the Veio Park, a natural protected area of about 15.000 ha situated in the north sector of Rome around the old Etruscan city of Veio. 787 taxa, mainly Mediterranean and Euroasiatic annuals and hemicriptophytes species, which represent the integration of climatic, geopedological and antropic factors. The importance of this area within the ecological network of the metropolitan area of Rome is also discussed.
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Some records of particular fitogeographical interest were found in Aspromonte mountains (S Calabria). New distribution data are presented about Osmunda regalis, Tilia platyphyllos subsp. pseudorubra, Woodwardia radicans, Quercus petraea subsp. austrothyrrenica, Pulmonaria apennina, Ptilostemon gnaphalioides, Tuberaria lignosa, Ulmus glabra, Polygala angelisii, Pteris vittata, Corydalis solida subsp. densiflora, Tricholaena teneriffae. Besides, 6 species are new about Aspromonte area (Consolida ajacis, Pteris cretica, Carpinus betulus, Saponaria calabrica, Delphinium staphisagria, Myriophyllum alterniflorum).
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Vengono qui presentati i risultati di una indagine storica ed erbariologica volta a chiarire il significato del binomio P. saccharata Miller. I dati raccolti hanno permesso di designare un lectotipo, la cui origine risulta essere nord-europea (Belgio), e non Svizzera, come indicato nel protologo, né Italiana, come vuole una tradizione derivata dalla classica monografia di Kerner. Queste conclusioni comportano modificazioni consistenti nel significato del binomio, poiché le popolazioni nord-europee vanno riferite ad una entità tassonomica distinta rispetto a quelle italiane, almeno a livello di sottospecie. Il nome P. saccharata compete alla pianta nord-europea; il più antico nome validamente pubblicato per le popolazioni italiane è P. pictà Rouy.
Article
IAHOPA, an overlay program package performing intersection analysis and information analysis has been applied to a large data set of relevs of beech woods in the Apennines (Italy) completed by several authors following the Braun Blanquet approach. The results have been treated by several numerical methods testing classification efficiency and predictivity. Ecological indicator values have been used to test for predictivity. The classification proposed by Gentile has been confirmed in its main lines. However 2 new associations (Polysticho-Fagetum and Digitali-Fagetum) and 12 new subassociations are described. Furthermore the Veronico-Fagetum Montacchini 1972 has been recognized also for the Apennines. The clusters corresponding to the association level could be classified in two main alliances: Geranio nodosi-Fagion and Geranio striati-Fagion as suggested by Gentile, however their syntaxonomical justification should be based on numerical comparisons of the data from the entire area of European beech woods.
La cartografia della vegetazione per la gestione del territorio”85–104
  • D Ubaldi
Plantarum historia universalis oxoniensis3 Oxonii
  • R Morison
Viterbo) (!). 2n = 26 + b: Appennino Ligure (SAUER, 1975)
  • Cimini
Cimini (Viterbo) (!). 2n = 26 + b: Appennino Ligure (SAUER, 1975). 2n = 28:
ltaly (Piemonte to Lazio). (fig. 3) SPECIMENS SEEN -Liguria: Scogna
  • General Distribution-France
GENERAL DISTRIBUTION-France (SE), ltaly (Piemonte to Lazio). (fig. 3) SPECIMENS SEEN -Liguria: Scogna (Sesta Godano), 600 m., 21.VI.1990, S. Ballelli (BOLO);
500 m., 21.VI.1990, 5, Ballelli (BOLO); a Romito, dintorni di La Spezia, su calcare, 30
  • Taglieto
Taglieto (Varese Ligure), 500 m., 21.VI.1990, 5, Ballelli (BOLO); a Romito, dintorni di La Spezia, su calcare, 30.III.1984, D. Marchetti (SIENA).
960 m., 1990, G. Cristofolini (BOLO)
  • Raticosa Passo Della
Passo della Raticosa (Bologna), 960 m., 1990, G. Cristofolini (BOLO).