A new butterwort species (Pinguicula, Lentibulariaceae) from Northern Apennine (Italy)

Abstract

A morphometric and taxonomic study of Pinguicula in Northern Apennine was carried out. A new species belonging to sect. Pinguicula, previously referred erroneously to P. vulgaris L. and/or P. leptoceras Rchb., is here described as P. christinae sp. nov. The taxonomic relationship of P. christinae with other octoploid species, such as P. apuana Casper and Ansaldi, P. fiorii Tammaro and Pace, and P. vulgaris s.l. (including also its central Italian endemic subspecies) is discussed.

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A new butterwort species (Pinguicula,
Lentibulariaceae) from Northern Apennine (Italy)
L. Peruzzia & G. Gestrib
a Dipartimento di Biologia, Unità di Botanica generale e sistematica, Università di Pisa, Italy
b Via Bonfiglioli, 30 59100 Prato, Italy
Accepted author version posted online: 05 Dec 2012.Published online: 10 Jan 2013.
To cite this article: L. Peruzzi & G. Gestri (2013) A new butterwort species (Pinguicula, Lentibulariaceae) from Northern
Apennine (Italy), Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the
Societa Botanica Italiana, 147:3, 692-703, DOI: 10.1080/11263504.2012.756073
To link to this article: http://dx.doi.org/10.1080/11263504.2012.756073
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A new butterwort species (Pinguicula, Lentibulariaceae) from Northern
Apennine (Italy)
L. PERUZZI
1
*& G. GESTRI
2
1
Dipartimento di Biologia, Unita
`di Botanica generale e sistematica, Universita
`di Pisa, Italy and
2
Via Bonfiglioli, 30 59100
Prato, Italy
Abstract
A morphometric and taxonomic study of Pinguicula in Northern Apennine was carried out. A new species belonging to sect.
Pinguicula, previously referred erroneously to P. vulgaris L. and/or P. leptoceras Rchb., is here described as P. christinae sp. nov.
The taxonomic relationship of P. christinae with other octoploid species, such as P. apuana Casper and Ansaldi, P. fiorii
Tammaro and Pace, and P. vulgaris s.l. (including also its central Italian endemic subspecies) is discussed.
Keywords: Emilia-Romagna, endemics, Italy, morphometrics, Pinguicula, Tuscany
Introduction
In recent years, increasing efforts in the taxonomic
study of Italian vascular flora allowed the description of
several new taxa (e.g. Conti 2010;Ernandes et al.
2010;Troia & Raimondo 2010;Barberis & Nardi
2011;Nardi 2011;Peruzzi 2011;Peruzzi & Carta
2011;Peruzzi et al. 2011;Bacchetta et al. 2012;
Castellano et al. 2012;Cataldo et al. 2012;Marino
et al. 2012;Selvi & Sutory
´2012), including Pinguicula
(Conti & Peruzzi 2006;Ansaldi & Casper 2009).
Pinguicula (butterworts) is the second most diverse
genus of the carnivorous family Lentibulariaceae, with
about 100 currently accepted species (Rodondi et al.
2010). A number of them were only recently
recognized, especially in consequence of taxonomic
studies in Central America (see literature cited in
Cieslak et al. 2005). Also in the Mediterranean area,
12 new taxa were described in the last 25 years
(Tammaro & Pace 1987;Romo et al. 1996;Zamora
et al. 1996;Casper & Steiger 2001;Conti & Peruzzi
2006;Ansaldi & Casper 2009;Yildirim et al. 2012).
The family Lentibulariaceae Richard, order
Lamiales Bromhead, has recently been proved to be
monophyletic (Jobson et al. 2003), as the whole
genus Pinguicula (Cieslak et al. 2005;Degtjareva
et al. 2006;Kondo & Shimai 2006;Shimai & Kondo
2007;Shimai et al. 2007). The latter authors
contributed much to explain also the phylogenetic
relationships within this genus, showing that many of
the infrageneric taxa recognized by Casper (1966)
are poly- or para-phyletic. Recently, Degtjareva et al.
(2004) provided useful new taxonomic information
on seed morpho-anatomy, Rodondi et al. (2010)
studied pollen morphology in detail, while Peruzzi
(2004) and Casper and Stimper (2009) summarized
the karyological knowledge of this genus.
At present, 12 species are listed for Italy (Conti &
Peruzzi 2006;Pascal et al. 2008;Ansaldi & Casper
2009;Compostella et al. 2010). P. alpina L., P. arvetii
Auct. Fl. Ital. (non Genty), P. grandiflora Lam.,
P. leptoceras Rchb. and P. p o l d i n i i Casper and Steiger, are
all limited or endemic (the latter) to the Italian Alps;
P. reichenbachiana Schindler, occurs in central-western
Liguria; P. apuana Casper and Ansaldi and P. m a r i a e
Casper, are both endemic to Apuan Alps (Tuscany),
P. fiorii Tammaro and Pace and P. vallis-regiae F. Conti
and Peruzzi, are both endemic to Abruzzo, P. hirtiflora
Ten., occurs onlyin Campania and Calabria (Southern
Italy). The latter species is also the only one
overwintering as a rosette (tropical growth-type) in
Italy (Peruzzi et al. 2004). Besides them, the species
with the widest distribution range in Italy is P. vulgaris
L., occurring in Alps throughout the peninsula up to
Central Italy (see Conti & Peruzzi 2006 for further
information on endemic subspecies at its Italian
southern limit of distribution). In this picture, a still
unsolved problem is represented by the identity of the
q2013 Societa
`Botanica Italiana
Correspondence: L. Peruzzi, Dipartimento di Biologia, Unita
`di Botanica, Universita
`di Pisa, via Luca Ghini 5, 56126 Pisa, Italy. Tel: þ39 (0)502211339.
Fax: þ39 (0)502211309. Email: lperuzzi@biologia.unipi.it
Plant Biosystems, 2013
Vol. 147, No. 3, 692–703, http://dx.doi.org/10.1080/11263504.2012.756073
Downloaded by [Universita Di Pisa] at 06:54 29 October 2013

butterworts from Northern Apennine (see also Peruzzi
2005,2007a): they were reported as P. leptoceras by
Casper (1966) and Pignatti (1982),asP. vulgaris by
Alessandrini et al. (2003) and Conti et al. (2005) and
finally as P. cfr. vulgaris by Casper and Stimper (2006,
2009). While P. l e p t o c e r a s Rchb. is always tetraploid
with 2n¼32 chromosomes (Casper 1966;Casper &
Stimper 2009), P. vulgaris L. is octoploid with 2n¼64
(Casper 1966;Caparelli et al. 2008;Casper & Stimper
2009). Northern Apennine plants resulted 8x(rarely
also 16x;Casper & Stimper 2006), but at first sight
looked somehow different from the typical P. vulgaris
and its known subspecies.
For this reason, we carried out a taxonomical
study of Northern Apennine butterworts, by means
of macro- and micromorphology, aiming (a) to verify
the identity of the plants and (b) to investigate their
taxonomic relationships.
Materials and methods
Plant general morphology
General observations (tens of individuals for each
studied population) were carried out on (a) growth
type, (b) rosette type, and (c) colouring patterns of
the corolla and indicative opening angles between
upper and lower lip. Moreover, we studied on 15–20
individuals for each studied APPEN population
(Table I): (a) shape, length and width of leaves;
(b) scape size; and (c) shape and length of capsule.
Floral morphometr y
Floral morphometry was studied by measuring in the
field 10 quantitative continuous floral characters on
15–20 individuals from each considered population
(Ta b l e I ): (a) corolla length; (b) spur length;
(c) upper lobes length; (d) upper lobes width; (e)
lower lip central lobe length; (f) lower lip central lobe
width; (g) lower lip lateral lobes length; (h) lower lip
lateral lobes width; (i) calyx upper lip length; and (j)
calyx lower lip length. Floral morphometric data
about Northern Apennine Pinguicula were compared
with those of other apparently close octoploid taxa
(P. apuana,P. fiorii,P. vulgaris s.l.; see Ansaldi &
Casper 2009;Casper & Stimper 2009). The data
about P. fiorii and P. vulgaris s.l. (a total of 8– 64
individuals, respectively) were derived from Conti
and Peruzzi (2006), while those about P. apuana
(32 individuals) are original. The sampling of
Table I. Sampled populations of Pinguicula from Northern Apennine (APPEN) and other octoploid similar taxa.
Locality Habitat Substrate Elevation Coordinates (m a.s.l.)
APPEN
Northern Apennine:
Val di Luce stream siliceous 1510 448070N108370E
Val di Luce bog siliceous 1620 448070N108380E
Foce di Campolino bog siliceous 1600 448070N108390E
Verginetta (Abetone) stream siliceous 1500–1700 448090N108400E
Lama Rossa bog siliceous 1460 448130N108220E
P. apuana (APU)
Apuan Alps:
Carcaraia meadow limestone 1710 448060N108160E
Pania della Croce/Omo Morto meadow limestone 1450 –1500 448020N108190E
Acquasparta meadow limestone 1100 448060N108060E
Lago Trombacco lake limestone 310 448020N108230E
Lago di Vagli lake limestone 550 448070N108140E
Foce di Pianza meadow limestone 1350 448070N108080E
Forno cliffs limestone 190 448040N108100E
Tre Fiumi cliffs limestone 800 448030N108160E
P. fiorii (FIO)
Majella:
Scrimacavallo cliffs limestone 2010–2040 428080N148070E
P. vulgaris (VULG)
Central Apennine:
[subsp. anzalonei ]
Piscicarello di Jenne cliffs limestone 495 418530N138080E
[subsp. ernica ]
Zompo Lo Schioppo cliffs limestone 730 418500N138240E
[suvsp. vestina]
Valle del Rio Arno bog limestone 1140 428300N138320E
Vado di Corno/Valle dell’Inferno cliffs limestone 1730 428270N138350E
[subsp. vulgaris ]
Fosso dell’Acero stream siliceous 1643 428350N138240E
A new species of Pinguicula 693
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P. vulgaris – a Circumboreal element – is certainly
limited in representativeness of the species, but
includes a central Italian population referred to
typical P. vulgaris and also populations from all the
three subspecies endemic to central Italy (Conti &
Peruzzi 2006)(Table I). Moreover, the obtained
morphological data are fully congruent with the
general description of the species provided by Casper
(1966).
The variables were processed singularly by means
of Kruskal – Wallis test comparing the same variable
among the four-considered species (N¼187, df
¼3) and t-test for independent samples comparing
the same variable among pairs of species (APPEN vs.
APU; APPEN vs. FIO; APPEN vs. VULG). Scatter
plots for selected pairs of variables and boxplots
expressing the variability of each character were
made. For morphometric multivariate analysis, we
considered simultaneously the 10 characters, and the
resulting data matrix (187 cases £10 characters) was
subjected to principal coordinate analysis (PCoA) by
using Euclidean distance as coefficient with no
standardization of data (data standardization by
range of variables or by standard deviation of variables
gave results identical to those here presented).
Seed morphology and anatomy
Seed general morphology was studied with binocular
lens on 10 seeds from each APPEN sample (Table I).
The following characters were considered: (a) total
length; (b) maximum width; (c) length of mycropilar
appendage; (d) ratio micropylar seed appendage/
seed length; and (e) presence/absence of chalazal end
appendage.
Cross and longitudinal sections were made by
hand razor and mounted in glycerine. The following
characters were considered: cotyledon features
(number, symmetry, type of aestivation).
Besides, four seeds from each sample were
studied with SEM. For SEM studies, the material
was coated by a gold thin layer, then observed and
photographed at 10 kV. The following further
characters were considered: exotesta cells features
(number, shape, size).
Results
Plant general morphology
All the investigated plants from the five APPEN
populations represent the same biological unit and
clearly belong to Pinguicula sect. Pinguicula, sharing
several features such as temperate growth type
(overwintering as hibernaculum), homophyllous
rosettes (spring and summer leaves similar in shape
and dimensions), five to nine ovate to obovate-
oblong-triangular leaves, scapes and capsules of
comparable shape and size. For what concerns corolla
features, Northern Apennine plants are marked by a
peculiar combination of colouring patterns and
opening angles (Figure 1): blue-violet corolla with
irregular central white spot with white hairs near the
throat, often well open (opening angle (45)608–
1208), lower lip with lobes not overlapping and the
median one often distinctly bending downwards in
the front part.
Floral morphometr y
Plant from Northern Apennine (APPEN) showed
relatively large corollas, whose length resulted
intermediate among those of P. apuana (APU)/P.
fiorii (FIO) and P. vulgaris (VULG) (Figure 2A).
These differences resulted statistically significant
(Kruskal –Wallis ¼46.893, p,0.001). Concerning
the spur length, APPEN plants are very variable, but
with values generally higher than VULG (Figure 2B).
Also the differences in spur length resulted statisti-
cally significant among all the units (Kruskal– Wallis
¼51.080, p,0.001). Calyx dimensions are the
smallest in APPEN, compared with others (Figures
2C–D; Kruskal–Wallis ¼41.160, p,0.001 for
Figure 1. Pinguicula christinae sp. nov. Front view of flower (A); lateral view of flower (B); front view of calyx (C) and capsule (D).
L. Peruzzi and G. Gestri694
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upper lobe; 82.173, p,0.001 for lower lobe). The
dimensions of the upper corolla lobes are more
similar, but statistically different, among APPEN,
APU and FIO (Figures 3A –B;Kruskal–
Wallis ¼121.976, p,0.001 for length; 79.212,
p,0.001 for width). Lateral corolla lobes are the
smallest in APPEN compared with others (Figures
3C–D; Kruskal–Wallis ¼114.409, p,0.001 for
length; 116.781, p,0.001 for width), while median
corolla lobe dimensions are similar among APPEN
and VULG, but still statistically different (Figures
3E–F; Kruskal–Wallis ¼80.596, p,0.001 for
length; 92.402, p,0.001 for width).
All these floral features taken together allow a
clear morphological discrimination of Apennine
plants from other considered species, as can be
seen by PCoA analysis (Figure 4).
Concerning the t-test comparison of characters
for pairs of systematic units, APPEN is distinct from
APU in having smaller calyces and corollas, with
smaller central and lateral lobes (Table II). APPEN is
also distinct from FIO in having again smaller
calyces, central and lateral corolla lobes and shorter
upper lobe of corolla (Table III). Finally, APPEN is
distinct from VULG in having larger corollas and
spur but with narrower corolla central lobe and
smaller corolla lateral lobes and calyces (Table IV).
The separation of APPEN respect with other
systematic units is further confirmed by plotting just
pairs of the most relevant characters against each other
(Figure 5). Indeed, APPEN has no overlap with others
when the following characters are considered: corolla
lower lip median lobe length against corolla median
lobe width (for comparison with APU; Figure 5A);
corolla lower lip lateral lobes width against corolla
lower lip lateral lobes length (for comparison with FIO;
Figure 5B); and corolla upper lip lobes length against
corolla lower lip lateral lobes length are considered (for
comparison with VULG; Figure 5C).
Seed morphology, anatomy and morphometry
Seed general morphology resulted the same for all
the studied APPEN populations, albeit no significant
morphological variation was noticed either at within-
population or within-individual level (data not
shown). Seeds are minute, like sawdust, with
reticulate surface, elliptic; their length ranges
Figure 2. Boxplots illustrating the variability, in the studied units, of corolla length (A), spur length (B), calyx upper lobes length (C) and
calyx lower lobes length (D). Values are expressed in mm. The outlined central box depicts the middle 50% of the data extending from upper
to lower quartile; the horizontal bar is at the median. The ends of the vertical lines (or “whiskers”) indicate the minimum and maximum data
values, unless outliers are present in which case the whiskers extend to a maximum of 1.5 times the inter-quartile range. Superimposed grey
areas indicate confidence interval bounds around its median (median ^1.58 times the inter-quartile range). Circles indicate outliers, unless
extreme outliers are present in which case the circles extend to a maximum of three times the inter-quartile range and the extreme outliers are
indicated as asterisks.
A new species of Pinguicula 695
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between 658 and 804 mm and their width between
202 and 342 mm. The ratio micropylar seed
appendage/seed length ranges between 0.11 and
0.15. The number of exotesta cells is 200 – 260, and
they resulted 55 – 90 mm long and 27–57 mm wide;
the depth of anticlinal walls of exotesta cells is 5–
10 mm. SEM analysis of seeds also shows that all
populations share rounded-polygonal exotesta cells,
divided more or less evidently by a furrow (Figure 6).
No chalazal end appendage was evidenced. All the
investigated populations have only one plicate
cotyledon, slightly asymmetric in section.
Discussion
According to our results, APPEN has a peculiar
combination of characters, which allow a clear
circumscription of this unit respect with P. apuana
(APU), P. fiorii (FIO) and P. vulgaris (VULG), even
considering biometric floral features only. In par-
ticular, a smaller lower lip of corolla marks a clear-cut
distinction with P. apuana and P. fiorii (the latter
shows also a shorter upper lip of corolla). APPEN is
also clearly distinct from P. vulgaris s.l. (including the
endemic central Italian subspecies) for the longer
and wider upper lip of corolla, which has also shorter
and narrower lateral lobes in the lower lip, together
with a tendency to have longer spurs. Also the –
generally – well open flowers are a distinctive feature
respect with P. v u l g a r i s with the exception of
P. vulgaris subsp. anzalonei Peruzzi and F. Conti
(endemic to Latium), which however shows bio-
metric floral features falling completely within the
variability of P. vulgaris s.l. (Conti & Peruzzi 2006). It
Figure 3. Boxplots illustrating the variability, in the studied units, of corolla upper lobes length (A) and width (B); corolla lateral lobes length
(C) and width (D); corolla median lobe length (E) and width (F). Values are expressed in mm (see Figure 2 for more explications).
L. Peruzzi and G. Gestri696
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must be noticed that some extreme individual in
APPEN populations may occasionally show, at first
sight, a P. vulgaris-like appearance (i.e. less open
corolla), but when the flower parts are measured a
safe attribution to APPEN is always possible.
The seed features obtained for APPEN are those
typical of other octoploid taxa of Pinguicula sect.
Pinguicula (Conti & Peruzzi 2006;Ansaldi & Casper
2009), and are therefore not useful as taxonomic
markers in this case.
Other species of sect. Pinguicula, which show
some resemblance with APPEN, are the Alpine
endemic P. leptoceras and the Balkan endemic
P. balcanica Casper. However, both species are
tetraploid (2n¼32) and easily distinct from
APPEN by some morphological features: corolla
lower lip with overlapping lobes and larger capsules,
the former (Casper 1966), corolla upper lip with
overlapping lobes, larger lateral lobes of corolla lower
lip, the latter (Casper 1966;Peruzzi 2007b).
The systematic unit from Northern Apennines
(APPEN), as circumscribed in our study, certainly
deserves taxonomic recognition and – at the present
state of knowledge – the specific status seems the
most appropriate, by considering the allopatric
occurrence of all the studied units and the peculiar
Table II. Mean differences between APPEN and APU popu-
lations, according to t-test. Those characters with df ¼113 were
assumed with equal variances (after Levene statistics); the others
were not assumed with equal variances. Significant differences are
evidenced in bold.
tdf p
difference
(mm)
Corolla length 23.669 113 0.000 22.806
Spur length 0.190 113 0.850 0.406
Upper lobes length 0.868 113 0.387 0.208
Upper lobes width 20.832 113 0.407 20.139
Central lobe length 28.157 33.158 0.000 23.945
Central lobe width 211.159 35.279 0.000 23.241
Lateral lobes length 213.864 45.970 0.000 22.447
Lateral lobes width 210.822 37.844 0.000 22.128
Calyx upper lip length 25.848 40.018 0.000 21.491
Calyx lower lip length 26.525 34.866 0.000 21.532
Table III. Mean differences between APPEN and FIO popu-
lations, according to t-test. Those characters with df ¼89 were
assumed with equal variances (after Levene statistics); the others
were not assumed with equal variances. Significant differences are
evidenced in bold.
tDf p
difference
(mm)
Corolla length 21.446 89 0.152 21.906
Spur length 1.473 89 0.144 1.108
Upper lobes length 5.810 89 0.000 2.174
Upper lobes width 0.438 89 0.662 1.293
Central lobe length 26.370 89 0.000 21.857
Central lobe width 25.538 89 0.000 21.342
Lateral lobes length 217.245 89 0.000 24.663
Lateral lobes width 29.681 7.413 0.000 23.525
Calyx upper lip length 23.598 89 0.001 21.070
Calyx lower lip length 25.801 89 0.000 21.094
Figure 4. Scatter plot of first two principal coordinate axes, expressing the 65.5% of the variability, based on PCoA of all the 10 quantitative
morphological characters considered for the flowers. Coloured areas unit individuals belonging to the same species/unit.
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and unique morphological features of APPEN.
Therefore, it is described as a species new to science.
However, a molecular phylogeny of all the involved
taxa would be desirable to clarify the origin of
APPEN in relation with related units (especially
P. vulgaris and its subspecies). Moreover, a hybrid
(either homoploid or allopolyploid) origin for
APPEN cannot be ruled out. This could
have involved P. vulgaris and a further octoploid
species (i.e. homoploid hybrid speciation) or a pair of
tetraploid taxa (i.e. allopolyploid hybrid speciation).
Hence, Italy is doubtlessly a European diversity
hotspot for butterworts, hosting at present a total of
16 taxa, 9 of them endemic. The larger Iberian and
Balkan peninsulae host only 9 taxa, 5 of them
endemic (Blanca et al. 1999;Blanca 2001) and
3 taxa, 2 of them endemic (Casper 1966;Shuka et al.
2007), respectively.
Interestingly, there are other species narrow
endemic to Northern Apennine (Peruzzi et al.
2012): Festuca riccerii Foggi and Rossi (Poaceae;
Foggi & Rossi 1996), Hieracium boreoapenninum
Gottschlich, H. faucis-jovis Gottschlich, H.
lanudae Gottschlich, H. umbrosoides Gottschlich
(Asteraceae; Gottschlich 2009), Primula apennina
Widmer (Primulaceae; Crema et al. 2009;Fisogni
et al. 2011), Taraxacum aemilianum Foggi and Ricceri
(Asteraceae; Foggi & Ricceri 1991)andMurbeckiella
zanonii (Ball) Rothm. (Brassicaceae), albeit the latter
Table IV. Mean differences between APPEN and VULG
populations, according to t-test. Those characters with df ¼145
were assumed with equal variances (after Levene statistics); the
others were not assumed with equal variances. Significant
differences are evidenced in bold.
tdf p
difference
(mm)
Corolla length 3.772 122.587 0.000 1.726
Spur length 7.054 104.139 0.000 1.741
Upper lobes length 16.021 145 0.000 2.602
Upper lobes width 10.136 143.785 0.000 1.281
Central lobe length 2.038 108.055 0.044 0.342
Central lobe width 24.733 145 0.000 20.519
Lateral lobes length 210.535 102.436 0.000 21.621
Lateral lobes width 210.800 102.547 0.000 21.331
Calyx upper lip length 24.428 142.708 0.000 20.559
Calyx lower lip length 28.367 119.073 0.000 20.816
9.00
8.00
7.00
6.00
(a)
5.00
4.00
3.00
2.00
1.00
0.00
0.00 2.00 4.00 6.00
Corolla bottom lip median lobe length (mm)
Corolla lower lip lateral lobes width (mm)
8.00 10.00 12.00 14.00 16.00 18.00
APU
APPEN
12.00
10.00
8.00
6.00
(b)
4.00
2.00
0.00
0.00 1.00 2.00 3.00
Corolla bottom lip lateral lobes width (mm)
Corolla lower lip lateral lobes length (mm)
4.00 5.00 6.00 7.00 8.00 9.00
FIO
APPEN
9.00
7.00
8.00
6.00
(c)
4.00
5.00
2.00
1.00
3.00
0.00
0.00 2.00
Corolla upper lip lobes len
g
th (mm)
Corolla lower lip lateral lobes length (mm)
4.00 6.00 8.00 10.00 12.00
VULG
APPEN
Figure 5. Scatter plots of measured individuals, against selected pairs of variables. A: APPEN (N¼83) and APU (N¼32), corolla lower lip
median lobe length (mm) (xaxis) plotted against corolla median lobe width (mm) (yaxis). B: APPEN (N¼83) and FIO (N¼8), corolla
lower lip lateral lobes width (mm) (xaxis) plotted against corolla lower lip lateral lobes length (mm) (yaxis). C: APPEN (N¼83) and VULG
(N¼64), corolla upper lip lobes length (mm) (xaxis) plotted against corolla lower lip lateral lobes length (mm) (yaxis).
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species was also recently recorded from France (J.-
M. Tison in litt.). A number of other taxa are
endemic of Northern Apennine and neighbouring
Apuan Alps: Armeria marginata (Levier) Bianchini
(Plumbaginaceae), Cirsium bertolonii Spreng. (Aster-
aceae; Zanotti & Cristofolini 1985), Festuca violacea
Schleich. ex Gaudin. subsp. puccinellii Foggi, Rossi
and Signorini (Poaceae; Foggi et al. 1999), Globu-
laria incanescens Viv. (Plantaginaceae), Leontodon
anomalus Ball (Asteraceae), Moltkia suffruticosa (L.)
Brand subsp. bigazziana Peruzzi and Soldano
(Boraginaceae), Pinguicula mariae Casper (Lentibu-
lariaceae), Polygala carueliana (Benn.) Burnat (Poly-
galaceae), Rhamnus glaucophylla Sommier
(Rhamnaceae; Bedini et al. 2011), Rhinanthus
apuanus Soldano (Orobanchaceae), Thesium som-
mieri Hendrych (Thesiaceae) and Viola ferrarinii
Moraldo and Ricceri (Violaceae; Moraldo et al.
2011). This accounts for a great biogeographic and
evolutionary relevance of this area.
Description of the new species
P. christinae Peruzzi and Gestri sp. nov. (Figure 1).
Diagnosis
Planta (29)64–150(200) mm alta. Rosula homo-
phylla; folia (16)20–50(75) mm longa. Corolla saepe
ampliata, (45)608–1208angulo aperturae, violaceo-
caerulea, centro saepe albo, (15)19 – 25.9(31) mm
longa; lobis labii inferi oblongis, inter se non tegentis;
lobis lateralis labii inferioris (2)2.1 – 3.9(4) mm latis
et 3–5(6) mm longis, lobis labii superioris (3)4 – 6(7)
mm latis et (4)5–7(10) mm longis, calcar (3)5–9.9
(13) mm longum; labium superior calycis triparti-
tum; lobis labii superi elliptico-triangularibus; Cap-
sula ovoidea.
Holotypus
Foce di Campolino (province of Pistoia), bog on
siliceous substrate, 1600 m a.s.l., 4 July 2011,
L. Peruzzi et G. Gestri (PI).
Paratypi
Val di Luce (province of Pistoia), along a stream on
siliceous substrate, 1510 m a.s.l., 13 June 2010,
G. Gestri (PI); Val di Luce (province of Pistoia),
along a stream on siliceous substrate, 1620 m a.s.l.,
13 June 2010, G. Gestri (PI); Lama Rossa (province
of Lucca), bog on siliceous substrate, 1460 m a.s.l., 2
June 2011, G. Gestri (PI); Verginetta (Abetone),
province of Pistoia, along a stream on siliceous
substrate, 1500 m a.s.l., 5 June 2011, G. Gestri (PI);
Verginetta (Abetone), province of Pistoia, along a
stream on siliceous substrate, 1550 –1700 m a.s.l., 17
July 2011, G. Gestri (PI).
Description
Herb perennial, small, rosette forming, scapose,
succulent. Stem short, with ascending, not branch-
ing rhizome and numerous adventitious fibrous
roots. Rosettes with few, 5–7(9) leaves lying more
or less flat on the ground, homophyllous. Over-
wintering as buds (hibernacula). Leaves in outline
obovate-oblong-triangular, (16)20–50(75) mm
long, (10)11 – 20(30) mm broad. The margin entire,
Figure 6. P. christinae sp. nov., examples of variability in seed shape and structure, by means of SEM microphotographs of seeds (general view
and particular of exotesta cells): Foce di Campolino population (A –B); Lama Rossa population (C–D).
A new species of Pinguicula 699
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slightly incurved; brittle; the upper surface densely
covered with mucilaginous sessile and stalked glands.
Scapes 1–3, erect, (29)64–150(200) mm tall, terete,
1-flowered, directly beneath the flower densely
covered with stipitate glands, to the base sparsely
glandular. Flowers relatively large, (15)19 – 25.9(31)
mm long (spur included). Calyx distinctly bilabiate,
densely covered on both surfaces and the margins
Figure 7. Pinguicula christinae sp. nov. Map, with 10 £10 Km UTM grid, showing the geographic distribution of the new species.
L. Peruzzi and G. Gestri700
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with stipitate glands, 2.5 – 7.5 mm in total diameter,
upper lip divided nearly to 1/2 –2/3 into 3 lobes
elliptic triangular, the central one sometimes bifid;
lower lip divided to 1/2 into 2 short lobes, sometimes
irregular. Throat relatively densely covered with
white clavate hairs. Spur slightly curved downwards,
thin, (3)5 – 9.9(13) mm long. Corolla distinctly
bilabiate, blue-violet, white at the centre, at the
throat white, spur blue-violet; upper lip with two
nearly identical lobes not overlapping, (4)5 – 7(10)
mm long and (3)4 –6(7) mm broad, erect to patent,
apex subtruncate-rounded to slightly emarginate;
lower lip much larger than the upper lip, with three
lobes not overlapping, the middle lobe distinctly
longer and broader than the lateral lobes, apex
rounded to slightly truncate, front part of the median
lobe often distinctly bending downwards: the whole
corolla appears often well open [opening angle (45)
608–1208]. Capsule ovoid-conical, (3)4–6(7) mm
long, 2 – 4(4.5) mm in diameter, 1-loculate. Seeds
minute, like sawdust, with reticulate surface, elliptic,
658 –804 mm long, 202 –342 mm wide. Ratio micro-
pylar seed appendage/seed length: 0.11 – 0.15. Num-
ber of exotesta cells 200–260, exotesta cells 55–
90 mm long, 27 –57 mm wide; depth of anticlinal
walls of exotesta cells 5 – 10 mm. Cotyledon 1, plicate,
asymmetric. Pollen spheroidal, 4 –6 colporate, 25–
33 mm. Flowering V–VI; fruiting VII –VIII.
Etymology
The name of this delicate species is dedicated to
Cristina Gavazzi, wife of one of the authors (GG)
and native from Northern Apennines, too.
Ecology
Marshes, bogs and humid grasslands, from 1000 to
1800 m a.s.l., on siliceous substrates.
Karyology
The populations from Lama Rossa and Verginetta
(also studied morphologically here) were checked
concerning chromosome number by Casper and
Stimper (2006; see also the website “Chrobase.it”,
Bedini et al. 2012). They resulted as 8xwith 2n¼64
and 16xwith 2n¼128 chromosomes, respectively.
Distribution
The new species is presently known only for Northern
Apennine (Figure 7), in both Tuscany (provinces of
Lucca and Pistoia) and Emilia-Romagna (provinces of
Parma, Reggio Emilia, Modena and Bologna). As far as
we are aware, this is the only butterwort species
occurring at high altitudes in Northern Apennine. On
the contrary, at lower altitude, on limestone cliffs
(Orrido di Botri gorges, Tuscany, province of Lucca),
also the morphologically and karyologically distant P.
mariae Casper occurs. The populations from Ligurian
Apennines (about 80km far from the known range of P.
christinae) will need further studies to clarify their
identity, and could pertain to the new species.
Presently, P. v u l g a r i s (subsp. vulgaris) is recorded with
reasonable certainty only for Alps and Central
Apennines.
Identification key for Italian octoploid species of
Pinguicula sect. Pinguicula.*See Conti and Peruzzi
(2006) for further criteria aimed for identifying the
subspecies of P. vulgaris.
1. Corolla opening angle 308–808....................... 2.
1. Corolla opening angle $908........................... 3.
2. Corolla upper lip $5 mm in length, with lateral
lobes #6 mm in length. .......................P. christinae
2. Corolla upper lip ,5 mm in length or, if above
5 mm, with lateral lobes .6 mm in length .......... 4.
3. Corolla uniformly blue-violet, generally with no
white spots (just showing whitish throat), lateral
lobes generally .7 mm in length ................. P. fiorii
3. Corolla with coloration pattern not as above, with
lateral lobes #7 mm in length ............................ 5.
4. Ratio of corolla lateral/upper lip length .1...
P. vulgaris s.l.*
4. Ratio of corolla lateral/upper lip
length #1......................... P. c h r i s t i n a e
5. Corolla with a white V-shaped dot, surrounded of
deeper blue-violet on median lower lip, with
upper lip #4 mm in length . ......... P. vulgaris
(subsp. anzalonei)
5. Corolla with coloration pattern not as above, with
upper lip $4.5 mm ........... ................................. 6.
6. Corolla with lower lip lateral lobes $4 mm wide; if
4 mm, median lobe .6mm.......... P. apuana
6. Corolla with lower lip lateral lobes #4 mm wide; if
4 mm, median lobe #6mm.........P. christinae
Acknowledgements
The authors wish to thank Maria Ansaldi (University
of Pisa) for collecting the data about P. apuana,
Andrea Vitali for logistic facilities and three
anonymous referees for the significant improvement
of an earlier version of the manuscript. Financial
funding (EX60%) from the University of Pisa is
gratefully acknowledged.
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