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Phylogenetic relationships among genera in Boraginaceae tribe Boragineae (simplified, after Hilger et al. 2004; Weigend et al. 2010). Taxa represented in the Italian flora are in bold. 

Phylogenetic relationships among genera in Boraginaceae tribe Boragineae (simplified, after Hilger et al. 2004; Weigend et al. 2010). Taxa represented in the Italian flora are in bold. 

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A synopsis of the Italian taxa of tribe Boragineae (Boraginaceae, subfam. Boraginoideae) is given as a second contribution to the treatment of the family for the Flora Critica d'Italia project. The work is mainly based on the critical study of herbarium material and extensive literature survey. All relevant floristic reports were examined and types...

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... the order “ Boraginales ”, the Boraginaceae s.s. ( Boraginaceae s.l. subfam. Boraginoideae ) form a well- supported monophyletic clade that can be easily distinguished by the gynobasic style and the eremocarpic fruit (Weigend et al. 2013; Hilger 2014). This clade includes 2/3 of the total amount of genera and species in the Boraginaceae s.l. (sensu APG 2009) and consists of four major tribes based on both morphological characters (mainly fruit) and DNA data: Boragineae, Cynoglosseae, Echiochileae and Lithospermeae (L ̊ngstro ̈ m & Chase 2002; Weigend et al. 2013; Figure 1). The members of tribe Boragineae are characterized by prominent faucal scales (in Pulmonaria L. replaced by patches or bands of trichomes), flat gynobase and, usually ovoid nutlets with non-calcified pericarp, provided with a basal thickening and lipidic elaiosome for myrmecochorous dispersal. They are mainly centered in the Old World, with main diversity centers in the Mediterranean and Irano-Turanian regions, but also include as basal groups the members of two genera from the Neotropics, i.e. Moritzia DC. ex Meisn. and Thaumatocaryon Baill. (Weigend et al. 2010). Follow- ing the description of new taxa, the appearence of new allochtonous species over the national territory and several taxonomic re-arrangements, the number of genera and species of Boragineae reported for Italy (see tab. I in Cecchi & Selvi 2014) has doubled from Flora Italica by Bertoloni (1835–1836; five genera with 18 species) to the most recent report in the Checklist by Conti et al. (2005; 12 genera with 35 species). Following our first contribution dealing with the “atypical”, basal groups of Boraginaceae s.l. (members of subfamilies Hydrophylloideae and Heliotropioideae ; Cecchi & Selvi 2014), we present here the first of three synopses dedicated to the Boraginaceae s.s. (i.e. Boraginaceae subfam. Boraginoideae sensu APG 2009), one for each tribe. As in the case of the previous one, our work is mainly based on a detailed literature survey and on the study of the herbarium material conserved in the great majority of the Italian collections, both public and private. Observations on natural populations made during numerous field excursions have provided additional elements to evaluate the extent and patterns of infraspecific variation in a number of taxa. All the relevant ...

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... & Puppi, another Italian endemic widely distributed along the Apennines. 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). ...
... This is also supported by the fact that all the populations showing intermediate features occur in those areas where P. hirta s.str. and P. apennina are reported to overlap (Puppi & Cristofolini, 1996;Vosa & Pistolesi, 2004;Cecchi & Selvi, 2015). Accordingly, we expected to find a similar pattern in AFLP data, reflecting a genetic differentiation between P. hirta s.str. ...
... Under the third scenario, the "vallarsoid" morph originated through descending dysploidy from "hirtoid" plants or through backcross of these latter plants with Pulmonaria officinalis. Once originated, "vallarsoid" plants spread across the Italian peninsula, maintaining connection (gene flow) with "hirtoid" plants in central Italy, so that pure "vallarsoid" morphs remained only at the southern and northern extremes of the range, where "hirtoid" plants have never been reported (Cecchi & Selvi, 2015;Bartolucci & al., 2018). The STRUC-TURE analysis shows that the genetic group dominant in the populations of P. hirta s.str. ...
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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.
... The experiments were carried out on two dozens of plants of Pulmonaria vallarsae Kerner subsp. apennina Cecchi et Selvi (Italian lungwort, family Boraginaceae), a perennial understory herb, relatively shade tolerant and widely distributed on the Italian peninsula along Apennine reliefs (Cecchi 2015). The plants were collected in several hilly sites of Content courtesy of Springer Nature, terms of use apply. ...
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Although many photosynthesis related processes are known to be controlled by the circadian system, consequent changes in photosynthetic activities are poorly understood. Photosynthesis was investigated during the daily cycle by chlorophyll fluorescence using a PAM fluorometer in Pulmonaria vallarsae subsp. apennina , an understory herb. A standard test consists of a light induction pretreatment followed by light response curve (LRC). Comparison of the major diagnostic parameters collected during day and night showed a nocturnal drop of photosynthetic responses, more evident in water-limited plants and consisting of: (i) strong reduction of flash-induced fluorescence peaks (FIP), maximum linear electron transport rate (J max , ETR EM ) and effective PSII quantum yield (Φ PSII ); (ii) strong enhancement of nonphotochemical quenching ( NPQ ) and (iii) little or no change in photochemical quenching qP , maximum quantum yield of linear electron transport ( Φ ), and shape of LRC ( θ ). A remarkable feature of day/night LRCs at moderate to high irradiance was their linear-parallel course in double-reciprocal plots. Photosynthesis was also monitored in plants subjected to 2–3 days of continuous darkness (“long night”). In such conditions, plants exhibited high but declining peaks of photosynthetic activity during subjective days and a low, constant value with elevated NPQ during subjective night tests. The photosynthetic parameters recorded in subjective days in artificial darkness resembled those under natural day conditions. On the basis of the evidence, we suggest a circadian component and a biochemical feedback inhibition to explain the night depression of photosynthesis in P. vallarsae .
... 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. ...
... 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.
... During the field work for determination of Aghdagh protected area and riversides floras in Ardabil province, some specimens of Nonea were collected. The collected specimens were crosschecked with various Flora and relevant literatures (Boissier 1879, Popov 1953, Riedl 1967, Baytop 1979, Chater 1972, Selvi & Bigazzi 2001, Khatamsaz 2002, Nejhad Falatoury & al. 2011, Ahmad 2014, Cecchi & Selvi, 2015and Mathieu 2019. As well the photos of the herbarium specimens of Nonea in E, P, B and W herbaria (acronyms according to Thiers 2019) were consulted in order to confirm the identity of the new species record. ...
... without specified specimen from Iran. According to latest literature (Cecchi & Selvi 2015, POWO 2019, Mathieu 2019) N. ventricosa is a synonym of N. echioides which has not previously been mentioned in the literature about Iran's flora. ...
Article
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Nonea echioides, a species native to the Mediterranean area, is reported from Aras and Qizil Üzan riversides as a rediscovered species for the flora of Iran. A detailed description and photographs of diagnostic features are provided as well as some notes on its micromorphology, distribution and the last state of nomenclature are presented.
... A specimen collected in Camaldoli (Toscana) was selected as the epitype for the name P. hirta by Selvi in Cafferty and Jarvis (2004). 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. ...
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.
... Smaller islets around Elba island were not considered because their flora was already studied and published by Foggi et al. (2009a). In addition to the papers cited in the Introduction, the following references were checked: Koestlin (1780), Thiébaut De Berneaud (1808), Caruel (1860, 1870), Bolzon (1893Bolzon ( , 1894, Baroni (1897-1908), Fiori (1923-1929, 1943, Corti (1940), Marcello (1951), Ottens (1967), Arrigoni (1976Arrigoni ( , 2016Arrigoni ( , 2017, Jalas and Suominen (1976), Paoli and Romagnoli (1976), Zangheri (1976), Viegi and Cela Renzoni (1981), Pignatti (1982a-b), Nardi (1984), Natali (1988), Banfi (1989Banfi ( , 2017, Landi (1989), Gatteschi and Arretini (1990), Alessandro et al. (1991), Corsi and Garbari (1991), Baldini (1993), Del Prete and Tosi (1994), Bussotti et al. (1997), Hofmann et al. (1998), Rizzotto (1999, Vagge and Biondi (1999), Rinaldi (2000), Breiner (2001, 2002), Del Prete (2001), Adamoli and Rigon (2003), Pignotti (2003), Ackermann and Ackermann (2004), Frangini (2004), Group of European Pteridologists (2004), Fanelli and Tescarollo (2005), Frangini et al. (2005Frangini et al. ( , 2006, Tavormina (2006), Atzori (2007), Barsotti (2008), Bedini et al. (2008), Guiggi (2008Guiggi ( , 2014, Hoffmann and Hoffmann (2008), Iamonico et al. (2008, Foggi and Venturi (2010), Arrigoni and Viegi (2011), Frignani and Iiriti (2011, , Lazzaro et al. (2013aLazzaro et al. ( -b, 2014a, Ardenghi et al. (2014Ardenghi et al. ( , 2015, Bagella et al. (2015), Cecchi and Selvi (2015), Iamonico (2015), Signorini and Tani (2015), Roma-Marzio et al. (2016, Scoppola et al. (2016), Benítez-Benítez et al. (2017), Martini and Viciani (2018), Troia et al. (2018). ...
Article
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We present an updated list of the vascular flora occurring on the Elba island (Tuscan Archipelago). The list is based on bibliographic analysis and field studies carried out in the years 2006–2018. With a total of 1,098 specific and subspecific taxa currently occurring on the island (including 101 naturalized aliens), plus 67 casual aliens and 16 hybrid taxa, Elba shows the highest number of species among the islands of the Tuscan Archipelago. Two taxa are new for Tuscany: Hieracium symphytaceum s.l. and Ophrys exaltata subsp. morisii; 22 taxa are new for the island, 34 have been confirmed, while 326 were reliably recorded previously by other authors, but not confirmed by our study. We excluded 41 taxa and considered doubtful the occurrence of 87. Life forms and chorotypes are in agreement with the Mediterranean climate of the island. Despite this, Elba also hosts a considerable proportion of Eurosiberian taxa. We detected significant differences in chorotypes and life forms spectra among different geographical portions of the island, paralleling distinct bioclimatic patterns. Despite the institution of the Tuscan Archipelago National Park, we are still far from an integrated protection of the island flora. Based on our results, it has been possible to arrange a geodatabase of the flora on the island, useful for its protection.
... In Italy, it certainly occurs in moist broadleaf woods of the northern regions, including NW Toscana, and in the central Apennines (Abruzzo) ( Bartolucci et al. 2012, Cecchi and. On the other hand, it was historically recorded from the Marche (Paolucci 1890-1891), but its presence in this administrative region was excluded by later authors (Pignatti 1982, Conti et al. 2005, Ballelli et al. 2010, Cecchi and Selvi 2015. Our finding confirms its presence in the Marche and particularly in one of the sites where it was historically recorded, i.e., Montefortino (Paolucci 1890-1891). ...
Article
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In this contribution new data concerning the Italian distribution of native vascular flora are presented. It includes new records, exclusions, and confirmations to the Italian administrative regions for taxa in the genera Arctostaphylos, Artemisia, Buglossoides, Convolvulus, Crocus, Damasonium, Epipogium, Ficaria, Filago, Genista, Heptaptera, Heracleum, Heteropogon, Hieracium, Myosotis, Ononis, Papaver, Pilosella, Polygonum, Pulmonaria, Scorzonera, Silene, Trifolium, Vicia and Viola.
... Taxonomic note:-The nomenclature follows Cecchi & Selvi (2015). Symphytum officinale L. ...
Article
The National Park of Gran Sasso and Monti della Laga is located in central Italy and covers an area of 143,311 ha, across three administrative regions (Abruzzo, Marche and Lazio). To date, there is not a comprehensive floristic study concernig the Park’s territory but only partial contributions regarding specific taxa or restricted sectors. An annotated checklist of the vascular plant species growing in this territory is here presented, based on field surveys carried out from 1993 to 2015, on extensive analysis of relevant literature and on a review of specimens kept in APP, FI, NAP and RO herbaria. The Park’s flora amounts to 2,642 species and subspecies grouped in 739 genera and 121 families. Two hundred twenty nine taxa are endemic to Italy. Seventy seven taxa are newly recorded for Gran Sasso and Monti della Laga National Park, 83 taxa are not confirmed in recent times and, 57 are doubtuful. The non-native plants amount to 137 taxa. The number of taxa recorded in the Gran Sasso and Monti della Laga National Park is the highest ever found in a protected area of the Mediterranean Basin.
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Background The genus Pulmonaria (Boraginaceae) represents a taxonomically complex group of species in which morphological similarity contrasts with striking karyological variability. The presence of different numbers of chromosomes in the diploid state suggests multiple hybridization/polyploidization events followed by chromosome rearrangements (dysploidy). Unfortunately, the phylogenetic relationships and evolution of the genome, have not yet been elucidated. Our study focused on the P. officinalis group, the most widespread species complex, which includes two morphologically similar species that differ in chromosome number, i.e. P. obscura (2n = 14) and P. officinalis (2n = 16). Ornamental cultivars, morphologically similar to P. officinalis (garden escapes), whose origin is unclear, were also studied. Here, we present a pilot study on genome size and repeatome dynamics of these closely related species in order to gain new information on their genome and chromosome structure. Results Flow cytometry confirmed a significant difference in genome size between P. obscura and P. officinalis, corresponding to the number of chromosomes. Genome-wide repeatome analysis performed on partial Illumina sequencing data showed that retrotransposons were the most abundant repeat type, with a higher proportion of Ty3/Gypsy elements, mainly represented by the Tekay lineage. Comparative analysis revealed no species-specific retrotransposons or striking differences in their copy number between the species. A new set of chromosome-specific cytogenetic landmarks, represented by satellite DNAs, showed that the chromosome structure in P. officinalis was more variable compared to that of P. obscura. Comparative karyotyping strongly supported the hybrid origin of putative hybrids with 2n = 15 collected from a mixed population of both species and outlined the origin of ornamental garden escapes, confirming their derivation from the P. officinalis complex. Conclusions Large-scale genome size analysis and repeatome characterization of the two morphologically similar species of the P. officinalis group improved our knowledge of the genome dynamics and differences in the karyotype structure. A new set of chromosome-specific cytogenetic landmarks was identified and used to reveal the origin of putative hybrids and ornamental cultivars morphologically similar to P. officinalis.
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We present an updated list of the vascular flora occurring in the Cerbaie hills (Tuscany), a site of high naturalistic interest. The list is based on a literature survey and on field studies carried out in the years 2010–2022. The Cerbaie hills host a flora of 1,107 specific and subspecific taxa (including 100 naturalized aliens), 32 casual aliens and 10 hybrid taxa. Two taxa are new for Tuscany: Carex oedipostyla and Thalictrum simplex subsp. galioides; 330 taxa are new for the study area. Concerning old records, 344 have been confirmed, while 47 were not confirmed, albeit considered reliable. Moreover, we considered three taxa as locally extinct, 19 as doubtfully occurring, and three as wrongly reported. Despite the low elevation of the study area, life forms and chorotypes show marked Eurosiberian affinities, in agreement with the temperate and continental climate.