By relating genetic divergence at neutral loci, phenotypic variation, and geographic and environmental distances, it is possible to dissect micro-evolutionary scenarios involving natural selection and neutral evolution. In this work, we tested the patterns of intraspecific genetic and phenotypic variation along an elevational gradient, using Dianthus virgineus as study system. We genotyped genome-wide SNPs through ddRAD sequencing and quantified phenotypic variation through multivariate morphological variation. We assessed patterns of variation by testing the statistical association between genetic, phenotypic, geographic, and elevational distances and explored the role of genetic drift and selection by comparing the Fst and Pst of morphometric traits. We revealed a weak genetic structure related to geographic distance among populations, but we excluded the predominant role of genetic drift acting on phenotypic traits. A high degree of phenotypic differentiation with respect to genetic divergence at neutral loci allowed us to hypothesize the effect of selection, putatively fuelled by changing conditions at different sites, on morphological traits. Thus, natural selection acting despite low genetic divergence at neutral loci can be hypothesized as a putative driver explaining the observed patterns of variation.
The herbarium digitization process is an essential first step in transforming the vast amount of data associated with a physical specimen into flexible digital data formats. In this framework, the Herbarium of the University of Pisa (international code PI), at the end of 2018 started a process of digitization focusing on one of its most relevant collections: the Herbarium of Michele Guadagno (1878-1930). This scholar studied flora and vegetation of different areas of southern Italy, building a large herbarium including specimens collected by himself, plus many specimens obtained through exchanges with Italian and foreign botanists. The Herbarium is composed by 547 packages of vascular plants. Metadata were entered into the online database Virtual Herbaria JACQ and mirrored into a personalized virtual Herbarium of the Botanic Museum. After the completion of the digitization process, the number of sheets preserved in the Herbarium amounts to 44,345. Besides Guadagno, who collected 42% of his specimens, a further 1,102 collectors are represented. Most specimens were collected in Europe (91%), but all the continents are represented. As expected, Italy is the most represented country (59%), followed by France, Spain, Germany, and Greece. The specimens cover a time span of 99 years, from 1830 to 1929, whereas the specimens collected by Guadagno range between 1889 and 1928. Furthermore, we traced 134 herbarium sheets associated with documents, among which 75 drawings handmade by Guadagno, 34 letters from various corresponding authors, 16 copies of publications, and 14 copies of published iconographies.
Sardinia and Corsica are two Mediterranean islands where the genus Armeria is represented by 11 taxa, 10 out of which are endemic. An integrative approach, using molecular phylogeny, karyology, and seed and plant morphometry was used to resolve the complex taxonomy and systematics in this group. We found that several taxa are no longer supported by newly produced data. Accordingly, we describe a new taxonomic hypothesis that only considers five species: Armeria leucocephala and A. soleirolii, endemic to Corsica, and A. morisii, A. sardoa, and A. sulcitana, endemic to Sardinia.
Systematics and taxonomy are basic sciences and are crucial for all applications dealing with living organisms [...].
Invasive alien species are among the main global drivers of biodiversity loss posing major challenges to nature conservation and to managers of protected areas. The present study applied a methodological framework that combined invasive Species Distribution Models, based on propagule pressure, abiotic and biotic factors for 14 invasive alien plants of Union concern in Italy, with the local interpretable model-agnostic explanation analysis aiming to map, evaluate and analyse the risk of plant invasions across the country, inside and outside the network of protected areas. Using a hierarchical invasive Species Distribution Model, we explored the combined effect of propagule pressure, abiotic and biotic factors on shaping invasive alien plant occurrence across three biogeographic regions (Alpine, Continental, and Mediterranean) and realms (terrestrial and aquatic) in Italy. We disentangled the role of propagule pressure, abiotic and biotic factors on invasive alien plant distribution and projected invasion risk maps. We compared the risk posed by invasive alien plants inside and outside protected areas. Invasive alien plant distribution varied across biogeographic regions and realms and unevenly threatens protected areas. As an alien's occurrence and risk on a national scale are linked with abiotic factors followed by propagule pressure, their local distribution in protected areas is shaped by propagule pressure and biotic filters. The proposed modelling framework for the assessment of the risk posed by invasive alien plants across spatial scales and under different protection regimes represents an attempt to fill the gap between theory and practice in conservation planning helping to identify scale, site, and species-specific priorities of management, monitoring and control actions. Based on solid theory and on free geographic information, it has great potential for application to wider networks of protected areas in the world and to any invasive alien plant, aiding improved management strategies claimed by the environmental legislation and national and global strategies.
Biodiversity , Ecosystem , Plants , Introduced Species , Species Specificity
Species richness is a fundamental property of biodiversity patterns and is properly expressed by the species-area relationship (SAR), namely the increase in the number of species with the area. Here, we studied and explored the species-area relationship with respect to vascular plant species in Italy and compared vascular plant richness among Italian administrative regions. Concerning the entire vascular flora (native and alien), the best-performing formula is the Arrhenius' Power function: S = c Az. The constants of this function are c = 241.2 and z = 0.281. The best-performing formula concerning just native (c = 245.2 and z = 0.263) and alien (c = 10.1 and z = 0.404) richness is the Power function as well. The floristically richest Italian regions considering the entire flora are Liguria, Friuli Venezia Giulia, and Trentino-Alto Adige, which are also the regions that are richest in alien flora unfortunately. Regions of particular naturalistic interest are Abruzzo, Valle d'Aosta, and Molise, because only these three regions exhibit native floristic richness that is higher than expected, and this is coupled with an alien floristic richness that is lower than expected. On the contrary, four regions (Lombardia, Veneto, Toscana, and Emilia-Romagna) show potentially severe conservation problems due to biological invasions since they experience native floristic richness that is lower than expected, with an alien floristic richness that is higher than expected. This study offers for the first time the 'c' and 'z' constants specifically calibrated at the national level for Italian vascular flora. The availability of such constants allows the calculation of the number of expected species for a given area to be investigated, providing a robust starting hypothesis for floristic studies.
A comparative karyotype analysis of four species of yellow-flowered Eranthis sect. Eranthis, i.e., E. bulgarica, E. cilicica, E. hyemalis, and E. longistipitata from different areas, has been carried out for the first time. All the studied specimens had somatic chromosome number 2n = 16 with basic chromosome number x = 8. Karyotypes of the investigated plants included five pairs of metacentric chromosomes and three pairs of submetacentric/subtelocentric chromosomes. The chromosome sets of the investigated species differ mainly in the ratio of submetacentric/subtelocentric chromosomes, their relative lengths, and arm ratios. A new oligonucleotide probe was developed and tested to detect 45S rDNA clusters. Using this probe and an oligonucleotide probe to 5S rDNA, 45S and 5S rDNA clusters were localized for the first time on chromosomes of E. cilicica, E. hyemalis, and E. longistipitata. Major 45S rDNA clusters were identified on satellite chromosomes in all the species; in E. cilicica, minor clusters were also identified in the terminal regions of one metacentric chromosome pair. The number and distribution of 5S rDNA clusters is more specific. In E. cilicica, two major clusters were identified in the pericentromeric region of a pair of metacentric chromosomes. Two major clusters in the pericentromeric region of a pair of submetacentric chromosomes and two major clusters in the interstitial region of a pair of metacentric chromosomes were observed in E. longistipitata. E. hyemalis has many clusters of different sizes, localized mainly in the pericentromeric regions. Summarizing new data on the karyotype structure of E. sect. Eranthis and previously obtained data on E. sect. Shibateranthis allowed conclusions to be formed about the clear interspecific karyological differences of the genus Eranthis.
In recent years, essential oils (EOs) have received interest due to their antibacterial properties. Accordingly, the present study aimed to investigate the effectiveness of the EOs obtained from seven species of Salvia on three strains of Listeria monocytogenes (two serotyped wild strains and one ATCC strain), a bacterium able to contaminate food products and cause foodborne disease in humans. The Salvia species analysed in the present study were cultivated at the Botanic Garden and Museum of the University of Pisa, and their air-dried aerial parts were subjected to hydrodistillation using a Clevenger apparatus. The obtained EOs were analysed via gas chromatography coupled with mass spectrometry for the evaluation of their chemical composition, and they were tested for their inhibitory and bactericidal activities by means of MIC and MBC. The tested Eos showed promising results, and the best outcomes were reached by S. chamaedryoides EO, showing an MIC of 1:256 and an MBC of 1:64. The predominant compounds of this EO were the sesquiterpenes caryophyllene oxide and ß-caryophyllene, together with the monoterpenes bornyl acetate and borneol. These results suggest that these EOs may possibly be used in the food industry as preservatives of natural origins.
The genus Santolina (Asteraceae, Anthemideae) includes 26 species of aromatic evergreen shrubs endemic to the western Mediterranean Basin. Santolina is widely used as ornamental plant, in xerigardening, and in ethnobotany. The Santolina chamaecyparissus complex, including about half of the known species diversity, has been properly investigated on systematic and taxonomic grounds only recently, and a complete morphometric study is still missing. Here we provide a morphometric characterization and comparison of all the 14 species of this complex, using both univariate and multivariate analyses. Our results suggest that species of this complex can be distinguished using combinations of quantitative and qualitative character-states, mostly related to the leaf morphology. The analysis of S. villosa, a tetraploid/hexaploid Spanish endemic, showed that the two cytotypes cannot be safely identified based on morphology. Coupling this evidence with available phylogenetic information, we conclude that there is no reason to split the two cytotypes of S. villosa in two distinct taxa. An identification key for all the species of the complex is presented.
Senecio squalidus (Asteraceae) currently includes nine subspecies distributed in North and Central Europe and in the Mediterranean basin. Within this taxonomic aggregate, many species have been described, but research on their nomenclatural types is incomplete. A complete nomenclatural survey of 19 names belonging to this taxonomically critical group was carried out. Fourteen lectotypes are here designated. The nomenclatural analysis, complemented by field investigations in the type localities of the taxa described in the Central Mediterranean, allowed us to accept 10 species. Accordingly, we proposed here a new name and a new missing combination at a specific level: S. aknoulensis and S. calabrus.
Three subspecies of Armeria arenaria are reported from Italy, two of which are considered endemic to the Apennines. The taxonomic value of these two taxa (A. arenaria subsp. marginata and A. arenaria subsp. apennina) is unclear. Moreover, the relationships between A. arenaria subsp. praecox and Northern Italian populations-currently ascribed to A. arenaria subsp. arenaria-have never been addressed. Accordingly, we used an integrated taxonomic approach, including morphometry, seed morpho-colorimetry, karyology, molecular systematics (psbA-trnH, trnQ-rps16, trnF-trnL, trnL-rpl32, and ITS region), and comparative niche analysis. According to our results, French-Northern Italian populations are clearly distinct from Apennine populations. In the first group, there is evidence which allows the recognition of A. arenaria s.str. (not occurring in Italy) and A. arenaria subsp. praecox. In the second group, the two putative taxa endemic to the Northern Apennines cannot be separated, so a single subspecies is here recognized: A. arenaria subsp. marginata.
Correlative ecological niche modelling (ENM) is a method widely used to study the geographic distribution of species. In recent decades, it has become a leading approach for evaluating the most likely impacts of changing climate. When used to predict future distributions, ENM applications involve transferring models calibrated with modern environmental data to future conditions, usually derived from Global Climate Models (GCMs). The number of algorithms and software packages available to estimate distributions is quite high. To experimentally assess the effectiveness of correlative ENM temporal projection, we evaluated the transferability of models produced using 12 different algorithms on historical and modern data. In particular, we compared predictions generated using historical data and projected to the modern climate (simulating a "future" condition) with predictions generated using modern distribution and climate data. The models produced with the 12 ENM algorithms were evaluated in geographic (range size and coherence of predictions) and environmental space (Schoener's D index). None of the algorithms shows an overall superior capability to correctly predict future distributions. On the contrary, a few algorithms revealed an inadequate predictive ability. Finally, we provide hints that can be used as guideline to plan further studies based on the adopted general workflow, useful for all studies involving future projections.
Santolina is a plant genus of dwarf aromatic shrubs that includes about 26 species native to the western Mediterranean Basin. In Corsica and Sardinia, two of the main islands of the Mediterranean, Santolina corsica (tetraploid) and S. insularis (hexaploid) are reported. Along with the cultivated pentaploid S. chamaecyparissus, these species form a group of taxa that is hard to distinguish only by morphology. Molecular (using ITS, trnH-psbA, trnL-trnF, trnQ-rps16, rps15-ycf1, psbM-trnD, and trnS-trnG), cypsela morpho-colorimetric, morphometric, and niche similarity analyses were conducted to investigate the diversity of plants belonging to this species group. Our results confute the current taxonomic hypothesis and suggest considering S. corsica and S. insularis as a single species. Moreover, molecular and morphometric results highlight the strong affinity between S. chamaecyparissus and the Santolina populations endemic to Corsica and Sardinia. Finally, the populations from south-western Sardinia, due to their high differentiation in the studied plastid markers and the different climatic niche with respect to all the other populations, could be considered as an evolutionary significant unit.
Existing global regionalization schemes for plants consider the compositional affinities among biotas, but these have not explicitly considered phylogenetic information. Here, we present for the first time, a phytogeographical delineation of the global vascular flora based on species-level evolutionary relationships. We analysed 8737 820 geographical occurrence records for vascular plants together with a time-calibrated phylogeny including 67 269 species. We constructed a global phylogenetic regionalization by estimating species composition and phylogenetic beta diversity among 200 km × 200 km grid cells across the world. We identified de novo 16 phytogeographical units that are deeply split into two clusters: Laurasian and Gondwanan. Our regionalization broadly matches previous schemes, but also highlights the separation of the Gondwanan biota into an Holotropical cluster and an Australian-Neozealandic-Patagonian cluster. In contrast, no clear split among Laurasian and Gondwanan biotas was retrieved when omitting phylogenetic information. The integration of phylogenetic and geographical information provides new insights into the delineation of phytogeographical areas and their historical relationships, enabling the identification of three large, clearly differentiated biotas, here referred to as kingdoms: Holarctic, Holotropical, and Austral. Our results provide further evidence for delineating transition zones and show a clear latitudinal pattern of increasing evolutionary distinctiveness towards the poles.
Biota , Tracheophyta , Australia , Biological Evolution , Phylogeny
Sileneneglecta has been misunderstood and confused with S.nocturna, although several morphological characters (petal shape, calyx indumentum, hairiness of stamen filaments, seed size, seed-coat surface and shape) allow separation of these species. Moreover, S.mutabilis (which has been considered conspecific with S.neglecta) and S.martinolii (an alleged endemic species to south-western Sardinia) are considered here as taxonomic synonyms of S.nocturna and S.neglecta, respectively. These taxonomic conclusions are strongly supported by multivariate morphometric analyses of 21 characters.
The nomenclature of 14 taxa from Central and Southern Europe within the Dianthusvirgineus group is discussed. Dianthusaggericola Jord., D.collivagus Jord., D.consimilis Jord., D.orophilus Jord., D.saxicola Jord., D.juratensis Jord. are here lectotypified by specimens from the Jordan herbarium in LY, while D.godronianus Jord. by a specimen in P. Dianthussubacaulis Vill. is neotypified by a specimen collected on Mont Ventoux (S. France) and housed in MPU. For D.sylvestris Wulfen, a lectotype is here designated and its previous neotypification is discussed. Dianthuscaryophyllusvar.tenuifolius Moris, D.caryophyllusf.minor Moris and D.sylvestrisvar.garganicus Ten. are lectotypified by specimens housed in herbarium Moris (TO) and herbarium Tenore (K). Dianthusvirgineusvar.tergestinus Rchb. is lectotypified by a drawing from the Icones florae Germanicae & Helveticae, while D.contractusvar.evolutus Lojac. is neotypified by a specimen in P. For each taxon the currently accepted name is provided including new synonymies. The type indication is followed by nomenclatural and taxonomic notes, in which the original material found is commented and the reasons for the identification of the types are discussed.
Comparative karyomorphological analyses of six out of the eight white-flowered species of Eranthissect.Shibateranthis have been carried out. All studied specimens of E.byunsanensis, E.lobulata, E.pinnatifida, and E.stellata had a somatic chromosome number 2n = 16 with basic chromosome number x = 8. On the contrary, E.tanhoensis and E.sibirica had a basic chromosome number x = 7. The specimens of E.tanhoensis were diploid with 2n = 14, while the specimens of E.sibirica were polyploid with 2n = 42. Monoploid chromosome sets of the investigated diploid species had 4-5 metacentric chromosomes and 2-4 submetacentric/subtelocentric/acrocentric chromosomes. The highest level of interchromosomal asymmetry, estimated via CVCL, was found in E.byunsanensis and E.pinnatifida. The highest levels of intrachromosomal asymmetry (MCA) and heterogeneity in centromere position (CVCI) were found in E.lobulata and E.byunsanensis, while E.sibirica had the most symmetric karyotype. A multivariate PCoA analysis of basic karyotype parameters (2n, x, THL, CVCL, MCA, and CVCI) highlighted no overlap among species accessions, which was also confirmed by LDA. The average absolute monoploid DNA content (1Cx) of the 23 investigated samples of six Eranthis species varied from 9.26 ± 0.25 pg in E.sibirica to 15.93 ± 0.32 pg in E.stellata. Overall karyological affinity was highlighted between E.lobulata and E.stellata, on one side, and between E.byunsanensis and E.pinnatifida, on the other side. Interestingly, there was no significant correlation between total haploid (monoploid) chromosome length (THL) and 1Cx values in these species.
Digital data concerning the flora of Italy are largely fragmented among different resources hosted on different platforms, and often with different data standards, which are neither connected by a common access point, nor by web services, thus constituting a relevant obstacle to data access and usage. Taxonomic incongruences add a further complication. This paper describes "FlorItaly", an online information system which allows to access and query updated information on the checklist of the flora of Italy, aiming at becoming an aggregator for Italian botanical resources. "FlorItaly" was developed in a collaborative effort by more than 50 taxonomists, with the support of the Italian Botanical Society, and of Project "Dryades" (University of Trieste), to provide a better and reliable organization of botanical knowledge in Italy, as well as a relevant simplification for data retrieval, and a further stimulus towards a more collaborative approach in botanical research.
This paper deals with the typification and taxonomy of five Mediterranean Cytisus species. Cytisus affinis, C. candidus, and C. spinescens nom. illeg., non Sieber ex Spreng. were described from Sicily by Karel Borivoj Presl, Cytisus spinescens was described from Apulia (southern Italy) by Curt Polycarp Joachim Sprengel, and C. villosus was described from southern France by Pierre André Pourret (1788). Lectotypes are here designated for Presl and Sprengel's names. A neotype is designated for C. villosus. The taxonomic revision of these five names confirmed that C. villosus Pourr. (= Cytisus affinis C.Presl) is the name to be used for the species occurring in the large part of the Mediterranean countries. Cytisus spinescens Sieber ex Spreng. (≡ C. candidus C.Presl = C. spinescens C.Presl, nom. illeg.) is the correct name for the amphi-adriatic species occurring in peninsular Italy, and along the NE coast of the Adriatic Sea. This species does not occur in Sicily and reference to this latter region in the protologues of both C. spinescens C.Presl and C. candidus C.Presl is a misinterpretation due, possibly, to exchange of labels.
Chromosome number and genome variation in flowering plants have stimulated growing speculation about the ancestral chromosome number of angiosperms, but estimates so far remain equivocal. We used a probabilistic approach to model haploid chromosome number (n) changes along a phylogeny embracing more than 10 000 taxa, to reconstruct the ancestral chromosome number of the common ancestor of extant angiosperms and the most recent common ancestor for single angiosperm families. Independently, we carried out an analysis of 1C genome size evolution, including over 5000 taxa. Our analyses revealed an ancestral haploid chromosome number for angiosperms of n = 7, a diploid status, and an ancestral 1C of 1.73 pg. For 160 families, inferred ancestral n are provided for the first time. Both descending dysploidy and polyploidy played crucial roles in chromosome number evolution. While descending dysploidy is equally distributed early and late across the phylogeny, polyploidy is detected mainly towards the tips. Similarly, 1C genome size also increases (or decreases) significantly in late-branching lineages. Therefore, no evidence exists of a clear link between ancestral chromosome numbers and ancient polyploidization events, suggesting that further insights are needed to elucidate the organization of genome packaging into chromosomes.
Magnoliopsida , Chromosomes , Evolution, Molecular , Genome Size , Genome, Plant/genetics , Magnoliopsida/genetics , Phylogeny , Polyploidy
