Massimo Bigazzi

Anchusa L. and allied genera (Boraginaceae) in Italy

ABSTRACT A revision of the Italian entities of Anchusa and of the related genera Anchusella, Lycopsis, Cynoglottis, Hormuzakia and Pentaglottis was carried out in view of the poor systematic knowledge of some entities of the national flora. The taxonomic treatment relies on a wide comparative basis, including macro- and micromorphological, karyological, chorological and ecological data. After a general description of some poorly known microcharacters of vegetative and reproductive structures, analytical keys, nomenclatural types, synonymies, descriptions, distribution maps and iconographies are provided for each entity. Pentaglottis (P. sempervirens) and Hormuzakia (H. aggregata) are monotypic genera, while Anchusella, Cynoglottis and Lycopsis each include two species of which only one is native to Italy: A. cretica, C. barrelieri and L. arvensis, respectively. The taxonomic autonomy of these “satellite” genera is supported by morphological, palynological and karyological evidence. In Italy Anchusa includ...

Leaf surface and anatomy in Boraginaceae tribe Boragineae with respect to ecology and taxonomy

Summary The main anatomical and surface characters of the leaves of 54 specific and subspecific taxa belonging to 14 Boraginaceae genera of tribe Boragineae were analysed and compared by means of light and scanning electron microscopy. Characters concerning the vascular system appeared essentially uniform, while stomata size and density, structure of individual trichomes, type of indumentum, leaf and epidermis thickness and palisade arrangement were variable in the examined taxa. This variation was discussed in relation to the morphological and ecological diversification exhibited by the members of the tribe. Most species had typical dorsiventral leaves, but some showed an incipient isobilateral symmetry with a layer of abaxial palisade tissue. Thickness of the lamina and of the outer wall of the adaxial epidermal cells were higher in the xerophytic taxa, especially in the psammophytic species of Anchusa . The mesophytic genera Brunnera, Pentaglottis, Trachystemon and, partly, Symphytum had typical hypostomatous leaves, while the xerophytic and psammophytic species were characterized by increased stomatal density on the adaxial surface. This was higher in the taxa with thicker leaves. Abaxial stomatal density and size were inversely correlated. Mesophytic species had more densely spaced and smaller stomata than the xerophytic species. Finally, size of the abaxial stomata was higher in the taxa showing a thicker outer wall of the adaxial epidermal cells. Seven types of trichomes were recognized, five of which were eglandular and two glandular. Some of these types were systematically useful (exclusive to one or two genera) e.g., the hooked hairs of Symphytum and the lithocysts of Trachystemon and Brunnera . Energy dispersive X-ray microanalysis (EDX) showed that silica and calcium are the most abundant elements in the mineral deposits of the trichome cells. No strict correlation occurred between surface elements, trichome structure, edaphic requirements and systematic position of the 14 examined taxa, although the presence of small amounts of sulphur and phosphorus in Nonea and a remarkable abundance of Potassium in Nonea pulla and Pulmonaria picta could have systematic relevance.

Pollen morphology in theBoragineae (Boraginaceae) in relation to the taxonomy of the tribe

The results of a comparative survey on pollen morphology in theBoragineae by means of light, scanning and transmission electron microscopy are presented and discussed in relation to the taxonomy of the tribe. Multivariate analyses lead to the recognition of 15 pollen morphotypes for which a discriminant key is proposed. The discriminatory characters concern mainly the stereostructural and ultrastructural features of the grains, such as tectum sculpture and aperture morphology, while the number of apertures appears variable within genera and even single species. Seven out of the 12 investigated genera, as currently circumscribed, are matched by palynological data:Anchusella, Borago, Brunnera, Elizaldia, Lithodora, Symphytum andTrachystemon. On account of aperture shape,Lithodora is however closer to theLithospermeae than to theBoragineae. Other genera, and in particularNonea, show a wide variation in tectum ornamentation, shape of grains and number of apertures. Palynological data do not support a broad concept of the genusAnchusa, and point to the autonomy of the ‘satellite’ generaHormuzakia, Gastrocotyle, Phyllocara, Pentaglottis andCynoglottis.

Molecular phylogeny, morphology and taxonomic re-circumscription of the generic complex Nonea/Elizaldia/Pulmonaria/Paraskevia (Boraginaceae-Boragineae)

We performed a molecular phylogenetic analysis of the generic complex Nonea/Elizaldia/Paraskevia/ Pulmonaria (Boraginaceae-Boragineae), using trn L UAA and ITS1 sequences from non-coding plastid and nuclear DNA, respectively. In the strict consensus of most parsimonious trees from combined ITS1/trnL sequences the ingroup forms a monophyletic group with two sister clades of Nonea/Elizaldia and Paraskevia/ Pulmonaria. The eastern Mediterranean Nonea ohtusifolia is sister to the rest of the ingroup and more distant to Nonea taxa than to those of Pulmonaria. Elizaldia is firmly nested in Nonea and forms a clade that includes the southern Mediterranean species N. vesicaria. Nonea as currently circumscribed is therefore paraphyletic. The molecular results are congruent with morphological, karyological and chorological features and suggest a rede-finition of Nonea s.l. based on monophyletic groups. We propose: (i) the institution of the new monotypic genus Melanortocarya gen. nov. for N. ohtusifolia, (ii) the inclusion of Elizaldia within Nonea, and (iii) the transfer of Paraskevia (Nonea) cesatiana to Pulmonaria. Nonea embergeri, previously treated at the infraspecific level, is recognized at species rank and E. calycina subsp. multicolor is placed in synonymy of N. calycina.

Systematics of Nonea (Boraginaceae-Boragineae): new insights from phenetic and cladistic analyses

Nonea, one of the largest but lesser known genera of Boraginaceae tribe Boragineae, has been investigated by means of phenetics and cladistics to analyse morphological diversity and phylogenetic relationships within the group. Cluster analysis of 29 taxa recognizes five groups of annual species placed in one large cluster and two groups of perennial taxa in another cluster. Principal Component Analysis shows that the enigmatic Greek species, Paraskevia cesatiana (= Nonea cesatiana), is morphologically intermediate between Nonea and Pulmonaria, but closer to the latter due to fruit characters. Successive weighting using 42 characters and Pulmonaria as outgroup indicates that Nonea is paraphyletic, because the two species of Elizaldia are nested within the group of annual taxa. The South Mediterranean species N. vesicaria is closer to the North African taxa of Elizaldia than to any other member of Nonea. The perennial species of the N. intermedia group growing in mesic habitats of the PonticCaucasian region and Eastern Anatolia are in basal position, whereas Paraskevia cesatiana is not nested within Nonea. Candolles sections of Nonea are not clearly supported. Nonea sect. Orthocaryum also appears paraphyletic due to the position of N. obtusifolia, a taxon with numerous autapomorphies that are difficult to interprete phylogenetically. Possible trends of character evolution are discussed also in the light of cytogeographical aspects. These aspects indicate x = 10 as the primary base number and either polyploidy or descending dysploidy associated with the adaptive shortening of the life-cycle as major driving forces of species formation

Anchusella, a new genus ofBoraginaceae from the Central-Eastern Mediterranean

The two closely related speciesLycopsis variegata andAnchusa cretica, formerly placed inAnchusa subg.Rivinia, were compared with the type species ofLycopsis andAnchusa, on the basis of a set of macro and microcharacters. The presence of only two fertile stamens as well as other peculiar characters in flower structure, androecium, gynoecium, pollen and fruit, supports the institution of the new genusAnchusella, consisting ofA. variegata andA. cretica. Karyological and eco-chorological aspects are consistent with morphological data in pointing to the autonomy of this genus, which appears characterized by autapomorphic, advanced traits.

Palynological contribution to the systematics of Rindera and the allied genera Paracaryum and Solenanthus (Boraginaceae-Cynoglosseae)

Abstract Bigazzi, M., Nardi, E. & Selvi, F.: Palynological contribution to the systematics of Rindera and the allied genera Paracaryum and Solenanthus (Boraginaceae-Cynoglosseae). — Willdenowia 36 (Special Issue): 37–46. — ISSN 0511-9618;

The Occurrence of Intranuclear Inclusions in the Labiatae, Verbenaceae and Scrophulariaceae

SUMMARYOur electron microscope observations on 106 species belonging to three different families of Angiosperms (Labiatae, Verbenaceae and Scrophulariaceae) showed the occurrence of nuclear inclusions in 39 of them. We can distinguish five different types of nuclear inclusions on the basis of their ultrastructure: a) with tubular structure (T-type): they are typical of the genus Linaria (Scrophulariaceae); b) with lamellar structure (L-type): occurring in the Scrophulariaceae, Labiatae and Verbenaceae and certainly in the Bignoniaceae and Lentibulariaceae as well; c) with fibrillar structure (F-type): they are typical of the Campanulaceae; d) with crystalline structure (C-type): they have been separated into two subtypes, C1 and C2 types, with thick and loose network respectively; e) with amorphous structure (A-type): they mainly occur in some Scrophulariaceae genera. Though nuclear inclusions have been observed in a limited number of Labiatae species (10, out of 65 examined species), they can be of great...

The phylogenetic relationships of Cynoglottis (Boraginaceae- Boragineae) inferred from ITS, 5.8S and trnL sequences

Abstract.A molecular phylogenetic analysis of Cynoglottis was performed to evaluate previous hypotheses based on non-molecular evidence concerning the position of this genus within Boraginaceae tribe Boragineae. ITS-5.8S and trnLUAA sequences from the nuclear and chloroplast non-coding genomes were obtained for four Cynoglottis taxa and selected members of the related genera Anchusa, Anchusella, Gastrocotyle, Brunnera and Pentaglottis. Cynoglottis is monophyletic, but neither trnL nor ITS support a close relationship with Brunnera, unlike previously supposed on morphological grounds. Brunnera is, instead, related to the southwestern European monotypic genus Pentaglottis, with which it forms a basal clade. ITS-5.8S sequences show that Anchusa thessala, a southeastern European annual species of Anchusa subg. Buglossellum, is sister to Cynoglottis and that the two taxa form a clade which also includes the Balkan endemic Gastrocotyle macedonica. Species of Anchusa subg. Anchusa form a separate lineage with high bootstrap support, suggesting that this heterogeneous genus is paraphyletic with respect to Cynoglottis. ITS sequences also discriminate between the Balkan-Apenninic diploid C. barrelieri and the Anatolian tetraploid C. chetikiana, albeit with low support. The molecular results are discussed in the light of karyological, morphological and chorological aspects.

Ultrastructure of nuclear inclusions and the separation ofVerbenaceae andOleaceae (incl.Nyctanthes)

TEM observations were carried out on 40 taxa of the familyVerbenaceae and 35 taxa of the familyOleaceae, in order to ascertain distribution, ultrastructure and morphology of the intranuclear proteinic inclusions in the mesophyll parenchymatic cells. The investigated genera amount to some 25% and 60% respectively of the genera of the two families. Inside theVerbenaceae, lamellar inclusions (L-type) occur in 6 out of 23 investigated genera: they are mostly present inside the tribesCitharexyleae andVerbeneae (both belonging toVerbenoideae), while they are absent in other subfamilies. All of the investigatedOleaceae genera show intranuclear crystalline inclusions (C 1-type) of three different shapes. Among theAsteridae this is a character peculiar toOleaceae. They appear to be a well defined natural group, including the controversial genusNyctanthes.