@article {122, title = {Historical biogeography of Boraginales: West Gondwanan vicariance followed by long-distance dispersal?}, journal = {Journal of Biogeography}, volume = {44}, year = {2017}, pages = {158{\textendash}169}, abstract = {

Aim To examine the historical biogeography of the Boraginales using molecular dating and ancestral area reconstruction. Location World-wide. Methods We constructed data sets that included all major clades of Boraginales and all orders of asterids using previously published sequences of four plastid markers (trnL-trnF, rps16, ndhF, rbcL). We estimated divergence times using a Bayesian uncorrelated, lognormal relaxed clock approach with four different fossil calibration schemes. Ancestral areas were reconstructed using maximum likelihood methods (Dispersal-Extinction-Cladogenesis). Results Boraginales originated during the Early to Late Cretaceous and started its diversification in the Late Cretaceous. The inferred ancestral area of Boraginales includes the Americas and Africa. The two major clades of Boraginales diversified during the Early Paleogene from African and American ancestors respectively. Early branching families in both clades (Codonaceae and Wellstediaceae in one clade and Hydrophyllacee and Namaceae in the other) may have remained restricted to their areas of origin. The other families started diversifying in several regions of the world during the Eocene (Boraginaceae s.str., Heliotropiaceae, Ehretiaceae) or later (Cordiaceae). Main conclusions Molecular dating and ancestral area reconstruction may be broadly consistent with the idea of a vicariant origin of the two major clades of Boraginales after the break-up of West Gondwana, followed by several independent trans-oceanic dispersal events into most areas of the world. However, uncertainty in both divergence times and ancestral area reconstruction do not rule out the possibility of an origin involving long-distance dispersal.

}, keywords = {Ancestral area reconstruction, Boraginaceae, historical biogeography, Molecular dating, vicariance, West Gondwana}, issn = {1365-2699}, doi = {10.1111/jbi.12841}, url = {http://onlinelibrary.wiley.com/doi/10.1111/jbi.12841/abstract}, author = {Luebert, Federico and Couvreur, Thomas L. P. and Marc Gottschling and Hilger, Hartmut H. and Miller, James S. and M. Weigend} } @article {hilger_pontechium:_2000, title = {Pontechium: a new genus distinct from Echium and Lobostemon (Boraginaceae)}, journal = {Taxon}, volume = {49}, number = {4}, year = {2000}, pages = {737{\textendash}746}, abstract = {

Plastid (trnL(UAA) intron, trnL(UAA)-trnF(GAA) spacer) and nuclear (ITS1) DNA sequence analyses do not only confirm the separate position of \„Echium maculatum L.\" which this species holds within the genus Echium due to the presence of a bicapitate rather than a bifid stigma, but necessitate the exclusion of the taxon from the genus Echium (Boraginaceae) and the description of a new genus, Pontechium.

}, keywords = {{DNA}, Boraginaceae, Echiostachys, Echium, Lithospermeae, Lobostemon, Pontechium gen. nov.}, author = {Hilger, HH and B{\"o}hle, UR} } @article {35, title = {Phylogenetic relationships of the monotypic genera Halacsya and Paramoltkia and the origins of serpentine adaptation in circum- mediterranean Lithospermeae (Boraginaceae): insights from ITS and matK DNA sequences}, journal = {Taxon}, volume = {58 }, year = {2009}, chapter = {700}, abstract = {

Halacsya and Paramoltkia are monotypic and partially sympatric genera in the Balkans, with no clear relation-
ships among extant Lithospermeae due to striking morphological autapomorphies and scarcity of phylogenetic
analyses in this group. The two species H. sendtneri and P. doerfleri show a strict selectivity for serpentine soils,
posing the question whether this edaphic specialization reflects a common ancestry or a parallel process of adap-
tive evolution in unrelated lineages. DNA sequences from the nuclear ITS and chloroplast matK regions were
generated from multiple accessions of Halacsya and Paramoltkia, and from representatives of 16 other genera
of Mediterranean Lithospermeae. SEM analyses of pollen morphology were also conducted to test relationships
indicated by molecular phylogenies. Parsimony analyses retrieved a clade of morphologically well differentiated
monotypic Lithospermeae including Halacsya and Paramoltkia. ITS provided a better resolution of relationships
and showed the two genera to be sistergroups close to Mairetis and Moltkiopsis, and no affinity to Moltkia as sup-
posed by past authors. Pollen characters corroborated the phylogenetic link between the two Balkan genera. Five
further monophyletic clades were recognised: Onosma-Echium, Moltkia, Lithospermum s.l., Arnebia-Macrotomia,
and Alkanna-Podonosma. Mapping the edaphic preferences of Lithospermeae onto molecular cladograms showed
that serpentinophytism as an obligate condition originated separately in the clade of monotypic genera and in
that of Onosma-Echium. In Halacsya and Paramoltkia it represents an early ecological synapomorphy which
probably originated in situ from non-serpentine ancestors related to Moltkiopsis and Mairetis.

}, keywords = {Boraginaceae, edaphic habitat, Halacsya, molecular phylogeny, Paramoltkia, serpentine adaptation}, url = { http://www.jstor.org/stable/27756939}, author = {L. Cecchi and Federico Selvi} } @article {33, title = {A synopsis of the genus Cynoglossum (Boraginaceae-Cynoglosseae) in Italy}, journal = {Plant Biosystems}, volume = {146}, year = {2012}, chapter = {461}, abstract = {

Based on field and herbarium investigations, a systematic synthesis of the critical genus Cynoglossum (Boraginaceae) in Italy is
provided. Types, diagnostic characters and original iconographies are given for each native species, together with a revised
analytical key. Regional distributions are outlined and a list of selected vouchers is included. Nine species are native to the
Italian territory: Cynoglossum barbaricinum, Cynoglossum cheirifolium, Cynoglossum clandestinum, C. columnae, C. creticum, C.
magellense, C. montanum, C. nebrodense and C. officinale. A new subspecies, named C. nebrodense ssp. lucanum, is described
from some calcareous massifs in S Basilicata and N Calabria. This taxon is morphologically and geographically distinct from
the typical C. nebrodense s.str. In addition, C. dioscoridis and C. germanicum, often reported from Italy, cannot be confirmed
and are apparently to be excluded from the Italian flora. Old herbarium records of C. amabile Stapf \& Drummond and C.
coelestinum Lindb. [: Paracaryopsis coelestina (Lindl.) R.R. Mill] from northeast Italy are reported for the first time, although
the presence of these two exotic species in Italy was probably occasional.

}, keywords = {Boraginaceae, Cynoglossum, Italian flora, karyology, systematics, Taxonomy}, doi = {10.1080/11263504.2012.667842}, author = {Federico Selvi and K. Sutor{\'y}} } @article {32, title = {Phylogeny, karyotype evolution and taxonomy of Cerinthe L. (Boraginaceae)}, journal = {Taxon}, volume = {58}, year = {2009}, chapter = {1307}, abstract = {

A phylogenetic and karyological analysis of the small and poorly known genus Cerinthe L. (Boraginaceae-
Lithospermeae) was performed using ITS sequences and standard chromosomal techniques. All taxa are
diploid with 2n = 16 or 2n = 18 and show a variable degree of infraspecific variation, in particular in the
polymorphic C. major and C. minor. Change in base number is associated with an early split between the
two well-supported clades of C. major, corresponding to Cerinthe sect. Cerinthe, and that of all other taxa
belonging to C. sect. Ceranthe, with the base x = 8 found only in the strictly annual C. major group, and
x = 9 in the other five species of the genus: C. minor, C. glabra, C. tenuiflora, C. retorta and C. palaestina.
The latter section is subdivided into the E Mediterranean, annual lineage of C. palaestina\–C. retorta and
the mainly continental, perennial group of C. minor\–C. glabra, the sister of which is the Corsican endemic
C. tenuiflora. The hypothesis that x = 9 represents the primary haploid number and x = 8 is derived through
descending aneuploidy, is discussed. A taxonomic revision of the genus is provided and the following formal
taxonomic changes are proposed: C. major L. subsp. oranensis (Batt.) Selvi \& Cecchi, stat. nov.; C. major
L. subsp. purpurascens (Boiss.) Selvi \& Cecchi, stat. nov.; C. minor L. subsp. cleiostoma (Boiss. \& Sprun.)
Selvi \& Cecchi, stat. nov. Cerinthe tenuiflora, C. retorta and the poorly known C. palaestina are well-defined
species with little internal variation.

}, keywords = {Boraginaceae, Cerinthe, chromosome evolution, molecular phylogeny, systematics, Taxonomy}, url = {http://www.jstor.org/stable/27757019}, author = {Federico Selvi and L. Cecchi and A. Coppi} } @article {29, title = {Flower morphology, nectar traits and pollinators of Cerinthe major (Boraginaceae-Lithospermeae)}, volume = {207}, year = {2012}, pages = {186{\textendash}196}, abstract = {

In April and May 2010, a natural population of Cerinthe major (Boraginaceae-Lithospermeae) was investigated with regard to: floral morphology, phenology, sexual receptivity during anthesis, the production of nectar and its sugar and amino acid content. The pollinator array of this species was also investigated. Cerinthe major has showy, pentamerous, hermaphroditic flowers with a deep yellow corolla tube arranged on numerous scorpioid inflorescences. Nectar and pollen are the main floral rewards for pollinators. The arrangement of stamens limits access to nectar and promotes pollen loading onto pollinator bodies as they force their way towards the source of nectar. By limiting the direct exposure of nectar to the environment, the corolla tube and the arrangement of anther filaments seemingly protect nectar from evaporation resulting from high temperatures and low relative humidity during flower anthesis. The green, annular nectary located at the base of the ovary lobes is composed of a very thin epidermis enclosing a dense mass of parenchyma. The epidermis lacks stomata, and the thin cuticle has an irregular surface. The parenchyma cells do not store starch during the pre-secretory stage, suggesting that immediate photosynthesis is the most likely source of nectar carbohydrates. Generally, anthesis lasts 4\–5 days; the gynoecium becomes fully receptive on the second day and this is synchronous with anther dehiscence. Since nectar production begins during the bud stage, a substantial volume is available for flower visitors at anthesis. Nectar production decreases drastically in senescent flowers and, if not consumed, the nectar can be re-absorbed. Analysis of sugars revealed a sucrose-dominant nectar (sucrose = 93.18 \± 1.35\%). Proline, GABA, taurine, leucine, citrulline and alanine were the main amino acids present and are probably implicated in pollinator flight performance. The bees Bombus hortorum, Anthophora sp. and Eucera sp. were constant and frequent visitors of C. major flowers. Moreover, the manner in which they interacted with the flower in order to obtain nectar and pollen loads was sufficient evidence for their role as pollinators. Their different foraging behaviours suggest that each species may promote differences in pollen flow both within and among individuals of the population.

}, keywords = {Boraginaceae, Cerinthe major, nectar, pollinators}, doi = {10.1016/j.flora.2012.01.004}, author = {Nocentini, D and Pacini, E and Guarnieri, M and Nepi, M} } @article {9, title = {Multiple origins for Hound{\textquoteright}s tongues (Cynoglossum L.) and Navel seeds (Omphalodes Mill.) {\textendash} The phylogeny of the borage family (Boraginaceae s.str.)}, journal = {Molecular Phylogenetics and Evolution}, volume = {68}, year = {2013}, month = {2013/09//}, pages = {604 - 618}, abstract = {AbstractRecent studies all indicated that both the affinities and subdivision of Boraginaceae s.str. are unsatisfactorily resolved. Major open issues are the placement and affinities of Boraginaceae s.str. in Boraginales and the major clades of the family, with especially the large tribes Cynoglosseae and Eritrichieae repeatedly retrieved as non-monophyletic groups, and the doubtful monophyly of several larger genera, especially Cynoglossum and Omphalodes. The present study addresses and solves these questions using two plastid markers (trnL{\textendash}trnF, rps16) on the basis of a sampling including 16 outgroup taxa and 172 ingroup species from 65 genera. The phylogeny shows high statistical support for most nodes on the backbone and on the individual clades. Boraginaceae s.str. are sister to African Wellstediaceae, Wellstediaceae{\textendash}Boraginaceae s.str. is sister to African Codonaceae. Echiochileae are retrieved as sister to the remainder of Boraginaceae s.str., which, in turn, fall into two major clades, the Boragineae{\textendash}Lithospermeae (in a well-supported sister relationship) and the Cynoglosseae s.l. (including Eritrichieae). Cynoglosseae s.l. is highly resolved, with Trichodesmeae (incl. Microcaryum, Lasiocaryum) as sister to the remainder of the group. Eritrichieae s.str. (Eritrichium, Hackelia, Lappula) are resolved on a poorly supported polytomy together with the Omphalodes-clade (incl. Myosotidium, Cynoglossum p.p.), and the Mertensia-clade (incl. O. scorpioides, Asperugo). The Myosotideae (Myosotis, Trigonotis, Pseudomertensia) are retrieved in a well-supported sister-relationship to the core-Cynoglosseae, the latter comprising all other genera sampled. Cynoglossum is retrieved as highly para- and polyphyletic, with a large range of generic segregates embedded in Cynoglossum, but other species of Cynoglossum are sister to Microula or to the American {\textquotedblleft}Eritrichieae{\textquotedblright} (Cryptantha and allied genera). Representatives of the genus Cynoglossum in its current definition are segregated onto six independent lineages, members of Omphalodes onto three independent lineages. At least 11 of the genera here sampled are deeply nested in other genera. The data show that individual details of nutlet morphology (e.g., winged margins, glochidia) are highly homoplasious. Conversely, a complex of nutlet characters (e.g., characters of the gynobase and cicatrix together with nutlet orientation and sculpturing) tends to circumscribe natural units. Geographical distribution of major clades suggests that the family originated in Africa and western Asia and radiated to eastern Eurasia, with several independent dispersal events into Australia and the New World. }, keywords = {Boraginaceae, Nutlet morphology, Phytogeography}, isbn = {1055-7903}, url = {http://www.sciencedirect.com/science/article/pii/S1055790313001590}, author = {M. Weigend and Federico Luebert and Federico Selvi and Grischa Brokamp and HH Hilger} }