02344nas a2200157 4500008004100000245016500041210006900206490000700275520168500282653010401967100001502071700001402086700001902100700002002119856004702139 2014 eng d00aNon-monophyly of Buglossoides (Boraginaceae: Lithospermeae): Phylogenetic and morphological evidence for the expansion of Glandora and reappraisal of Aegonychon0 aNonmonophyly of Buglossoides Boraginaceae Lithospermeae Phylogen0 v633 a
The phylogeny of the small Old World genus Buglossoides and its position in tribe Lithospermeae was investigated
using nrDNA and cpDNA sequences and morphology. Maximum parsimony and Bayesian analyses of ITS-5.8S and trnL-trnF
IGS datasets consistently show that this group is close to Glandora and Lithospermum but not monophyletic. Of the seven spe-
cies usually included, two were retrieved in the genus Glandora, i.e., B. goulandrisiorum from northern Greece and B. gastonii
from the western Pyrenees. Based also on morphology and ecology, the placement of these two rare, rupicolous endemics in
Glandora is here advocated and new combinations are made. The rest of Buglossoides includes two early-diverging clades,
one with annual taxa of section Buglossoides and one with the three perennials of section Margarospermum. Morphological,
palynological and ecological data support the separation of these two groups in distinct genera, Buglossoides s.str. and the
old but largely neglected Aegonychon. Within Buglossoides, two main clades correspond to the B. arvensis and B. incrassata
complexes. These show a largely sympatric distribution from the south Mediterranean to central and northern Europe. Com-
bined with their strong phenotypic polymorphism, this causes difficulties in the distinction between taxa of the two clades,
especially without characteristic cotyledons or fruiting material. Molecular and morphological evidence clearly support the
transfer of the west Mediterranean B. arvensis subsp. permixta to the B. incrassata complex.
An annotated English translation of a German early 19th century text including Latin diagnoses is presented with a high-quality scan of the original publication and direct links to the cited pages with taxon and literature citations (including TL-2 entries).
1 aHilger, HH1 aCole, TCH1 aSelvi, Federico uhttp://biodiversitydatajournal.com/articles.php?id=106401269nas a2200133 4500008004100000245009000041210006900131490000700200520083100207100001601038700001401054700002001068856004701088 2012 eng d00aNonea dumanii sp. nov. (Boraginaceae) from the Taurus mountains (south Turkey)0 aiNonea dumaniii sp nov Boraginaceae from the Taurus mountains so0 v303 aThe new species Nonea dumanii, endemic to the mountains of the western Taurus in south Turkey (C4 Antalya), is described
an illustrated based on original collections by the authors. Karyological observations and analysis of ITS1 DNA sequences
showed that the species is probably hexaploid with 2n = 6x = 60 and has phylogenetic affinity to the diploid N. monticola
from the Paphlagonian mountains as well as to the tetraploid N. anchusoides from northwest Iran and southeast Turkey.
From these allopatric species it is morphologically distinct in characters of the indumentum, flower and fruit. Polyploidy
is supported as a major driving force for speciation in Nonea, especially in the group of Anatolian mountain species with
primary base number x =10.
Aim The presence of numerous reliable fossils and the occurrence of many
endemic island species make the Boraginales particularly suitable for integrative
biogeographical studies. In this paper we aim to elucidate the time frame
and events associated with the origin of selected borages endemic to the
Mediterranean climate zone. More specifically, we describe and examine the
alternative palaeo- and neoendemic hypotheses for their origin.
Location Corsica and Sardinia (continental fragment islands) and the Canary
Islands (an oceanic island archipelago).
Methods Eighty-nine accessions, representing 30 genera from five families
ascribed to the Boraginales, were examined for six chloroplast DNA regions. We
used an integrative approach including phylogenetic analyses (MrBayes),
Bayesian molecular dating (T3 package) with four fossil constraints on nodes,
and biogeographical reconstructions (diva) to elucidate the temporal and spatial
origins of the Corso-Sardinian and Canary Island endemics.
Results Species of Echium endemic to the Canary Islands diverged from their
continental sister clade during the Miocene (15.3 ± 5.4 Ma), probably after the
rise of the oldest islands (c. 20 Ma). Corso-Sardinian endemics of Borago
diverged from their primarily North African sister clade during the late
Miocene-Pliocene (c. 6.9 ± 3.6 Ma), well after the initial fragmentation of the
islands (c. 30 Ma). Similarly, Corso-Sardinian endemics of Anchusa diverged
from the South African Anchusa capensis during the Pliocene–Pleistocene (c. 2.7 ± 2.1 Ma).
Main conclusions The present study reveals an Anatolian origin for Anchusa,
Borago and Echium and underlines the importance of the Eastern Mediterranean
region as a possible reservoir for plant evolution in the Mediterranean Basin. For
Anchusa and Borago, the divergence from their respective sister clades on the two
types of islands post-dated the formation of the islands, thus supporting the neo-
endemic hypothesis, whereas the dating results for the origin of Echium endemics
were less conclusive.
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.
Fruit morpho-anatomy and DNA sequence diversity in Euro-Mediterranean taxa of Cynoglossum and the closely
related genera Solenanthus and Pardoglossum were analysed to assess the structural traits that promote dispersal through
transport via the fur of mammals, and to evaluate the phylogenetic value of carpological variation in the group. Electron and
light microscopy showed striking epizoochorous adaptations in characters of the pluricellular projections of the epicarp (glo-
chids), such as multiple apical hooks, conical shape, and finely tuberculate surface, as well as the heavy mineralization of the
cell walls with especially silicon and calcium revealed by X-ray microanalysis. The attachment potential of nutlets to sheep
fleece, estimated through a General Linear Model, was species-specific and relatively high in the taxa with small and light
diaspores (> 60%). ITS sequences from 29 specific and infraspecific accessions were poorly variable, with pairwise genetic
distances ranging from 0.002 to 0.097 (mean 0.044). A comparative analysis of ITS sequence diversity in relation to the differ-
ent dispersal strategies in the four main Boraginaceae tribes revealed substantially higher levels of variation and interspecific
genetic distances in the non-epizoochorous groups, including the Cynoglosseae genera Myosotis and Omphalodes. Bayesian and
maximum parsimony tree construction suggested paraphyly of Cynoglossum due to the nested position of Solenanthus apen-
ninus and Pardoglossum. Species-level relationships remained largely unresolved, preventing an estimation of the phylogenetic
significance of fruit characters. Rapid spread across the Mediterranean region via epizoochory by wild mammals is possibly
the primary cause for the lack of genetic divergence among species. During the Holocene, nomadic and transhumant pasture
by domesticated herbivores has likely contributed to further spreading and mixing of previously isolated taxa, possibly causing
events of homoploid hybridization and introgression. More variable molecular markers should be tested to better understand
the impact of these processes in the evolutionary history of Cynoglossum and to solve relationships within Cynoglosseae.
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.
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.