@article {36, title = {Origin of Mediterranean insular endemics in the Boraginales: integrative evidence from molecular dating and ancestral area reconstruction}, journal = {Journal of Biogeography}, volume = {36}, year = {2009}, chapter = {1282}, abstract = {

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.



}, keywords = {Anchusa, Borago, Canary Islands, continental fragment island, Corsica, Echium, endemism, oceanic island, Sardinia.}, doi = {10.1111/j.1365-2699.2009.02082.x}, author = {G. Mansion and Federico Selvi and A. Guggisberg and E. Conti} } @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 {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} }