Taxonomy and systematics of the genus Pinus based on morphological, biogeographical and biochemical characters

For a long time, systematists have subdivided the genus Pinus into Diploxylon and Haploxylon according to morpho-anatomy and the number of needles. Nevertheless, divergent views remain regarding the structure of these two subgenera, mainly at the section and subsection levels. We propose to clarify some of these uncertainties by studying 45 Pinus taxa of different origins. Our results, based on morphometric and biochemical (flavonoids) parameters, complement those obtained from classical anatomical and morphological studies, and also modern macro-molecular markers (proteins, DNA). We confirm the subdivision of the genus into Pinus = Diploxylon versus Strobus = Haploxylon and the further sectioning of the first subgenus into sections Pinus and Trifoliae. Moreover, we specify the different subsections, whereby the contents of the methylated flavonol isorhamnetin coupled with needle morphometry play a significant role (subsections Pinus vs. Pinaster in section Pinus, Australes + Ponderosae vs. Contortae in section Trifoliae). Given that isorhamnetin proceeds from quercetin by the irreversible action of an O-methyl-transferase, this methylated flavonol becomes a dynamic marker in such way that the taxa rich in isorhamnetin can be considered as more “derived = evolved”. In addition, there exists a highly significant negative correlation between methylation index and number of needles. Consequently, the pines from the Holarctic Strobus group (with five needles and low isorhamnetin contents) can be considered as “ancestral”, in reference to a Laurasian origin of the genus. In the subgenus Pinus, the Nearctic group (=section Trifoliae) remains near the ancestral base. On the other hand, the Holarctic subset “densiflorae” is connected to the other members (mainly European) of the polyphyletic subsection Pinus, in particular with series “sylvestres”. Because of their very high contents of isorhamnetin, the Mediterranean pines result from an accentuation of this evolutionary trend (=subsection Pinaster). In fact, the pines growing under hot and dry climates (Mediterranean region) are highly evolved compared to those from cold and/or wet regions (Eurasia and North America but also, to a lesser extent, the south-eastern USA and East Asia). Our dynamic propositions based on plant phenolics data complete those from more modern macromolecular (DNA, proteins) studies.