Abstract
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.
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References
Bergmann F, Gillet EM (1997) Phylogenetic relationships among Pinus species (Pinaceae) inferred from different numbers of 6PGDH loci. Pl Syst Evol 208:25–34
Critchfield WE, Little EL (1966) Geographic distribution of the Pines of the world. USDA Forest Service 991:1–97
Farjon A (1984) Pines: drawings and descriptions of the genus, 2nd edn. Brill and Backhuys, Leiden (ibid. 2005, 235 p)
Farjon A (2005) A bibliography of conifers: selected literature on taxonomy and related disciplines of the coniferales, 2nd edn. Royal Botanical Gardens, Kew
Gaussen H (1960) Les Gymnospermes actuelles et fossiles. Genre Pinus. Trav Lab Forest Fac Sci Toulouse VI(11):1–272
Gernandt DS, Lopez GG, Garcia SO, Liston A (2005) Phylogeny and classification of Pinus. Taxon 54:29–42
Govindaraju D, Lewis P, Cullis C (1992) Phylogenetic analysis of pines using ribosomal DNA restriction fragment length polymorphisms. Pl Syst Evol 179:141–153
Idrissi-Hassani M (1985) Etude de la variabilité flavonique chez deux conifères mediterranéenne: le pin maritime Pinus pinaster Ait. et le genévrier thurifère Juniperus thunifera L. PhD Thesis, Université Claude Bernard Lyon 1, Lyon
Idrissi-Hassani M, Lebreton P (1992) Les flavonoïdes du Pin maritime: une approche chimiotaxonomique. Forêt méditerr 13:3–8
Jalas J, Suominen J (1973) Atlas Florae Europaeae. Finnish Museum of Natural History, Helsinki
Kaundun SS (1995) Contribution biochimique à la connaissance systématique et génétique du Pin d’alep Pinus halepensis Mill. et du Douglas Pseudotsuga menziesii Mirb. Université Claude Bernard Lyon 1, Lyon
Kaundun SS, Fady B, Lebreton P (1997) Genetic differences between Pinus halepensis, Pinus brutia and Pinus eldarica based on needle flavonoids. Biochem Syst Ecol 25:553–562
Klaus W (1989) Mediterranean pines and their history. Pl Syst Evol 162:133–163
Krupkin AB, Liston A, Strauss SH (1996) Phylogenetic analysis of the hard pines (Pinus subgenus Pinus, Pinaceae) from chloroplast DNA restriction site analysis. Am J Bot 83:489–498
Krüssmann G (1972) Handbuch der Nadelgehölze. Parey, Berlin
Kutil BL, Williams CG (2001) Triplet-repeat microsatellites shared among hard and soft pines. J Heredity 92:327–332
Laracine C (1984) Etude de la variabilité flavonique infraspécifique chez deux Conifères: le Pin sylvestre Pinus sylvestris L. et le genévrier commun Juniperus communis L. PhD Thesis, Université Claude Bernard Lyon 1, Lyon
Laracine-Pittet C, Lebreton P (1988) Flavonoid variability within Pinus sylvestris. Phytochemistry 27:2663–2666
Lauranson-Broyer J (1989) Exploration de la diversité biochimique chez les Conifères: contributions à l’étude de l’hybridation Pinus uncinata Ram. x Pinus sylvestris L., et à la connaissance du complex spécifique Pinus nigra Arn. Université Claude Bernard Lyon 1, Lyon
Lauranson-Broyer J, Lebreton P (1995) Flavonic chemosystematics of the specific complex Pinus nigra Arn. In: Baradat P, Adams WT, Müller-Starck G (eds) Population geenetics and genetic conservation of forest trees. Academic Publishing, Amsterdam, pp 181–188
Lebreton P (1990) La chimiotaxonomie des Gymnospermes. Bull Soc Bot Fr 137:35–46
Lebreton P (1995) Genetics and biodiversity of flavonoids in conifers. In: Baradat P, Adams WT, Müller-Starck G (eds) Population geenetics and genetic conservation of forest trees. Academic Publishing, Amsterdam, pp 41–54
Lebreton P, Laracine-Pittet C, Bayet C, Lauranson J (1990) Variabilité polyphénolique et systématique du pin sylvestre Pinus sylvestris L. Ann Sci For Fr 47:117–130
Liston A, Robinson WA, Pinero D, Alvarez-Buylla ER (1999) Phylogenetics of Pinus (Pinaceae) based on nuclear ribosomal DNA internal transcribed spacer region sequences. Mol Phylogenet Evol 11:95–109
Little EL, Critchfield WB (1969) Subdivisions of the genus Pinus. Misc. Publ. 1144. USDA, Washington DC
Lopez GG, Kamiya K, Harada K (2002) Phylogenetic relationships of Diploxylon pines (subgenus Pinus) based on plastid sequence data. Int J Pl Sci 163:737–747
Mirov NT (1967) The genus Pinus. Ronald Press, New York
Norin T (1972) Some aspects of the chemistry of the order Pinales. Phytochemistry 11:1231–1242
Piovesan G, Pelosi C, Schirone A, Schirone B (1993) Taxonomic evaluations of the genus Pinus (Pinaceae) based on electrophoretic date of salt soluble and insoluble seed storage proteins. Pl Syst Evol 186:57–68
Price RA, Liston A, Strauss SH (1998) Phylogeny and systematics of Pinus. In: Richardson DM (ed) Ecology and biogeography of Pinus. Cambridge University Press, Cambridge. Cited by Gernandt et al. 2005, pp 49–68
Schirone B, Piovesan G, Bellarosa R, Pelosi C (1991) A taxonomic analysis of seed proteins in Pinus spp. (Pinaceae). Pl Syst Evol 178:43–53
Shaw GR (1914) The genus Pinus, vol 5. Arnold Arbor, Boston
Syring J, Willyard A, Cronn R, Liston A (2005) Evolutionary relationships among Pinus (Pinaceae) subsections inferred from multiple low-copy nuclear loci. Am J Bot 92:2086–2100
Wang X-R, Tsumura Y, Yoshimaru H, Nagasaka K, Szmidt AE (1999) Phylogenetic relationships of Eurasian pines (Pinus, Pinaceae) based on chloroplast rbcL, matK, rpl20-rps18 spacer, and trnV intron sequences. Am J Bot 86:1742–1753
Yazdani R, Lebreton P (1991) Inheritance pattern of the flavonic compounds in Scots pine (Pinus sylvestris L.). Silvae Genet 40:57–59
Acknowledgments
We would like to express our gratitude to Mr Paul Berthet and Mr Frédéric Pautz, former and present directors of the Parc de la Tête d’Or botanical garden (Lyon, France), who have given us free access to the plant samples and their institute. Our thanks also go to Mrs Christine Bayet who analysed the flavonoids from Asian and American pines and anonymous reviewers for their critical advice on the manuscript.
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Kaundun, S.S., Lebreton, P. Taxonomy and systematics of the genus Pinus based on morphological, biogeographical and biochemical characters. Plant Syst Evol 284, 1–15 (2010). https://doi.org/10.1007/s00606-009-0228-y
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DOI: https://doi.org/10.1007/s00606-009-0228-y