Genetic structure and phylogeography of Juniperus phoenicea complex throughout Mediterranean and Macaronesian regions: different stories in one Pedro Sánchez-Gómez, Juan F. Jiménez, Jose Luis Cánovas, Juan Bautista Vera, Isabell Hensen, Miloud Aouissat To cite this version: Pedro Sánchez-Gómez, Juan F. Jiménez, Jose Luis Cánovas, Juan Bautista Vera, Isabell Hensen, et al.. Genetic structure and phylogeography of Juniperus phoenicea complex throughout Mediterranean and Macaronesian regions: different stories in one. Annals of Forest Science, Springer Nature (since 2011)/EDP Science (until 2010), 2018, 75 (3), pp.75. 10.1007/s13595-018-0741-7. hal-02190096 HAL Id: hal-02190096 https://hal.archives-ouvertes.fr/hal-02190096 Submitted on 22 Jul 2019 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Annals of Forest Science (2018) 75: 75 https://doi.org/10.1007/s13595-018-0741-7 RESEARCH PAPER Genetic structure and phylogeography of Juniperus phoenicea complex throughout Mediterranean and Macaronesian regions: different stories in one Pedro Sánchez-Gómez1 & Juan F. Jiménez1,2 & Jose Luis Cánovas1 & Juan Bautista Vera1 & Isabell Hensen3 & Miloud Aouissat4 Received: 22 November 2017 /Accepted: 4 May 2018 /Published online: 18 July 2018 # INRA and Springer-Verlag France SAS, part of Springer Nature 2018 Abstract & Key message The genetic structure of Juniperus phoenicea in the Mediterranean Basin is inferred using amplified fragment length polymorphism markers (AFLP) markers. As other Mediterranean conifers, J. phoenicea populations show moderate levels of genetic diversity and interpopulational differentiation. The pattern of distribution of genetic diversity seems highly influenced by the climatic fluctuations which occurred in the Pleistocene. & Context It has been stated that the genetic structure of Mediterranean conifers is mediated by the historical climatic changes and the geological rearrangements which occurred in the Mediterranean Basin. J. phoenicea provides an excellent example to test how its genetic structure is influenced by these events. & Aims In this work, we study the amount and distribution of genetic diversity of J. phoenicea complex, in order to evaluate its taxonomic status and to reveal underlying phylogeographic patterns. & Methods The molecular diversity was analyzed for 805 individuals from 46 populations throughout its distribution range using AFLP markers. Principal coordinate analysis, analysis of molecular variance (AMOVA), and Bayesian-based analysis were applied to examine the population structure. Handling Editor: Ricardo Alia Contribution of the co-authors J. F. Jiménez has contributed as co- supervisor of the project, running the data analysis and editing the man- uscript and figures. P. Sánchez-Gómez is responsible of designing the work and editing the manuscript. J. L. Cánovas has contributed running the data and editing manuscript, tables and figures. J. B. Vera, I. Hensen and M. Aouissat have contributed designing the work and running several analyses. All authors are also responsible of collecting samples in the field and writing the manuscript. Electronic supplementary material The online version of this article (https://doi.org/10.1007/s13595-018-0741-7) contains supplementary material, which is available to authorized users. * Juan F. Jiménez Miloud Aouissat [email protected] [email protected] Pedro Sánchez-Gómez 1 Departamento de Biología Vegetal (Botánica), Universidad de [email protected] Murcia, Campus de Espinardo s/n, 30100 Murcia, Spain Jose Luis Cánovas 2 [email protected] Departamento de Botánica, Facultad de Ciencias, Universidad de Granada, C/Fuentenueva s/n, 18001 Granada, Spain Juan Bautista Vera 3 [email protected] Institut für Biologie, Martin-Luther Universitat, 06099 Halle, Germany Isabell Hensen [email protected] 4 Centre Universitaire Salhi Ahmed Naama, Naâma, Algeria 75 Page 2 of 12 Annals of Forest Science (2018) 75: 75 & Results AFLP markers revealed moderate levels of intrapopulation genetic diversity, pairwise genetic differentiation, and a clear pattern of isolation by distance. Bayesian analysis of population structure showed five clusters related to the taxonomic status of J. phoenicea and J. turbinata, and a geographic pattern of genetic structure in J. turbinata. & Conclusion All the analysis separate J. phoenicea from J. turbinata.ForJ. turbinata, up to four groups can be distinguished from a phylogeographic point of view. The genetic structure of J. turbinata seems highly influenced by climatic and geologic fluctuations occurring since the Oligocene. Keywords AFLP markers . Biogeography . Cupressaceae . Genetic diversity . Inter-populational differentiation . Taxonomy 1 Introduction 2011). Alternative interpretations define two independent spe- cies, J. phoenicea s. str. and J. turbinata Guss., with J. The Mediterranean Basin is considered a biodiversity “hot turbinata being discussed by several authors (see Farjon spot” (Médail and Quézel 1997), because it contains approx- 2005), and its recognition as a species supported by recent imately 10% of all higher plant species on Earth, despite only molecular studies (Adams 2014; Adams et al. 2013;Adams covering an area equivalent to less than 2% of the world’s land and Schwarzbach 2013). As such, J. phoenicea ≡ J. masses (Fady-Welterlen 2005).Themaincausesofthisspe- phoenicea subsp. phoenicea is generally considered as being cies richness are the great diversity of habitats that it shows endemic to the Iberian Peninsula, south of France and (Blondel and Aronson 1999;Thompson2005) arising from Norhtwestern Italy, while J. turbinata ≡ J. phoenicea subsp. the drastic climatic and geologic changes that have occurred turbinata is widespread throughout the Mediterranean Basin since the Tertiary, which has led to population fragmentation, and Macaronesia. However, Macaronesian populations were the creation of glacial refugia for Tertiary species and subse- formerly described as J. canariensis Guyot, but were later quent speciation, and the creation of a great number of endem- combined as a new subspecies, J. turbinata subsp. canariensis ic species (Médail and Diadema 2009). The diversity of by Rivas-Martínez et al. (1993), with the subspecific range Mediterranean woody species is also very high, representing being widely accepted in recent years. However, its taxonomic three times the number of European temperate tree species status has recently been challenged based on several molecu- (Fady-Welterlen 2005). With this background in mind, a study lar markers (RAPDs, Adams et al. 2006; cpDNA, Adams et al. of the genetic diversity of Mediterranean gymnosperms is of 2010; nrDNA, and cpDNA, Adams et al. 2013), leading to the great interest, because it provides and explanation of the evo- conclusion that J. turbinata subsp. canariensis should not be lutionary strategies followed until now. Unlike conifers in supported at the subspecific level. general, Mediterranean conifers show a much larger span of Although several studies exist about the genetic structure of gene diversity and genetic differentiation (Fady-Welterlen J. phoenicea based on different molecular markers (e.g., 2005). Furthermore, they exhibit some characteristics that pro- Boratyński et al. 2009; Dzialuk et al. 2011; Meloni et al. vide a great opportunity for understanding how geological and 2006), there is no one in which the whole distribution area climate changes might have influenced the evolution of dif- has been sampled. For this purpose, we have chosen amplified ferent lineages, because they are long-lived outcrossing spe- fragment length polymorphism markers (AFLP) (Vos et al. cies that arose mostly in ancient times and dominated the 1995). Since its development, this technique has been applied surface of the Earth before the last palaeoclimate changes that to address questions regarding genetic relatedness among in- occurred in the Mediterranean Basin. dividuals, population structure, phylogenetic relationships, Juniperus grex phoenicea (family Cupressaceae) is a small and mapping of quantitative trait loci (QTL) (Mueller and dioecious or rarely monoecious Mediterranean tree whose dis- Wolfenbarger 1999; Meudt and Clarke 2007) The power of tribution encompasses the Mediterranean Basin and AFLP analysis derives from its ability to quickly generate Macaronesian regions, from Madeira and Canary Islands in large numbers of marker fragments for any organism, without the west to Jordan and Saudi Arabia in the east, although it is prior knowledge of genomic sequence. In addition, AFLP most prevalent throughout the Iberian Peninsula and North requires only small amounts of starting template and it ex- Africa (Browicz and Zieliński 1982; Christensen 1997;do hibits high reproducibility. Nevertheless, it still suffers from Amaral Franco 1986;Farjon2005; Greuter et al. 1984;Jalas several weaknesses, such as its dominant nature, making dif- and Suominen 1973; Quézel and Pesson 1980). The species ficult distinguish between homozygotes and heterozygotes,