Fagaceae) En Algérie (Morphological and Taxonomic Analysis of Quercus Faginea (Fagaceae) Complex in Algeria)

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Fagaceae) En Algérie (Morphological and Taxonomic Analysis of Quercus Faginea (Fagaceae) Complex in Algeria) Botany Analyse morphologique et taxonomique du complexe Quercus faginea (Fagaceae) en Algérie (Morphological and taxonomic analysis of Quercus faginea (Fagaceae) complex in Algeria) Journal: Botany Manuscript ID cjb-2020-0075.R2 Manuscript Type: Article Date Submitted by the 10-Aug-2020 Author: Complete List of Authors: Aissi, Abdeldjalil; Institut des sciences vétérinaires et des sciences agronomiques, des sciences agronomiques BEGHAMI, DraftYassine; D�partement dagronomie, universit� de Biskra Lepais, Olivier; INRAE Vela, Errol; University of Montpellier morphometry, indumentum, Q. canariensis Willd, Q. faginea subsp. Keyword: faginea, Q. f. subsp. broteroi Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : © The Author(s) or their Institution(s) Page 1 of 38 Botany Analyse morphologique et taxonomique du complexe Quercus faginea (Fagaceae) en Algérie Morphological and taxonomic analysis of Quercus faginea (Fagaceae) complex in Algeria Abdeldjalil Aissi1,*, Yassine Beghami1, Olivier Lepais2, Errol Véla3 1 LAPAPEZA, Université Batna 1 Hadj Lakhdar, ISVSA, Batna, Algérie. E-mail : [email protected] 2 BIOGECO, INRA, Univ. Bordeaux, 33610 Cestas, France. E-mail : [email protected] 3 AMAP, Université de Montpellier/CIRAD/CNRS/INRA/IRD, Montpellier, France. E-mail : [email protected] *Auteur correspondant : Abdeldjalil Aissi, N.V. Hamla, Batna, Algérie. E-mail: [email protected] Draft © The Author(s) or their Institution(s) Botany Page 2 of 38 Summary The systematics of Quercus faginea Lam. (s.l) (Fagaceae) remains obscure, as it tends to crossbreed with other species and show signs of polymorphism. Four subspecies were identified and currently recognized in the Iberian Peninsula and Morocco. However, the literature to date does not allow the distinction between the taxa present in Algeria. To remedy the lack of required data, morphological and taxonomic analysis of thirteen tree populations has been carried out in the country. In the process, twenty leaves and twenty buds from ten mature trees were collected at each population to produce a more detailed description of 23 particular morphological traits. MCA (multiple correspondence analysis) and AHC (agglomerative hierarchical clustering) were used to analyze the data and to identify homogeneous groups. The results obtained show high intra and inter-population variability, with leaf size diminishing from east to westDraft and from north to south. Though the findings were remarkable, they alone could not be used to differentiate between populations under examination; the traits relating to limb shape and indumentum type on the leaf prove to be more useful. It is clear from this study that the Q. faginea Lam. group in Algeria is represented by two species: Quercus canariensis Willd. and Q. faginea Lam., the latter being represented by two taxa: subsp. faginea and subsp. broteroi. Keywords: morphometry, indumentum, Q. canariensis Willd, Q. faginea subsp. faginea and Q. f. subsp. broteroi. Résumé La systématique de Quercus faginea Lam. (s.l) (Fagaceae) reste encore indéfinie, d’autant plus qu’elle s’hybride avec d’autres espèces et qu’elle est polymorphe. Quatre sous-espèces ont été identifiées et présentement reconnues en péninsule ibérique et au Maroc, or, les données disponibles ne permettent pas de trancher entre les taxons présents en Algérie. Pour remédier au manque de données nécessaires, une analyse morphologique et taxonomique de treize populations du complexe dans le pays a été effectuée. Dans chacune, vingt feuilles et vingt bourgeons sur dix arbres matures ont été prélevés pour © The Author(s) or their Institution(s) Page 3 of 38 Botany une description détaillée de 23 traits morphologiques. L’ACM (analyse des correspondances multiples) et la CAH (classification ascendante hiérarchique) ont été utilisées pour analyser les données et identifier des groupes homogènes. Une variabilité significative intra et inter-populationnelle est remarquable, avec une taille des feuilles qui tend à se diminuer en allant de l’est vers l’ouest et du nord au sud du pays. Cette variabilité ne permet pas, à elle seule, de différentier entre les populations étudiées, en revanche, les traits liés à la forme du limbe et aux types du tomentum sur la feuille s’avèrent plus utiles. Cette étude confirme que le complexe Q. faginea Lam. est représenté en Algérie par deux espèces : Quercus canariensis Willd. et Q. faginea Lam., lui-même représenté par deux taxons, subsp. faginea et subsp. broteroi. Mots clés : Morphométrie, tomentum, Q. canariensis Willd, Q. faginea subsp. faginea et Q. f. subsp. broteroi. Draft 1. Introduction Le genre Quercus L. (Fagaceae) est réputé pour sa complexité et la difficulté qu’ont les taxonomistes à classer les taxons qu’ils observent. Selon Nixon (1993), il comprend environ 500 espèces distribuées dans l’hémisphère nord, dont les origines remontent au début du Tertiaire. Sa richesse taxonomique et sa grande variabilité ont depuis longtemps fasciné les botanistes (cf. Camus 1936-1954) et sont encore aujourd’hui un sujet très controversé. Malgré de très nombreuses descriptions entre le milieu du 18ième et le milieu du 20ième siècle en Europe et en Méditerranée, de nouveaux taxons continuent d’être formellement décrits comme espèces ou sous-espèces, chez les chênes caducifoliés (Mossa et al. 1999) mais aussi sempervirents (Capelo et Costa 2001 ; 2005). Pourtant, de nombreux taxons anciennement décrits sont encore aujourd’hui considérés comme synonymes taxonomiques d’espèces antérieures, alors qu’ils n’ont pas fait l’objet d’un réexamen attentif. Chez les chênes comme dans d’autres genres et familles, les taxons initialement décrits dans les pays du Maghreb, et l’Algérie en particulier, © The Author(s) or their Institution(s) Botany Page 4 of 38 demeurent très peu connus et cela rend difficile la distinction et la comparaison avec les taxons européens. La plupart des espèces de chênes ont une histoire évolutive très complexe, liée à l’existence d’hybridations interspécifiques (Müller 1952) et d’introgressions (Hardin 1975 ; Rushton 1993 ; Van Valen 1976 ; Leroy et al. 2017). Ces phénomènes limitent l’interprétation de représentations strictement dichotomiques obtenues avec l’essentiel des logiciels de reconstruction phylogénétique, en particulier lorsque seul l’ADN chloroplastique est utilisé. L’analyse conjointe des lignées d’ADN nucléaire et des approches génétiques des populations peuvent aussi être plus pertinentes à cette échelle (Manos et al. 1999). Parmi les critères utilisés pour la classification infragénérique du genre Quercus, l’étude morpho- anatomique des organes floraux est la plus Draftutile pour la reconnaissance des sous-genres ou des sections (Maire 1961). Cependant, pour l’approche taxonomique des complexes d’espèces et de leurs sous- espèces, les critères les plus utiles demeurent la description morpho-anatomique et biométrique des feuilles (Cristofolini et Crema 2005). Les glands et leur cupule peuvent être utilisés pour reconnaître, à différents niveaux, des taxons dans le genre, mais aident surtout à délimiter les sections ou les complexes d’espèces. Autour de l’appellation Quercus faginea Lam., on regroupe des arbres ou arbustes à feuillage caduc ou marcescent appartenant à un complexe de taxons endémiques de la région ibéro-maghrébine (Espagne, Portugal, Maroc, Algérie, Tunisie). Au nord comme au sud, les populations appartenant à ce groupe sont d’un intérêt écologique, socioéconomique et patrimonial considérable. Elles dévoilent également un polymorphisme foliaire très remarquable, qui se manifeste sous nombreuses formes dont certaines d’entre elles paraissent des produits d’hybridation avec d’autres espèces (Q. petraea (Matt.) Liebl, Q. lusitanica Lam., Q. pubescens Willd., Q. robur L., Q. ilex L. Q. coccifera L., Q. suber L., Q. afares © The Author(s) or their Institution(s) Page 5 of 38 Botany Pomel., & Q. pyrenaica Willd.), rendant délicate et instable la délimitation taxonomique (Trabut 1892 ; Maire 1961 ; Amaral Franco 1990 ; Bussotti et Grossoni 1998). Les changements au fil du temps de statut taxonomique du complexe (voir Matériel supplémentaire 1) illustre la difficulté de rattacher la variabilité phénotypique inter-individuelle à des populations voire à des sous-ensembles géographiques cohérents, tant dans la péninsule ibérique que dans sa distribution nord-africaine, l’étude de cette dernière restant inachevée. De surcroît, le caractère de classification lié aux dimensions des poils du tomentum sur les feuilles avancé par Amaral Franco (1990), qui distingue entre Q. faginea subsp. faginea (caractérisé par des feuilles de faible dimensions généralement dentées avec un tomentum à poils courts de 122-155µm) et Q. faginea subsp. broteroi (caractérisée par des feuilles larges généralement crénelée-dentées avec un tomentum à poils longs de 175-200 µm) semble être à lui seul insuffisant pour un diagnosticDraft taxonomique autonome, en raison du chevauchement des valeurs de la longueur des poils du tomentum des deux taxons constaté par Tschan et Denk (2012). En complément, Vàzquez et Coombes (2018) reprennent désormais l’usage de Q. faginea subsp. maroccana (Braun-Blanq & Maire) F.M Vàzquez & A. Coombes, et proposent ainsi Q. faginea subsp. oscensis (P. Monts) F.M Vàzquez & A. Coombes. A contrario des deux autres sous-espèces, ces dernières présentent des caractéristiques particulières ; la première étant limitée au seul Maroc (Jahandiez et Maire 1932) (Fig. 1) et ne
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