Leaf morphological variation in common yew (Taxus baccata L.) at the species’ southern range margin in North Africa

Chafiaa Hamidouche-Si Mohammed; Tassadit Dib; Fazia Krouchi; Wadi Badri; Arezki Derridj

doi:10.24193/subbbiol.2026.1.06

Authors

Chafiaa Hamidouche-Si Mohammed Laboratory of Ecology, Biotechnilogy and Health, Mouloud Mammeri University, Tizi-Ouzou, Algeria. ✉Corresponding author, E-mail: chafiaa.hamidouche@ummto.dz https://orcid.org/0009-0006-4397-830X
Tassadit Dib Laboratory of Ecology, Biotechnilogy and Health, Mouloud Mammeri University, Tizi-Ouzou, Algeria. https://orcid.org/0000-0003-2136-7795
Fazia Krouchi Laboratory of Ecology, Biotechnilogy and Health, Mouloud Mammeri University, Tizi-Ouzou, Algeria. https://orcid.org/0009-0003-1271-5660
Wadi Badri University Hassan II of Casablanca, Faculty of Sciences Ben M'Sik, Laboratory of Ecology and Environment, Casablanca, P.O. Box 7955, Morocco. https://orcid.org/0000-0003-1529-9474
Arezki Derridj Laboratory of Ecology, Biotechnilogy and Health, Mouloud Mammeri University, Tizi-Ouzou, Algeria. https://orcid.org/0000-0002-9670-4226

DOI:

https://doi.org/10.24193/subbbiol.2026.1.06

Keywords:

Taxus baccata L., leaf morphology, morphometric variability, North Africa

Abstract

The main aim of this study was to provide the first morphometric characterization of Taxus baccata L. leaves in North Africa. Phenotypic variability was investigated in nine natural populations from Algeria and Morocco using 11 quantitative and qualitative traits. analysis of variance (ANOVA) followed by Fisher’s post hoc test revealed significant differences at multiple hierarchical levels, including trees within populations, among populations, regions and mountain ranges for all quantitative variables. In parallel chi-square analysis indicated significant variation in qualitative traits. Hierarchical clustering classified the studied populations into two major clusters for both types of morphological traits. The Chrea population displayed the largest leaves, whereas the Saharan Atlas populations showed the smallest. Principal component analysis (PCA) clearly separated populations and geographic groups in relation to both ecological and leaf morphological factors. Overall, the observed morphological differentiation suggested that phenotypic plasticity together with possible genetic differentiation, is driven more by local ecological conditions than by latitude alone. North African yews are characterized by relatively narow leaves and region specific variation in petiole and apex morphology, reflecting high degree of ecological adaptability to various ecological and geographical conditions. These findings have important implications for conservation of southern marginal population of this relic threatened conifer.

Article history: Received 8 January 2026; Revised 29 April 2026;
Accepted 16 June 2026; Available online 25 June 2026.

References

Abdessemed K. (1981). Le cèdre de l'Atlas dans les massifs de l'Aurès et du Belezma Etude phytosociologique et problèmes de conservation et d'aménagement [in French]. Thèse Doc-Ing., Univ. Aix-Marseille III, 199p.

Angert, A. L. (2006). Growth and leaf physiology of monkey flowers with different altitude ranges. Oecologia, 148(2),183-194. https://doi.org/10.1007/s00442-006-061-z

Barbero, M. & Quézel, P. (1994). Place, rôle et valeur historique des éléments laurifoliés dans les végétations préforestières et forestières ouest-méditerranéennes [in French]. Annal de Botanique Volume, 52. 81-133.

Beghami, Y. (2010). Contribution à l’étude de l’influence des facteurs édaphiques, orographiques et biologiques sur la régénération naturelle du cèdre de l’Atlas (Cedrus atlantica. M) dans les monts des Ouled Yagoub [in French]. These doc. Univ. de biskra, Algerie 288 p

Boeger, M., Alves, L. & Negrelle, R. (2004). Leaf morphology of 89 tree species from a lowland tropical rain forest (Atlantic Forest) in South Brazil. Brazilian Archives of Biology and Technology 47. https://doi.org/10.1590/S1516-89132004000600013

Boudy, P. (1952). Guide forestier en Afrique du Nord. La maison rustique, Paris. 505 p.

Bresson, C. Vitasse, Y. Kremer, A. & Delzon, S. (2013). To what extent is altitudinal variation of functional traits driven by genetic adaptation in European oak and beech? Tree Physiology. 31, 1164-1174. https://doi.org/10.1093/treephys/tpr084

Bruschi, P., Grossoni, P. & Bussotti, F. (2003). Within- and among-tree variation in leaf morphology of Quercus petraea (Matt.) Liebl. natural populations. Trees 17, 164–172. https://doi.org/10.1007/s00468-002-0218-y

Caudullo, G., Welk, E. & San-Miguel-Ayanz, J. (2017). Chorological maps for the main European woody species. Data in Brief, 12, 662-666. https://doi.org/10.1016/j.dib.2017.05.007

Cedro, A. & Iszkuło, G. (2011) Do females differ from males of european yew (taxus baccata l.) in dendrochronological analysis? Tree-Ring Research, 67(1), 3-11 https://doi.org/10.3959/2009-9.1

Cope, S., Corney, D., Clark, Y., Remagnino,P., Wilkin, P. & Debeaux O. (2012); Plant species identification using digital morphometrics: A Review Expert Systems with Applications 39, 7562–7573. https://doi.org/10.1016/j.eswa.2012.01.073

Dempsey, D. & Hook, I. (2000). Yew (Taxus) species - Chemical and morphological variations. Pharmaceutical Biology 38(4), 274-280. https://doi.org/10.1076/1388-0209(200009)3841-aft274

Derridj, A. (1990). Etude des populations de Cedrus atlantica Manetti en Algérie[in French]. Thèse Doc., Univ. Paul Sabatier de Toulouse, 288 p.

Garcia, D., Zamora, R., Hódar, J. A., Gómez, J. M. & Castro, J. (2000). Yew (Taxus baccata L.) regeneration is facilitated by fleshy-fruited shrubs in Mediterranean environments. Biological Conservation, 95, 31-38. https://doi.org/10.1016/S0006-3207(00)00016-1

Hamidouche-Si, M. C., Krouchi, F., Bouamed, A. & Vessella, F. (2014). Geographic distribution and morphological variation of Taxus baccata in Algeria. Der Eibenfreund, 20, 39–53.

Iszkuło, G., Jasińska, A., Giertych, M. & Boratyński, A. (2009). Do secondary sexual dimorphism and female intolerance to drought influence the sex ratio and extinction risk of Taxus baccata? Plant Ecology 200(2), 229-240. https://doi.org/10.1007/s11258-008-9447-5

Iszkuło, G. & BoratyńskI, A. (2006). Analysis of the relationship between photosynthetic photon flux density and natural Taxus baccata seedlings occurrence. Acta Oecologica, 29, 78 84. https://doi.org/10.1016/j.actao.2005.08.001

Iszkuło, G., Didukh, Y., Giertych, M. J., Jasińska, A. K., Sobierajska, K. & Szmyt, J.(2012). Weak competitive ability may explain decline of Taxus baccata. Annals of Forest Science, 69, 705-712. https://doi.org/10.1007/s13595-012-0193-4

Kenzo, T. (2012). Variations in leaf photosynthetic and morphological traits with tree height in various tree species in a Cambodian tropical dry evergreen forest. Japan Agricultural Research Quarterly, 46, 167-180. https://doi.org/10.6090/jarq.46.167

Kostrakiewicz, K. (2009). The influence of shadow created by adjacent plantson phenotypic plasticity of endangered species Trollius europaeus l (Ranunculacae). Polish Journal of Ecology: 57(4), 625-634.

Lapie G. & Maige A. (1914). Flore forestière illustrée, comprenant toutes les espèces ligneuses de l'Algérie et les espèces ligneuses les plus répandues en Tunisie, au Maroc et dans le Midi de la France[in French]. Édition Orlhac E., Paris. 357p.

Laribi, M. (2000). Contribution à l’étude phytosociologique des formations caducifoliées à Quercus canariensis Willd et Quercus afares Pomel du massif forestier d’Ath-Ghobri et d’Akfadou (Kabylie) [in French]. Mémoire de Magister, Universite Mouloud Mammeri, Tizi ouzou, 140p.

Li, T., Shen-Tu, X., Xu, L., Zhang, W., Duan, J., Song, Y. & Dong, M. (2022). Intraspecific and sex-dependent variation of leaf traits along altitude gradient in the endangered dioecious tree Taxus fuana Nan Li & R.R. Mill. Frontiers in Plant Science, 13, 996750. https://doi.org/10.3389/fpls.2022.996750

Linares, J. C. (2013). Shifting limiting factors for population dynamics and conservation status of the endangered English yew (Taxus baccata L., Taxaceae). Forest Ecology and Management, 291, 119-127. https://doi.org/10.1016/j.foreco.2012.11.009

Maire R., (1916).Deuxième contribution à l'étude de la flore du Djurdjura. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord, Tome. 07(2), 49-61.

Mayol,M., Riba, M., Gonzalez-Martınez, S., Bagnoli,F., de Beaulieu, J., Berganzo, E., Burgarella, Dubreuil, M., Krajmerov D., Paule, L., Romsakova, I., Vettori, C., Vincenot, L. & Vendramin, G. (2015). Adapting through glacial cycles: Insights from a long-lived tree (Taxus baccata). New Phytologis, 208, 973–986. https://doi.org/10.1111/nph.13496

Meddour R. (1994). La cédraie de l'Atlas Blidéen (Algérie): Valeur bioclimatique, syntaxonomique et dynamique[in French]. Annals de Recherches Forestieres, Maroc, T. 27(1), 105-27.

Medina, E., Garci, V. & Cuevas, E. (1990). Sclerophylly and oligotrophic environments : relationships between leaf structure, mineral nutrient content, and drought resistance in tropical rain forests of the upper rio negro region. Biotropica, 22(1), 51-64. https://doi.org/10.2307/2388719

Metslaid, M., Jogiste, K., Nikinmaa, E., Moser, W. & Porcar-Castell, A. (2007). Tree variables related to growth response and acclimation of advance regeneration of Norway spruce and other coniferous species after release. Forest Ecology and Management: 250, 56–63. https://doi.org/10.1016/j.foreco.2007.03.009

M'hirit O. & Blerot P. (1999). Le grand livre de la forêt marocaine[in French]. Edition Mardaga, 280 p.

Mitchell, A. K. & Arnott, J.T. (1995). Effects of shade on the morphology and physiology of amabilis fir and western hemlock seedlings. New Forests 10, 79-98. https://doi.org/10.1007/BF00034177

Mitchell, A. K. (1998). Acclimation of Pacific yew (Taxus brevifolia) foliage to sun and shade. Tree Physiology 18, 749-757. https://doi.org/10.1093/treephys/18.11.749

Moller M., Gao, L., Mill, R., Li, D., Hollingsworth, M. & Gibby, M. (2007). Morphometric analysis of the Taxus wallichiana complex (Taxaceae) based on herbarium material. Botanical Journal of the Linnean Society 155(3), 307-335. https://doi.org/10.1111/j.1095-8339.2007.00697.x

Omarova, P. (2018). Age intra population variability of generative bodies of Taxus baccata in foothill of Dagestan. 12, 129-135. https://doi.org/10.17513/use.36916.

Paridari, I. C., Jalali, S. G., Sonboli, A., Zarafshar, M. & Bruschi, P. (2013). Leaf macro-and micro-morphological altitudinal variability of Carpinus betulus in the Hyrcanian forest (Iran). Journal of Forestry Research, 24, 301-307. https://doi.org/10.1007/s11676-013-0353-x

Pasini D. & Mirjalili V. W. (2006). The optimized shape of a leaf petiole transactions on Ecology and the Environment, 87, 35. https://doi.org/10.2495/dn060041

Perrin, P. & Mitchell, F. (2013). Effects of shade on growth, biomass allocation and leaf morphology in European yew (Taxus baccata L.). European Journal of Forest Research, 132, 211-218. https://doi.org/10.1007/s10342-012-0668-8

Pescador, D., Bello, F., Valladares, F. & Escudero, A. (2015). Plant trait variation along an altitudinal gradient in mediterranean high mountain grasslands: controlling the species turnover effect. Plos One 10(3), e0118876. https://doi.org/10.1371/journal.pone.0118876

Pfennigwerth, A., Bailey, J.K. & Schweitzer, J.A. (2017). Trait variation along elevation gradients in a dominant woody shrub is population-specific and driven by plasticity. https://doi.org/10.1093/aobpla/plx027

Read, Q.D., Moorhead, L.C., Swenson,N.G., Bailey,J.K. & Sander, N.J. (2014). Climate change and species range shifts convergent effects of elevation on functional leaf traits within and among species. The Annals of Botany Plants, 9(4), plx027, https://doi.org/10.1111/1365-2435.12162

Robakowski, P., Pers-Kamczyc, E., Ratajczak, E, Thomas P. A., Ye Zi-Piao, Rabska M. & Iszkuło, G. (2018). Photochemistry and antioxidative capacity of female and male Taxus baccata L. acclimated to different nutritional environments. Frontiers in Plant Science, Plant Physiology, 9, 742 https://doi.org/10.3389/fpls.2018.00742

Roche, P., Diaz Burlinson, N. & Gachet, S. (2004). Congruency analysis of species ranking based on leaf traits: Which traits are the more reliable. Plant Ecology, 174, 37-48. https://doi.org/10.1023/b:vege.0000046056.94523.57

Romo, A., Iszkuło, G., Seghir Taleb, M., Walas, Ł. & Boratyński, A. (2017). Taxus baccata in Morocco: a tree in regression in its southern extreme. Dendrobiology, 78, 63-74. https://doi.org/10.12657%2Fdenbio.078.007

Sarmoum, M., Navarro-Cerrilo, R. & Guibal, F. (2019). Bilan actuel et rétrospectif du dépérissement du cèdre de l’Atlas dans le Parc national de Theniet El Had (Algérie). Bois et Forêts des Tropiques, 342. https://doi.org/10.19182/bft2019.342.a31636

Sattarian, A., Akbarian, M., Zarafshar, M., Bruschi, P & Fayyaz, P. (2011). Phenotypic variation and leaf fluctuating asymmetry in natural populations of Parrotia persica (Hamamelidaceae), an endemic species from the Hyrcanian forest (Iran). Acta Botanica Mexicana, 97, 65-81.

Schirone, B., Caetano Ferreira, R., Vessella, F., Schirone, A. Piredda, R. & Cosimosi Simeone, M. (2010). Taxus baccata in the Azores: A relict form at risk of imminent exticinction. Biodiversity and Conservation 19(6), 1547-1565. https://doi.org/10.1007/s10531-010-9786-0

Shah, A., Li, D.Z., Möller,M., Gao,L., Hollingsworth, M.L. & Gibb, M. (2008). Delimitation of Taxus fuana Nan Li & R.R. Mill (Taxaceae) based on morphological and molecular data. Taxon, 57(1), 211-222. https://doi.org/10.2307/25065961

Spjut R. W. (2007 a). A phytogeographical analysis of Taxus (Taxaceae) based on leaf anatomical characters. Journal of the Botanical Research Institute of Texas, 1(1), 291-332.

Spjut R. W. (2007 b). Taxonomy and nomenclature of Taxus (Taxaceae). Journal of the Botanical Research Institute of Texas, 1(1),: 203-289.

Stefanović M., Nikolic, B., Matic, R., Popović, Z., Vidaković, V. & Bojovic, S. (2017). Exploration of sexual dimorphism of Taxus baccata L. needles in natural populations. Trees, 31, 1697–1710. https://doi.org/10.1007/s00468-017

Svenning J.C. & Magard E. (1999). Population ecology and conservation status of the last natural population of English yew Taxus baccata in Denmark. Biological Conservation, 88(2), 173-182. https://doi.org/10.1016/s0006-3207(98)00106-2

Thomas, P. A. & Polwart A. (2003). Taxus baccata L. Journal of Ecology, 91, 489-524. https://doi.org/10.1046/j.1365-2745.2003.00783.x

Tumpa, K., Liber, Z., Šatović, Z., Medak, J., Idžojtić, M., Vidaković, A., Vukelić, J.,Šapić, I., Nikl, P. & Poljak, I. (2022). High level of phenotypic differentiation of common yew (Taxus baccata L.) populations in the north-western part of the Balkan Peninsula. Forests, 13, 78. https://doi.org/10.3390/f13010078

Vessella, F., Simeone, M.C., Fernandes, F.M., Schirone, A., Pires Gomes, M. & Schirone, B. (2013). Morphological and molecular data from Madeira support the persistence of an ancient lineage of Taxus baccata L. in Macaronesia and call for immediate conservation actions. Caryologia, 66, 162–177. https://doi.org/10.1080/00087114.2013.821842

Vivian, M., Robert, C., Godfree, M. J., Colloff, C., Ellery M. & Marshall, D. J.; (2014). Wetland plant growth under contrasting water regimes associated with river regulation and drought: implications for environmental water management. Plant Ecology, 215, 997–1011. http://doi.org/10.1007/s11258-014-0357-4.

Wyka, T., Robakowski, P. & Żytkowiak, R. (2007). Leaf age as a factor in anatomical and physiological acclimative responses of Taxus baccata L. needles to contrasting irradiance environments. Photosynthesis Research, 95(1), 87-99. http://doi.org/10.1007/s11120-007-9238-1

Yahi N. (2007). Les cédraies d’Algérie : phytoécologie, phytosociologie, dynamique et conservation des peuplements. Thèse Doctorat, UniversiteSTHB, Alger, 265p.

Zarek, M. (2007). Variability of morphological features of needles, shoots and seeds of Taxus baccata L. in nature reservations of southern Poland. Electronic Journal of Polish Agricultural Universities, 10. https://doi.org/10.1007/s11270-020-04932-0

Leaf morphological variation in common yew (Taxus baccata L.) at the species’ southern range margin in North Africa

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