Technical guidelines for genetic conservation and use and Brutia /

Bruno Fady1, Hacer Semerci2 and Giovanni G. Vendramin3 1 INRA, Mediterranean Forest Research Unit, Avignon, EUFORGEN 2 Forest Seeds and Tree Breeding Research Directorate, Gazi- Ankara, 3 CNR, Institute of Genetics, Firenze, These Technical Guidelines are intended to assist those who cherish the valuable Aleppo and Brutia pine genepools and wish to ensure their sustainability, through conserving important seed sources or use in practical . The focus is on conserving the genetic diversity of the species at the European scale. The recommendations provided in this module should be regarded as a commonly agreed basis to be complemented and further developed in local, national or regional conditions. The Guidelines are based on the available knowledge of the species and on widely accepted methods for the conservation of forest genetic resources.

Biology and

Aleppo pine (Pinus halepensis Mill.) has a pedunculate cone, and fine, flexible, light green nee- dles 5–10 cm long. Brutia pine (Pinus brutia Ten.) has a sessile cone and strong, dark green needles 10–18 cm long. Both species are wind-pollinated and allogamous. Male and female flowers are located on different parts of a tree (monoecy). Both species are extremely prolific seed dispersers and can colo- nize open and disturbed areas easily. Aleppo and Brutia pine forests can grow on all substrates and almost all bioclimates of the Mediterranean region. They can be found at altitudes of 0–600 m in the northern Mediterranean and 0–1400 m in the southern Mediterranean (thermo- and meso-Mediterranean levels). Locally, they can reach higher altitudes, e.g. 2600 m for Pinuseppo and Brutia pinesPinus halepensis halepensis Pinus brutiaAleppo and Brutia pinesPinus halepensis P

P. halepensis in the Higher Atlas Distribution Importance and use of and 1650 m for P. brutia in the Taurus Mountains of Turkey. At the upper limit of Pinus halepensis and P. brutia Aleppo and Brutia pines repre- their distribution, they often con- form a group of related species sent the only or main source of stitute a pre-forest colonizing that can intercross, but occupy wood and forest cover in many stage or are part of a mixed pine- different geographical ranges Mediterranean countries. Eco- oak forest. and bioclimates. nomically, P. brutia is the most Optimal development of Aleppo pine forests cover important species in P. halepensis forests occurs at extensive areas in the western Turkey; P. halepensis is the most annual rainfalls of 350–700 mm Mediterranean: , France, important forest species of North and absolute mean minimum Italy, , , , Africa, and has high ecological temperatures between –2 and Morocco, , , Libya importance in southern France +10°C (semi-arid and sub-humid and . A few natural and arti- and Italy, especially at the urban- bioclimates). Optimal develop- ficial populations can be found in forest interface. Mean productiv- ment of P. brutia forests requires the eastern Mediterranean in ity is approximately 1–2 m3 higher rainfalls but accepts a Turkey, , , and ha-1 year-1 for Aleppo pine, and wider range of temperatures . Total forest cover is 2–3 m3 ha-1 year-1 for Brutia pine. (absolute mean minimum temper- estimated to be approximately Maximum yield can reach atures between –5 and +10°C, 3.5 million hectares. 12–15 m3 for both species. The sub-humid and humid biocli- Brutia pine forests cover wood of these Mediterranean mates). extensive areas in the Eastern pines is used for many purposes: Mediterranean: Greece, Turkey, construction, industry, carpentry, , Syria and Lebanon. A firewood and pulp. Seeds are few small populations can be also used for making pastry. found in and . Other related taxo- nomic groups are present in Ukraine (Crimea, P. stankewiczii Sukaczew), around the Black Sea (, Russian Federation, Ukraine, P. pithyusa Stevenson) and in the Caucasus (, Georgia, Iran, Turkey, P. eldarica Medw.). Total forest cover is estimated to be over 4 million hectares, of which 3.8 million hectares are in Turkey. Pinuss Pinus brutiaAleppo and Brutia pinesbrutiaPinus halepensis Pinus brutia AleppoPinus and Brutia pinesP

Genetic knowledge P. halepensis. The easternmost Threats to P. halepensis provenances tend genetic diversity to have higher juvenile growth. Genetic inventories using bio- Pinus halepensis is better adapt- Aleppo and Brutia pines are not chemical and DNA markers have ed to drought but less adapted considered ecologically threat- demonstrated that genetic diver- to cold than P. brutia. However, ened as a whole. However P. sity is geographically structured. under severe water-stress condi- pityusa is considered vulnerable Most P. halepensis diversity was tions, P. halepensis has an because of population size found in Greek and Spanish pop- increased sensitivity to the fun- reduction linked to habitat ulations although other popula- gus Sphaeropsis sapinea. Both decline (IUCN red list). Pinus tions had lower diversity than species are sensitive to the pine eldarica has a patchy distribution other conifer species. This is procecionnary moth Thaume- and its genetic diversity is lowest consistent with the hypothesis of topoea pityocampa which can among the taxa of this group. a recent expansion of the cause severe defoliation. Pinus such as Matsucoccus species (in the last 10 000 years) halepensis is sensitive to the pine josephii represent a major threat from these two refugial areas, bast scale Matsucoccus josephii, in the Eastern Mediterranean. with colonizing populations although P. brutia is resistant to Thaumatopea pityocampa can establishing by migration of a it. also induce severe defoliation limited number of individuals These, along with ecological throughout the distribution area (founder effect) and/or popula- studies, have been used to of both pines although it does tion dynamics regulated by fire define seed-collection zones and not often lead to mortality. (population bottlenecks). Genetic seed stands in several countries Recently, the canker Crumenu- diversity is higher for P. bru- (e.g. 29 P. halepen- lopsis sororia has started to tia and roughly separates sis seed stands cause severe defolia- Western and Eastern popu- in France). tion and dieback on lations. P. halepensis in Controlled pollination France. The impact of experiments and monitoring forest fires is ambivalent. of sympatric east Mediter- Although they actually pro- ranean populations using mote regeneration, they could be molecular markers have responsible for rare allele demonstrated that unidirectional changes over generations, gene flow is possible from explain the very low diversity P. halepensis to P. brutia result- found in P. halepensis and pro- ing in the emergence of hybrids. mote the spreading of Hybridization is not possible P. halepensis genes into P. brutia using P. brutia as pollen parent. forests. Among-region seed Provenance and laboratory transfers have led to significant tests have revealed significant frost or water-stress damage geographic patterns in adaptive after planting when ill-adapted trait variability. Although both material was used. Reducing species demonstrate polycyclic local population adaptability annual growth patterns, initial through gene flow from planta- shoot units are smaller in tions is also a risk. Finally, sPinus halepensishalepensis Pinus brutiaAleppo and Brutia pinesPinus halepensis PinusPin brutiaAleppo and

because these species (and Guidelines for genetic Population under recurrent especially P. halepensis) are conservation and use forest fires. Because they are highly resistant to drought, they adapted to forest fires, both are often the last forest species Current conservation measures pines usually regenerate well to be found at desert or steppe undertaken at national levels after fire, using the seed bank margins. Global warming and its most commonly include in situ released from serotinous cones. collateral modification of rainfall gene conservation networks If regeneration happens to be regimes may dramatically modify specifically designed for the tar- poor in the first 2 years after fire, their distribution ranges. get species (e.g. in Turkey, 52 and if only a few isolated seed P. brutia conservation units) and remain in the burnt area, forest reserves or national parks artificial regeneration should be which include the target species. used to counteract the risk of Ex situ measures include clonal genetic erosion in the juveniles. archives, cold storage seed In this case, seed lots collected banks and DNA banks. from large genepools should be To increase the efficiency used (e.g. at least 30 trees per of in situ genetic resource population from at least three conservation, a concerted populations from a single seed management effort should zone). be carried out range-wide. Although transfer of seed Populations where hybridiza- material is often legally pos- tion may occur. Planting Aleppo sible, it should be avoided pine where Brutia pine is present across zones and countries should be avoided in areas with different ecological where frost and potential pest requirements, notably damage are limiting factors, or because of cold, drought and strictly monitored in areas where damage risks. drought is the limiting factor. Locally, some populations Owing to the anisotropy of require specific attention and between-species gene flow, the appropriate forestry practice. impact should be reduced when planting Brutia pine in the vicini- Marginal populations. As popu- ty of Aleppo pine forests. lations at high altitudes, in desert margins and mixed forests may contain valuable genes (resist- ance to drought, cold, pests) for adaptation under global warm- ing, efforts such as gene reserves should be made to conserve them. PinusAleppo and Brutia pinesPinus halepensishalepensis Pinus brutiaAleppo and Brutia pinesPinus h

Distribution range of Aleppo pine

Distribution range of Brutia pine shalepensis PinusPinus brutiaAleppo and Brutia pinesbrutiaPinus halepensis Pinus brutiaAleppo Pi and Bruti EUFORGEN

These Technical Guidelines were Selected bibliography produced by members of the EUFORGEN Network. The objective of the Network is to Bariteau, M., R. Huc and G.G. Vendramin (coordinators). 2001 Adaptation and selection of Mediterranean Pinus and Cedrus for sustainable afforestation identify minimum genetic conser- of marginal lands. Final report of EU Project FAIR CT95-0097, 173 p. vation requirements in the long Bucci, G., M. Anzidei, A. Madaghiele and G.G. Vendramin. 1998. Detection of term in Europe, in order to reduce haplotypic variation and natural hybridization in halepensis-complex pine the overall conservation cost and species using chloroplast simple sequence repeat (SSR) markers. Molec- to improve the quality of stan- ular Ecology 7(12):1633-1643. dards in each country. Conkle, M.T., G. Schiller and C. Grunwald. 1988. Electrophoretic analysis of diversity and phylogeny of Pinus brutia and closely related taxa. System- atic Botany 13(3):411-424. Kaundun, S.S., B. Fady and Ph. Lebreton. 1997. Genetic differences between Pinus halepensis, Pinus brutia and Pinus eldarica based on needle flavonoids. Biochemical Systematics and Ecology 25(6):553-562. Citation: Fady, B., H. Semerci and Ne’eman, G. and L. Trabaud (eds.). 2000. Ecology, biogeography and manage- G.G. Vendramin. 2003. EUFOR- ment of Pinus halepensis and P. brutia forest ecosystems in the Mediter- GEN Technical Guidelines for ranean basin. Backhuys Publishers, Leiden, The Netherlands. 407 pages. genetic conservation and use for The distribution map was compiled by members of the EUFORGEN Aleppo pine (Pinus halepensis) Conifers Network based on an earlier map published by W.B.Critchfield & and Brutia pine (Pinus brutia). E.L.Little, Jr. in 1966 (Geographic Distibution, of the Pines of the World, International Plant Genetic USDA Forest Service Misc. Publication, 991 pages). Resources Institute, Rome, Italy. 6 pages.

Drawings: Pinus halepensis, Clau- dio Giordano. © IPGRI, 2003.

ISBN 92-9043-571-2

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