Technical guidelines for genetic conservation and use European black pine Pinus nigra V. Isajev1, B. Fady2, H. Semerci3 and V. Andonovski4 1 Forestry Faculty of Belgrade University, Belgrade, Serbia and Montenegro EUFORGEN 2 INRA, Mediterranean Forest Research Unit, Avignon, France 3 Forest Tree Seeds&Tree Breeding Research Directorate, Ankara,Turkey 4 Faculty of Forestry, Skopje, Macedonia FYR These Technical Guidelines are intended to assist those who cherish the valuable European black pine genepool and its inheritance, through conserving valuable seed sources or use in practical forestry. 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 ecology although seed yield is abundant only every 2–4 years. Trees reach sexual maturity at 15–20 years in The European black pine (Pinus their natural habitat. Flowers nigra Arnold) grows up to 30 appear in May. Female inflores- (rarely 40–50) m tall, with a trunk cences are reddish, and male that is usually straight. The bark catkins are yellow. Fecundation is light grey to dark grey-brown, occurs 13 months after pollina- deeply furrowed longitudinally on tion. Cones are sessile and hori- older trees. The crown is broadly zontally spreading, 4–8 cm long, conical on young trees, umbrel- 2–4 cm wide, yellow-brown or la-shaped on older trees, light yellow and glossy. They especially in shallow soil ripen from September to Octo- on rocky terrain. ber of the second year, and open Branch tips are in the third year after pollination. slightly ascending Cones contain 30–40 seeds, of on young trees; on which half can germinate. Seeds older trees only are grey, 5–7 mm long, with a branches at the top wing 19–26 mm long. Germina- part of the crown tion can occur without stratifica- have upturned tips. tion although this technique is Needles are rather often used in forest nurseries stiff, 8–16 cm long, (30–60 day moist +5°C treat- 1–2 mm in diameter, ment). Black pine is an obligate straight or curved, finely seeder under natural conditions. serrated. Resin ducts are Most black pine subspecies median. Leaf sheath is persist- (see Distribution) grow in a ent, 10–12 mm long. Mediterranean-type climate, Black pine is a monoecious except P.n. nigra which is more wind-pollinated conifer, and its typically temperate. Bioclimatic seeds are wind dispersed. conditions range from humid Flowering occurs every year, (800–1000 mm annual rainfall) as Pinusuropean black pinePinus nigranigraEuropean black pine PinusPinus nigraEuropean black pine P

in P.n. mauretanica or P.n. laricio, Distribution northern Greece through the to subhumid (600–800 mm) as in Julian Alps and the Balkan P.n. pallasiana in Cyprus, to mountains, covering more than semi-arid (400–600 mm) as Black pine extends over more 800 000 ha. in P.n. pallasiana in Ana- than 3.5 million hectares from Pinus nigra dalmatica (Vis.) tolia. western North Africa through Franco, the Dalmatian pine, is The optimal altitudinal southern Europe to Asia found on a few islands off the range of black pine is Minor. Owing to this large coast of Croatia and on the between 800 to 1500 m. albeit discontinuous range southern slopes of the Dinaric However, a considerable alti- and its large genetic and Alps. tudinal variation can be phenotypic variability, it Pinus nigra pallasiana (Lamb.) observed: from 350 to 1000 m is regarded as a collec- Holmboe covers extensive areas, in Italy (P.n. nigra) and on the tive species. Although mostly in Greece and Turkey (2.5 Croatian coast (P.n. dalmatica), there is no definite con- million ha, 8% of total forest from 500 to 900 m in the French sensus on its taxonomy, area) and possibly as far west as Pyrenees and 1600 to 2000 m in six main subspecies can Bulgaria. It can also be found in Spain (P.n. salzmannii), from be recognized from North Africa Cyprus and the Crimea. It is 1000 to 1600 m in Corsica (P.n. to the Crimea. sometimes referred to as the laricio), from 1000 to 2200 m in Pinus nigra mauretanica Crimean pine. the Taurus mountains and 1400 (Maire et Peyerimh.) Heywood to 1800 m in Cyprus (P.n. pal- covers only a few hectares in the lasiana) and from 1600 to 1800 m Rif mountains of Morocco and the in North Africa (P.n. mauretanica). Djurdjura mountains of Algeria. Black pine can grow on a Pinus nigra salzmannii (Dunal) variety of substrates: limestone Franco (syn: P. n. clusiana, P. n. (e.g. P.n. mauretanica, P.n. dal- pyrenaica) covers extensive matica, P.n. pallasiana in Central areas in Spain (over 350 000 ha Greece), dolomites (e.g. P.n. from Andalucia to Catalunia and nigra in northern Italy and Aus- on the southern slopes of the tria, P.n. salzmanni in the Pyrenees) and is found in a few Cévennes, France), acidic soils isolated populations in the Py- (P. n. laricio, P.n. pallasiana in renees and Cévennes in France. Anatolia, P. n. salzmanni in the It is sometimes referred to as the French Pyrenees) or volcanic Pyrenean pine. soils (P.n. laricio in Sicily). Pinus nigra laricio (Poiret) is Black pine is a light-demand- found in Corsica (Corsican pine) ing species, intolerant of shade over 22 000 ha, in Calabria but resistant to wind and (where it is also recognized as P. drought. It grows in pure stands n. l. calabrica, the Calabrian pine) or more rarely in association with and in Sicily. other pines such as P. sylvestris Pinus nigra nigra (syn: P.n. or P. uncinata. austriaca Höss, P.n. nigricans Host, the Austrian pine) is found from Italy in the Apennines to Pinusnigra nigraEuropean black pinePinusPinus nigraEuropean black pinenigraPinus nigraEuropean b

Importance and use species in southern England as Genetic knowledge well as in some French regions (e.g. Loire valley). Black pine is one of the most Black pine is also valued for The first black-pine-type fossils economically important native landscaping, both in parks (iso- date to the Miocene, about 20 conifers in southern Europe. Ear- lated trees or in groups) and in million years ago. The ice cycles ly growth is rather fast. It is wide- urban and industrial contexts that shaped the Quaternary peri- ly planted outside its natural because of its tolerance to pol- od in Europe are believed to have range. Wood is durable and rich lution. It is one of the most been responsible for the currently in resin, easy to process. common introduced orna- very discontinuous range of black P.n. laricio is appreci- mentals in the USA. Other uses pine. This geographic separation ated for building and include Christmas trees, fuel did not result in mating barriers, roofing because of its wood and poles. and all subspecies are interfertile straightness and thin branches. Black pine is included in under experimental conditions. If properly thinned, its low the European Council Direc- Studies using morphological and amount of duramen makes it a tive 1999/105/CE (Decem- genetic markers have confirmed fine carpentry and cabinetry ber 22 1999) on the mar- the common phylogenetic origin wood. The same use can be keting of forest reproduc- of all black pines. The most diver- made of Calabrian pine, although tive material. Minimum gent and genetically original Euro- it is more branchy. Wood of P.n. requirements have to pean groups are P.n. salzmanii nigra is of lower quality and thus be met before black and P. n. laricio, although P.n. restricted to lower-grade building pine seed can be nigra, dalmatica and pallasiana wood and the making of crates. sold for reforesta- appear quite similar. The amount Black pine has a mean produc- tion. of genetic diversity is also high tivity of 8–20 m3 ha-1 yr-1 when within populations. Experiments grown as a monoculture on measuring adaptive traits have fertile soils. In natural revealed strong within- and conditions, productivity among-population variability for is 6–10 m3 ha-1 yr-1 and traits such as vigour, form and down to less then 3 m3 ha-1 drought, frost and disease resist- yr-1 on the driest sites. ance. It is this huge adaptive plas- Because of its ability to ticity that has made black pine develop well on open lands such a favourite for reforestation and in ecologically demanding projects over a wide range of situations, Austrian pine was environments. intensively used during 19th and In the middle of the 20th cen- early 20th century reforestation tury, several provenance trials programmes, e.g. in the French were established independently southern Alps for landslide con- in Europe, the USA and New trol and land rehabilitation and in Zealand. The Corsican and Cal- England and the USA for sand- abrian black pine provenances dune fixation and as a wind- were found to be the best in break. Currently P.n. laricio is the almost every respect on siliceous most important reforestation soils. They had consistently lackPinus pinePinus nigraEuropean black nigra pinePinus nigraEuropean blackPinus pinePinus nigraEu

excellent stem form and branch- Threats to Damaging insects include ing habit, gave the greatest vol- genetic diversity European black pine shoot moth ume production and were hardy (Rhyacionia buoliana), pine pro- against winter and late frosts Black pine is not recognized as a cessionary caterpillar (Thaume- (except in north-central USA). threatened species although topoea pityocampa), especially The major defect reported is some of its sub-Mediterranean in warm and dry climates, and tip branch forking, which is both endemic populations constitute blight (Sphaeropsis sapinea), heritable and highly correlated priority habitats under the EU which has been particularly with polycyclism and branch Natura 2000 directive (Habitat active in France and Turkey in angle. On calcareous soils, P.n. Directive n° 92/43/CEE, May 21 1990’s. Other pests such as laricio does not perform well and 1992). Acantholyda hieroglyphica, Dipri- is to be replaced by the slower- Extensive plantations were on pini, Pissodes validirostis and growing but more Ca-tolerant often made across Europe in the Monophlebus hellenicus have P.n. nigra. In dry climates (as in past two centuries with material been active in Turkey. Most inner Anatolia, Turkey), black from unknown and/or very dis- recently, an increase in the pine is slow growing and breed- tant sources for which no histor- impact of a needle blight known ing programmes for such zones ical traces currently exist. This as the ‘red band disease’ (Doth- are focused on improving growth has probably resulted in exten- istroma septospora) has been rate and increasing drought and sive mixing of local and imported reported. frost tolerance through within- genepools all over the distribu- population selection. tion area of black pine. Intraspecific hybridization is easily performed among all black pine subspecies (a further proof of phylogenetic relatedness), but has not contributed any out- standing genotypes to breeding programmes so far. Interspecific crosses seem to be possible at a low survival rate with P. sylvestris. Black pine seed orchards have been established in several European countries, e.g. in France there are one Calabrian pine and two Corsican pine seed orchards. Current experiments in vegetative propagation include micropropa- gation of zygotic embryos and brachyblasts as well as somatic embryogenesis. Propagation by grafting has been known since 1820; the method generally used is lateral grafting. PinusEuropean black pinePinus nigranigraEuropean black pinePinus Pinu nigraEuropean black

Distribution range of European black pine

In areas where black pine is Guidelines for genetic encouraged separately as seed widespread and very important conservation and use stands and gene conservation for forestry, factors such as forest forests. Those do not serve the fires and illegal cutting cause Because black pine of different same goal and should not serious damage. In areas where it origins has been extensively always be identical, especially to occurs in small isolated popula- planted, it is now important to make the conservation of mar- tions, major risks come from any identify authochthonous popula- ginal populations possible. An factor that may provoke local tions. This undertaking should be international in situ network of extinction, either through illegal carried out at the international 100–120 stands would seem cutting and fires or through level. In each country, an inven- appropriate to represent the nat- hybridization (‘genetic pollution’) tory should be made to define ural ecological and genetic vari- from planted black pines belong- the geographical distribution of ability of black pines. ing to other subspecies. Original the species, its conservation sta- As intraspecific hybridization and rare varieties such as P. nigra tus, threats and potential uses. is easy among black pines, exot- var. pyramidalis or P. nigra var. Breeding activities provide valu- ic or improved black pines should sheneriana in Turkey are under able information by defining not be planted in the vicinity of identical threats. potential plantation, seed collec- autochthonous and naturalized tion and transfer zones. In situ stands. This is particularly true for conservation activities should be localized and fragmented sub- usk pinePinus nigra nigraEuropean black pine Pinus nigraEuropean black pine nigPinus nigraEuro EUFORGEN

These Technical Guidelines were species such as P.n. laricio, and is of extreme importance for sub- produced by members of the species that are threatened, such as P.n. salzmanii in France and P.n. EUFORGEN Conifers Network. mauretanica in North Africa. For these subspecies and other varieties The objective of the Network is to of rare occurrence, ex situ conservation is appropriate and urgent. As identify minimum genetic conser- a step in that direction, in 1999 a gene conservation forest was select- vation requirements in the long ed in Turkey for the rare P. nigra var. pyramidalis. term in Europe, in order to reduce Information on the provenance and progeny trials established the overall conservation cost and throughout Europe should be entered in a database. This network of to improve the quality of stan- experimental sites could be used for ex situ conservation of black dards in each country. pine. Marginal areas might need to be further sampled to strengthen this network and possibly planted as ex situ seed orchards to re-install depleted resources.

Citation: Isajev, V., B. Fady, H. Selected bibliography Semerci and V. Andonovski. 2004. EUFORGEN Technical Guidelines for genetic conserva- Lauranson-Broyer, J. and Ph. Lebreton. 1995. Flavonic chemosystematics of the specific complex Pinus nigra Arn. Pp. 181-188 in Population genetics tion and use for European black and genetic conservation of forest trees (P. Baradat, W.T. Adams and G. pine (Pinus nigra). International Müller-Starck, eds.). SPB Academic Publishing, Amsterdam. Plant Genetic Resources Insti- Nikolic, D. and N. Tucik. 1983. Isoenzyme variation within and among popula- tute, Rome, Italy. 6 pages. tions of European black pines (Pinus nigra Arnold). Silvae Genetica 32(3- 4):80-89. Drawings: Pinus nigra, Claudio Quézel, P. and F. Médail. 2003. Ecologie et biogéographie des forêts du bassin Giordano. © IPGRI, 2003. méditerranéen. Elsevier, Paris. Tutin, T.G., V.H. Heywood, N.A. Burgess, D.M. Moore, D.H. Valentine, S.M. Walters and D.A. Webb, eds. 1983. Flora Europaea, Vol 1, 2nd edition, pp. ISBN 92-9043-659-X 40-44. Cambridge University Press, Cambridge, UK.

Vidakovic,` M. 1974. Genetics of European black pine (Pinus nigra Arn.). Ann. Forest. 6/3 JAZU Zagreb:57-86.

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