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Mediterranean Pines (Pinus Halepensis Mill. and Brutia Ten.)

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Mediterranean Pines (Pinus Halepensis Mill. and Brutia Ten.) Chapter 5 1 Mediterranean Pines (Pinus halepensis 2 Mill. and brutia Ten.) 3 [AU1] Maria Regina Chambel, Jose Climent, Christian Pichot, and Fulvio Ducci 4 5.1 Introduction 5 Pinus halepensis Mill. (Aleppo pine) and P. brutia Ten. (Brutia pine or Turkish red 6 pine) together cover more than 7 million hectare around the Mediterranean Basin 7 and play major ecological and economical roles in low- to mid-elevation Mediter- 8 ranean forests. Both species are well adapted to dry summer conditions and are able 9 to successfully colonize abandoned arable lands and burned areas. Apart from 10 the high ecological value of natural stands, these species can create highly resilient 11 forest covers in limiting dry conditions for both production and protection purposes. 12 Although large adaptive genetic diversity was demonstrated in both species, Aleppo 13 pine materials are globally more tolerant to water stress while Brutia pine is, in 14 general, less susceptible to frost damage. 15 Even when genetic improvement of these species is less developed compared to 16 other species of the genus in Europe, such as Scots pine or Maritime pine, provenance 17 testing has been carried out in most countries thanks to international collaborative 18 With contributions of: Ernesto Fusaro (CRA PLF, IT), Eduardo Notivol (CITA, ES), Francisco Auñón (INIA, ES), Paraskevi Alizoti (AUTH, GR), Şükran Gökdemir (CAFRD, TUR), Leonid Korol (Volcani C., IL), Mohamed Larbi Khouja (INRGREF, TUN), Hassan Sbay (FRC, MOR). M.R. Chambel • J. Climent (*) Instituto Nacional de Investigacion y Tecnologia Agraria y Alimentaria (INIA), 28080 Madrid, Spain e-mail: [email protected] C. Pichot Institut National de la recherche agronomique (INRA), Avignon, France F. Ducci Consiglio per la ricerca e Sperimentazione in Agricoltura – Istituto Sperimentale per la Selvicoltura (CRA), 52100 Arezzo, Italy e-mail: [email protected] L.E. Pâques (ed.), Forest Tree Breeding in Europe: Current State-of-the-Art and Perspectives, Managing Forest Ecosystems 25, DOI 10.1007/978-94-007-6146-9_5, © Springer Science+Business Media Dordrecht 2013 M.R. Chambel et al. 19 initiatives (FAO Silva Mediterranea, IUFRO and EU projects). Standard breeding 20 programmes based on seedling and clonal seed orchards have been developed in 21 Spain (among TreeBreedex partners) and particularly in Greece for Aleppo pine 22 and in Turkey for Brutia pine, outwith TreeBreedex. The target of these programmes 23 has been not only yield improvement, but especially adaptability and endurance 24 under limiting environmental conditions, particularly summer drought. 25 5.2 Taxonomy, Biology and Ecology of Aleppo Pine 26 5.2.1 Taxonomy and Species Distribution 27 Pinus halepensis Mill. and Pinus brutia Ten. are indeed two very close taxa, formerly 28 included in a separate section or subsection Halepenses (Price et al. 1998; López 29 et al. 2002). However, recent classifications of subgenus Pinus (Dyploxylon or 30 “hard” pines) tend to group these two species with P. heldreichii, P. pinaster, P. pinea, 31 P. canariensis and P. roxburghii within subsection Pinaster, also called the 32 Mediterranean pine group (Gernandt et al. 2005, 2008). 33 Both species have circum-Mediterranean ranges of distribution, sometimes 34 consisting of isolated populations, but occupy different geographical ranges and 35 bioclimatic niches along the Mediterranean basin (Panetsos 1981; Nahal 1986; 36 Vidakovic 1991; Fig. 5.1). Nevertheless, natural hybrids exist through unidirec- 37 tional pollen flow from P. halepensis as the pollen donor to P. brutia acting as the 38 female tree. This occurs in some Greek stands (Chalkidiki peninsula, where Pinus 39 halepensis Mill. meets the easternmost limit of its distribution in the Mediterranean 40 basin, Panetsos 1975; Panetsos et al. 1997) and in several Turkish stands. Distinction 41 between P. halepensis, P. brutia and their natural hybrids has long been discussed, 42 both in distant and sympatric populations (Panetsos 1975; Panetsos et al. 1997; 43 Tozkar et al. 2009). Controlled pollination experiments suggest that partial repro- 44 ductive barriers exist between the two species (Panetsos 1975). 45 P. halepensis Mill. occupies the southernmost area of the Mediterranean pines 46 (with the exception of P. canariensis) and it is widespread in the western part of the 47 Mediterranean Sea (ranging from 45° to 31°N), including North Africa (Morocco, 48 Algeria, Tunisia, Libya), the southern regions of France, Italy, eastern Spain, Greece 49 and Malta. It is also present in coastal areas of Croatia and Albania. There are some 50 natural and artificial populations in Turkey, Jordan, Israel, Lebanon and Syria (hence 51 the name Aleppo pine). 52 The natural range of Pinus brutia Ten. is the Eastern Mediterranean (ranging 53 from 44° to 35°N), from Greece to the eastern Crimea, Georgia, northern Iraq, west- 54 ern Syria, Lebanon and Cyprus. It is particularly abundant in Turkey, justifying its 55 English common name (Turkish red pine). In Italy, P. brutia is not indigenous, 56 although it is naturalized in some hilly areas of southern Calabria and Salento where 57 it was probably introduced by the Romans (who called it Calabria: Bruttium). 58 In the margins of its distribution area, three varieties (besides the type) and 59 one subspecies are recognized (Frankis 1999): P. brutia var. pityusa (Steven) Silba 5 Mediterranean Pines (Pinus halepensis Mill. and brutia Ten.) Fig. 5.1 Range of distribution of Pinus halepensis (green) and P. brutia (blue) (adapted from Euforgen) (syn: P. pityusa Steven), is widespread in Georgia and along the Russian coasts of 60 the Black Sea; P. brutia var. stankewiczii (Sukaczev) Frankis, is present in the 61 Crimea and the Ukraine; P. brutia var. pendulifolia Frankis. is widespread in coastal 62 regions of southern Turkey and P. brutia subsp. eldarica (Medw.) Nahal (syn: 63 P. eldarica Medw.) is typical of the Caucasus (mainly in Azerbaijan, but also in 64 Georgia, Iran and Turkey). It is considered a separate species (Pinus eldarica) by 65 some authors, adapted to drier and colder climates, with rainy summers. It was 66 probably introduced from Azerbaijan into the Mediterranean during the rule of 67 Alexander the Great. 68 5.2.2 Biology 69 Secondary needles of P. brutia are darker, longer (10–16 cm) and thicker (1.5 mm) 70 than those of Aleppo pine, and with a significantly higher leaf mass per area (Pardos 71 et al. 2009). They have three lines of hypodermal cells and resin canals, while in 72 P. halepensis resin canals are marginal (Vidakovic 1991). Primary needles, very 73 similar in both species, are retained longer in Aleppo pine than in Brutia pine. 74 The more precocious formation of secondary needles and first bud set in Brutia pine 75 is one of the traits distinguishing both species more clearly at early developmental 76 stages (Climent et al. 2011). 77 M.R. Chambel et al. Fig. 5.2 Extremely different winter buds of Brutia pine (left) and Aleppo pine (right) taken at the same moment in 3-year-old plants at the nursery. Shoot elongation in Aleppo pine does not actually stop during mild winters (J. Climent) 78 P. brutia varieties are not very distinct from the typical form. Var. pityusa has 79 shorter needles; var. stankewiczii has solitary cones and seeds with shorter wings 80 and var. pendulifolia has 20–29 cm long weeping needles. By contrast, the Eldar 81 pine (P. brutia subsp. eldarica) has a more slender growth habit, 5–6 cm long cones 82 with a short stalk and shorter and thicker needles (8.5–10 cm) (Vidakovic 1991). 83 Both species have the ability to produce multiple flushes during the growing 84 period (polycyclic shoot growth), but this phenomenon is more extreme in Aleppo 85 pine (Isik et al. 2002; Pardos et al. 2003). Shoots of Aleppo pine can restart elonga- 86 tion very early in winter, and continue growing well into late autumn in mild climates 87 (Fig. 5.2). Multiple female flowering in the same year is not rare, increasing the 88 amount of cones of the same yearly cohort (Climent et al. 2008; Pardos et al. 2003) 89 (Fig. 5.3a). 90 Cones of P. brutia are sessile or sub-sessile, while the cones of P. halepensis have 91 a 2–3 cm long petiole (Fig. 5.3b). Seeds of Brutia pine have a length of between 6 92 and 9.5 mm, with a 15–20 mm wing, while Aleppo pine seeds are 5–6 mm long with 93 a wing of 20 mm. Seeds of Aleppo pine are the lightest among Mediterranean pines: 94 between 45,000 and 65,000 seeds/kg, versus about 20,000 seeds/kg in Brutia pine. 95 Aleppo pine is one of the most reproductively precocious pine species, starting 96 to produce female cones as early as 3 years of age (Tapias et al. 2001, 2004). Besides, 97 high female fecundity and partial serotiny or xeriscence ensure the precocious 98 maintenance of an important aerial seed bank (up to one million seeds per ha; 99 Ne’eman et al. 2004; Tapias et al. 2001) (Fig. 5.3c). Brutia pine is less precocious 100 and serotinous, but it is still able to keep a variable aerial seed bank. Both species 101 start their reproductive life as females, while male strobili appear in lower or 102 secondary branches when the crown attains a certain complexity level (Ne’eman 103 et al. 2004; Shmida et al. 2000). 5 Mediterranean Pines (Pinus halepensis Mill. and brutia Ten.) Fig. 5.3 Reproductive features of Aleppo pine: (a) multiple female flowering cycles of the same vegetative period, (b) mature cones, (c) serotinous cone in a young tree showing the characteristic thin light grey bark and (d) male strobili (L.
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