Species breeding monograph: Aleppo pine (Pinus halepensis Mill.) and Brutia pine (Pinus brutia Ten.)

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

5.1 Introduction

Pinus halepensis Mill. (Aleppo pine) and P. brutia Ten. (Brutia pine or Turkish red pine) together cover more than 7 million hectare around the Mediterranean Basin and play major ecological and economical roles in low- to mid-elevation Mediter-ranean forests. Both species are well adapted to dry summer conditions and are able to successfully colonize abandoned arable lands and burned areas. Apart from the high ecological value of natural stands, these species can create highly resilient forest covers in limiting dry conditions for both production and protection purposes. Although large adaptive genetic diversity was demonstrated in both species, Aleppo pine materials are globally more tolerant to water stress while Brutia pine is, in general, less susceptible to frost damage.

Even when genetic improvement of these species is less developed compared to other species of the genus in Europe, such as Scots pine or Maritime pine, provenance testing has been carried out in most countries thanks to international collaborative

Chapter 5Mediterranean Pines (Pinus halepensis Mill. and brutia Ten.)

Maria Regina Chambel, Jose Climent, Christian Pichot, and Fulvio Ducci [AU1]

M.R. Chambel • J. Climent (*)Instituto Nacional de Investigacion y Tecnologia Agraria y Alimentaria (INIA),28080 Madrid, Spaine-mail: [email protected]

C. PichotInstitut National de la recherche agronomique (INRA), Avignon, France

F. DucciConsiglio per la ricerca e Sperimentazione in Agricoltura – Istituto Sperimentale per la Selvicoltura (CRA), 52100 Arezzo, Italye-mail: [email protected]

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).

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

M.R. Chambel et al.

initiatives (FAO Silva Mediterranea, IUFRO and EU projects). Standard breeding programmes based on seedling and clonal seed orchards have been developed in Spain (among TreeBreedex partners) and particularly in Greece for Aleppo pine and in Turkey for Brutia pine, outwith TreeBreedex. The target of these programmes has been not only yield improvement, but especially adaptability and endurance under limiting environmental conditions, particularly summer drought.

5.2 Taxonomy, Biology and Ecology of Aleppo Pine

5.2.1 Taxonomy and Species Distribution

Pinus halepensis Mill. and Pinus brutia Ten. are indeed two very close taxa, formerly included in a separate section or subsection Halepenses (Price et al. 1998; López et al. 2002). However, recent classifications of subgenus Pinus (Dyploxylon or “hard” pines) tend to group these two species with P. heldreichii, P. pinaster, P. pinea, P. canariensis and P. roxburghii within subsection Pinaster, also called the Mediterranean pine group (Gernandt et al. 2005, 2008).

Both species have circum-Mediterranean ranges of distribution, sometimes consisting of isolated populations, but occupy different geographical ranges and bioclimatic niches along the Mediterranean basin (Panetsos 1981; Nahal 1986; Vidakovic 1991; Fig. 5.1). Nevertheless, natural hybrids exist through unidirec-tional pollen flow from P. halepensis as the pollen donor to P. brutia acting as the female tree. This occurs in some Greek stands (Chalkidiki peninsula, where Pinus halepensis Mill. meets the easternmost limit of its distribution in the Mediterranean basin, Panetsos 1975; Panetsos et al. 1997) and in several Turkish stands. Distinction between P. halepensis, P. brutia and their natural hybrids has long been discussed, both in distant and sympatric populations (Panetsos 1975; Panetsos et al. 1997; Tozkar et al. 2009). Controlled pollination experiments suggest that partial repro-ductive barriers exist between the two species (Panetsos 1975).

P. halepensis Mill. occupies the southernmost area of the Mediterranean pines (with the exception of P. canariensis) and it is widespread in the western part of the Mediterranean Sea (ranging from 45° to 31°N), including North Africa (Morocco, Algeria, Tunisia, Libya), the southern regions of France, Italy, eastern Spain, Greece and Malta. It is also present in coastal areas of Croatia and Albania. There are some natural and artificial populations in Turkey, Jordan, Israel, Lebanon and Syria (hence the name Aleppo pine).

The natural range of Pinus brutia Ten. is the Eastern Mediterranean (ranging from 44° to 35°N), from Greece to the eastern Crimea, Georgia, northern Iraq, west-ern Syria, Lebanon and Cyprus. It is particularly abundant in Turkey, justifying its English common name (Turkish red pine). In Italy, P. brutia is not indigenous, although it is naturalized in some hilly areas of southern Calabria and Salento where it was probably introduced by the Romans (who called it Calabria: Bruttium).

In the margins of its distribution area, three varieties (besides the type) and one subspecies are recognized (Frankis 1999): P. brutia var. pityusa (Steven) Silba

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

5 Mediterranean Pines (Pinus halepensis Mill. and brutia Ten.)

(syn: P. pityusa Steven), is widespread in Georgia and along the Russian coasts of the Black Sea; P. brutia var. stankewiczii (Sukaczev) Frankis, is present in the Crimea and the Ukraine; P. brutia var. pendulifolia Frankis. is widespread in coastal regions of southern Turkey and P. brutia subsp. eldarica (Medw.) Nahal (syn: P. eldarica Medw.) is typical of the Caucasus (mainly in Azerbaijan, but also in Georgia, Iran and Turkey). It is considered a separate species (Pinus eldarica) by some authors, adapted to drier and colder climates, with rainy summers. It was probably introduced from Azerbaijan into the Mediterranean during the rule of Alexander the Great.

5.2.2 Biology

Secondary needles of P. brutia are darker, longer (10–16 cm) and thicker (1.5 mm) than those of Aleppo pine, and with a significantly higher leaf mass per area (Pardos et al. 2009). They have three lines of hypodermal cells and resin canals, while in P. halepensis resin canals are marginal (Vidakovic 1991). Primary needles, very similar in both species, are retained longer in Aleppo pine than in Brutia pine. The more precocious formation of secondary needles and first bud set in Brutia pine is one of the traits distinguishing both species more clearly at early developmental stages (Climent et al. 2011).

Fig. 5.1 Range of distribution of Pinus halepensis (green) and P. brutia (blue) (adapted from Euforgen)

60

61

62

63

64

65

66

67

68

69

70

71

72

73

74

75

76

77

M.R. Chambel et al.

P. brutia varieties are not very distinct from the typical form. Var. pityusa has shorter needles; var. stankewiczii has solitary cones and seeds with shorter wings and var. pendulifolia has 20–29 cm long weeping needles. By contrast, the Eldar pine (P. brutia subsp. eldarica) has a more slender growth habit, 5–6 cm long cones with a short stalk and shorter and thicker needles (8.5–10 cm) (Vidakovic 1991).

Both species have the ability to produce multiple flushes during the growing period (polycyclic shoot growth), but this phenomenon is more extreme in Aleppo pine (Isik et al. 2002; Pardos et al. 2003). Shoots of Aleppo pine can restart elonga-tion very early in winter, and continue growing well into late autumn in mild climates (Fig. 5.2). Multiple female flowering in the same year is not rare, increasing the amount of cones of the same yearly cohort (Climent et al. 2008; Pardos et al. 2003) (Fig. 5.3a).

Cones of P. brutia are sessile or sub-sessile, while the cones of P. halepensis have a 2–3 cm long petiole (Fig. 5.3b). Seeds of Brutia pine have a length of between 6 and 9.5 mm, with a 15–20 mm wing, while Aleppo pine seeds are 5–6 mm long with a wing of 20 mm. Seeds of Aleppo pine are the lightest among Mediterranean pines: between 45,000 and 65,000 seeds/kg, versus about 20,000 seeds/kg in Brutia pine.

Aleppo pine is one of the most reproductively precocious pine species, starting to produce female cones as early as 3 years of age (Tapias et al. 2001, 2004). Besides, high female fecundity and partial serotiny or xeriscence ensure the precocious maintenance of an important aerial seed bank (up to one million seeds per ha; Ne’eman et al. 2004; Tapias et al. 2001) (Fig. 5.3c). Brutia pine is less precocious and serotinous, but it is still able to keep a variable aerial seed bank. Both species start their reproductive life as females, while male strobili appear in lower or secondary branches when the crown attains a certain complexity level (Ne’eman et al. 2004; Shmida et al. 2000).

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

79

80

81

82

83

84

85

86

87

88

89

90

91

92

93

94

95

96

97

98

99

100

101

102

103


Bark aspect and thickness is another major feature distinguishing Pinus halepensis and P. brutia. While Aleppo pine bark is extremely thin; silver grey at early ages and later ash-grey, Pinus brutia shows a thicker reddish bark from young ages.

5.2.3 Ecology

Pinus halepensis is a thermophilic, drought-tolerant species; in fact it is probably one of the pines most tolerant to high temperatures and drought (Magini 1955; Scarascia-Mugnozza 1986).

In the northern part of the natural distribution range it grows in coastal areas (from sea level up to 600 m), while in the south it can climb up to 1,400 m (the absolute maximum is 2,600 m in the Atlantic mountains of Morocco; Fady et al. 2003). The optimum climatic conditions for this species are 350–700 mm annual rainfall (semi-arid and sub-humid climate) and between −2 and 10 °C absolute minimum temperatures. P. brutia is less thermophilic (minimum temperature between −5 and 10 °C), but more water-demanding (it needs humid/sub-humid climates, with annual rainfall ranging from 400 to 1,300 mm). The altitude limits of this species are between 0 and 600 m, but in the southern part of its natural range it can reach 1,200–1,400 m (1,650 m on the Taurus Mountains in Turkey).

Pinus halepensis is one of the European tree species able to survive in shallower soils, particularly on limestone bedrocks, provided it has a favourable temperature range. In Spain, France, Italy, Balkans and Greece 90 % of Aleppo pine stands grow on calcareous soils, chalk and limestone shallow soils; however, it can grow well on moderately acid soils, although on these soils it is frequently outcompeted by other species, such as stone pine or maritime pine.

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. Santos del Blanco)

104

105

106

107

108

109

110

111

112

113

114

115

116

117

118

119

120

121

122

123

124

125

126

M.R. Chambel et al.

The relationship between Pinus halepensis and fire has been the subject of intensive research (Daskalakou and Thanos 1996; Ne’eman et al. 1992, 2004; Pausas et al. 2004; Tapias et al. 2001; Thanos et al. 1996) following widespread concern after the highly destructive fires that occurred in some European Mediterranean countries over recent decades. Briefly, Aleppo pine is capable of having a very intense recruitment after stand-replacing fires, with seedling densities as high as 50,000–200,000 seedlings/ha (Ne’eman et al. 1992); but, on the other hand, it is very flammable and its capability of withstanding low-intensity fires is very low compared to, for example, Pinus pinea (Pausas et al. 2008; Rigolot 2004). In fact, Aleppo pine can be considered a model of an evader strategy in the face of fire, characterized by a low investment in adult endurance combined with the previ-ously mentioned precocious high fertility (Keeley and Zedler 1998; Tapias et al. 2004). This extreme behaviour is less marked in Pinus brutia, but this species is also capable of recruiting successfully after destructive crown fires; for example, densi-ties of 20,000 seedlings/ha 5 years after a stand-replacing fire have been reported in Greece (Spanos et al. 2000).

Aleppo pine is one of the first species used for afforestation in dry conditions. The most emblematic programme is probably the “Green Belt” programme in South Algeria where 100,000 ha were planted with Aleppo pine over 20 years. However, the extensive use of Aleppo pine for afforestation in highly degraded soils in the western Mediterranean has been the object of strong criticism by ecologists and environmentalists for various reasons; the species has been blamed for increasing the risk of fire, reducing biodiversity and diminishing water resources (Bellot et al. 2004; Maestre et al. 2003). This negative view starts with the lack of recognition of the Aleppo pine as a true native species out of very narrow limits, a gross scientific error that still lingers within some collectives (see a discussion in Gil et al. 1996).

Aleppo pine can rapidly colonize burned areas covered by other species, less adapted to frequent fires, like Pinus pinaster or Pinus nigra. This high colonizing capacity, based on the already mentioned reproductive traits, makes this species one of the major invasive alien trees in South Africa where, together with P. pinaster and P. radiata, it constitutes a conservation problem in large areas of the Cape peninsula covered with fynbos, the highly diverse native shrubland (Richardson 1998; Richardson et al. 1990). In south-western Australia, Aleppo pine has been declared as an invader weed (www.environment.sa.gov.au) to be eradicated but, at the same time, it has been promoted for low-input forestry farming.

5.3 Aleppo and Brutia Pines in Treebreedex Countries

5.3.1 Native and Cultivated Area

Considering the figures for natural and cultivated areas of Aleppo pine within TreeBreedex countries (Table 5.1), it could be considered as a minor species with the exception of Spain, where it represents as much as 45 % of the total forest

127

128

129

130

131

132

133

134

135

136

137

138

139

140

141

142

143

144

145

146

147

148

149

150

151

152

153

154

155

156

157

158

159

160

161

162

163

164

165

166

http://www.environment.sa.gov.au


area. In all three countries, Aleppo pine remains essentially a wild species, with a low percentage of cultivated forests (22 % in Spain, 14 % in France and 8.8 % in Italy).

5.3.2 Products and Major Uses

Intrinsic wood quality of Aleppo and Brutia pines is very similar to that of other Mediterranean pines, for example P. pinaster (Giordano 1976; Thibault et al. 1992). However, the wood of both species is of variable interest in different countries depending on the availability of other pine wood of better quality. In general, the wood provided by P. brutia has a higher quality due to the better stem form and branching, but this depends largely on site quality. Mean productivity is approximately 1–3 m3 ha−1 year−1 for Aleppo pine, and 2–3 m3 ha−1 year−1 for Brutia pine. According to the French Forest Inventory Service (Anonymous 2010), mean productivity in France averages 2.3 m3 ha−1 year−1 but it can reach 5 m3 ha−1 year−1 in favourable conditions. It must be stressed that this productivity was strongly increased (+40 %) over the last 25 years. Maximum yield can reach 12–15 m3 ha−1 year−1 in high-quality stands for both species.

Some non-wood products are also obtained from these species. Seeds are used for making pastry (particularly in Tunisia). Resin was extracted in some countries during the twentieth century, but this use is currently practically abandoned, except in Greece, were it is still used for wine production (retsina). Bark was used for tannin extraction, but nowadays this use is limited to North Africa. A curious, yet economically relevant, non-wood production of these pines is the honey produced in Greece and Turkey from the honeydew released by the sap-sucking insect Marchalina hellenica (Bacandritsos et al. 2004; Santas 1983).

In Spain, Pinus halepensis is widely used for land protection and afforestation of degraded mountain areas. Extensive planting of this species took place from the early twentieth century for land reclamation and soil protection both within and outside its natural range. In particular, moderate success was achieved in the refor-estation of gypsum-limestone marls at areas of extreme Continental climate in the

[AU2]

Table 5.1 Native and cultivated area covered by Aleppo pine in Treebreedex countries

CountryNative range (ha)

Cultivated range (ha)

Total range (ha)

Total area of conifers (ha)

Total forest area (ha)

Spain 2,345,663 649,211 2,994,874 6,374,650 (+ 3,498,643 mixed conifer and broadleaf stands)

27,872,829

France 250,000 ~0 250,000 4,800,000 16,100,000Italy 226,101a 20,000 143,300 1,388,000 8,860,701aArea occupied by Mediterranean pine forests: P. halepensis, P. pinaster and P. pinea

167

168

169

170

171

172

173

174

175

176

177

178

179

180

181

182

183

184

185

186

187

188

189

190

191

192

193

194

195

t1.1

t1.2

t1.3

t1.4

t1.5

t1.6

t1.7

t1.8

t1.9

M.R. Chambel et al.

Duero basin (northern Castillian plateau). Annual fellings are about 206,250 m3 (2007 data). Timber of this species is not appreciated because the density is too high, and there are numerous knots and resin pockets, even when other mechanical properties (MOE) are adequate. There is some use for pallets and chipping for par-ticleboards; it is also used for boat making at a local scale. Aleppo pine is frequently planted in rain-fed suburban parks and road lines.

In France this species is mainly used for soil preservation in erodible calcareous slopes and for recreation forestry. However, there are very few plantations and the increase of Aleppo pine forests is mostly due to natural regeneration in abandoned agricultural lands. Although it is sometimes used as fuel or in the pallet industry, its wood is mostly used for paper production (200,000 tm year−1). The species is also used as an ornamental tree and for road line plantations.

In Italy, afforestation with Aleppo pine is carried out in coastal areas and in the adjacent hills. This species was widely planted between 1930 and 1970 in Mediterranean areas for soil protection and wind breaking near the coasts and for resin extraction, staining products and wood manufacture. Indeed, the wood of this species was used for boat construction; nowadays it is mainly used for dune protec-tion, for ornamental proposes, for road line plantations and as raw material for the pulp and paper industry. Resin production, tannin extraction for leather and fishing nets were also important traditional uses. However, the recreational use of this species is nowadays the dominant use along the coasts.

5.3.3 Plant Needs for Reforestation

There is a marked lack of statistics within TreeBreedex countries concerning the actual number of plants consumed for afforestation with Aleppo pine, due to the highly dispersed information between countries and regions. The EU Common Agricultural Policy has increased the use of this species for afforestation of private former arable lands. For example, in Spain between 2003 and 2006 a total of 29,113 ha were planted with Aleppo pine. This gives us about 28 million plants (considering an average use of 800 plants/ha and replanting rates of 20 %, Fig. 5.4). In France and Italy plant production is presently very low due to the lack of affores-tation with this species and the limited use for ornamental purposes.

In Italy, during the forest fires of 2007, about 10,000 ha of natural and afforested areas planted with Pinus halepensis and P. brutia were destroyed in the Abruzzo and Puglia regions. Reforestation programmes to recover these areas have started at least in the Puglia region, due to the touristic and economic importance of forests in this area.

To complete reforestation of these and other burned areas as in Greece (Greece is certainly the country the most affected by the great fires of recent years), the global potential demand for seedlings could be to the tune of several million in the next years. Intensive exchange of reproductive materials occurs between Italian and Greek nurseries, but there are no statistics on these figures.

196

197

198

199

200

201

202

203

204

205

206

207

208

209

210

211

212

213

214

215

216

217

218

219

220

221

222

223

224

225

226

227

228

229

230

231

232

233

234

235

236


5.4 The Current Situation as a Starting Point for Breeding

Within the TreeBreedex consortium, there are no active breeding programmes of Aleppo or Brutia pine at the same level of those of major species (Norway spruce, Sitka spruce, Scots pine, etc.). Provenance research can be considered as part of low-intensity breeding programmes in France, Italy and Spain aimed mainly at assuring self-maintenance of low-production and protective forests.

5.4.1 Provenance Research Within TreeBreedex Countries

5.4.1.1 FAO International Network

Between 1977 and 1982, FAO/Silva Mediterranea, within the project FAO/SCM/ CRFM/4 bis, promoted a large network of provenance trials with species of the halepensis group. This programme allowed the collection of seeds from 57 prove-nances (33 of Aleppo pine, 17 of Brutia pine and seven of Eldarica pine) covering the natural range of the species (Fig. 5.5) and supported the establishment of several field trials across the Mediterranean and in other areas with a Mediterranean climate like Australia (Spencer 1985) or New Mexico (Fisher et al. 1986). Twelve of these trials are located in TreeBreedex countries (10 in Italy and 2 in the south-east of France; Tables 5.2 and 5.3; Fig. 5.6).

Fig. 5.4 Aleppo pine plant stock at TRAGSA nurseries, Spain (J. Climent)

237

238

239

240

241

242

243

244

245

246

247

248

249

250

251

252

253

M.R. Chambel et al.

Considering the age of these trials, the results can now be regarded as definitive. In general, growth traits displayed high plasticity and significant species × site interaction across trial-sites; however, provenance rankings tended to remain rather stable (i.e. there is quantitative but not qualitative g × e interaction). On the other hand, traits such as stem form and frost resistance showed lower levels of interaction both between species and between provenances within species.

In Italy, in the trial sites situated at higher altitudes, a long and hard frost occurred in 1985 which had devastating consequences for most of the Aleppo pine provenances, with the French provenance A24 – Gemenos being the least affected (Bariteau 1992; Eccher et al. 1987; Ducci and Guidi 1998). In sites where temperatures reached −24 °C for as long as 2 weeks, only Pinus brutia provenances managed to survive. Therefore, Pinus halepensis provenances should be planted only in truly Mediterranean conditions, while P. brutia should preferably be used in interior and more continental sites. In these conditions, the best performing and most stable provenances were those form the Chalkidiki (A4 and A5) peninsula (North-eastern Greece), A27-Vico del Gargano and A28-Patemisco (Italy), A6-Shaharia and A7-Elkosh (Israel). Furthermore, these provenances, together with the French provenance Gemenos, were also characterized by good stem quality.

Fig. 5.5 Location of the provenances sampled for the FAO/Silva Mediterranea trial network (dots) and location of the trials in TreeBreedex countries (flags). Red dots represent provenances of Pinus halepensis and orange dots represent P. brutia. Dark dots represent provenances not included in TBX countries trials

254

255

256

257

258

259

260

261

262

263

264

265

266

267

268

269

270

271

272


Tabl

e 5.

2 Pr

oven

ance

tria

ls in

stal

led

with

in th

e fr

amew

ork

of F

AO

/Silv

a M

edite

rran

ea b

etw

een

1977

and

198

2 in

TB

X c

ount

ries

Site

Prov

ince

Reg

ion

Lat

. (N

)L

ong.

(E

)A

ltitu

de (

m)

Plan

ting

year

Mea

n an

nual

ra

infa

ll (m

m)

Mea

n an

nual

te

mp.

(°C

)

A. C

olle

mez

zano

tePe

scar

aA

bruz

zo41

° 13

¢13

° 52

¢70

019

7997

612

.0B

. Len

tisce

teC

ampo

bass

oM

olis

e41

° 49

¢14

° 50

¢17

519

8168

515

.5C

. Pix

inam

anna

Cag

liari

Sard

inia

38°

58¢

8° 5

7¢16

519

8262

516

.7D

. Cas

tiada

sC

aglia

riSa

rdin

ia39

° 10

¢9°

33¢

140

1982

502

17.7

E. A

rezz

oA

rezz

oT

usca

ny43

° 27

¢11

° 56

¢30

019

7782

812

.7F

. Ovi

leR

ome

Laz

io41

° 54

¢12

° 21

¢60

1975

834

16.0

G. C

aste

l di G

uido

Rom

eL

azio

41°

53¢

12°

09¢

4019

7571

414

.7H

. For

nari

Tara

nto

Apu

lia40

° 28

¢17

° 14

¢50

1976

544

16.6

I. C

astig

lionc

ello

Gro

sset

oT

usca

ny42

° 21

¢11

° 07

¢22

519

7665

514

.8J.

Mon

tebe

lloFo

rlì

Em

ilia-

Rom

agna

44°

07¢

11°

48¢

575

1976

1,02

011

.8K

. Cey

rest

eB

ouch

es-D

u-R

hône

Prov

ence

Côt

e d’

Azu

r43

° 13

¢5°

40¢

446

1976

786

15.0

L. V

itrol

les

Bou

ches

-Du-

Rhô

nePr

oven

ce C

ôte

d’A

zur

43°

28¢

5° 1

5¢20

019

7856

915

.5

Test

A. C

olle

mez

zano

tte w

as d

estr

oyed

by

fire

in 2

003

t2.1

t2.2

t2.3

t2.4

t2.5

t2.6

t2.7

t2.8

t2.9

t2.1

0

t2.1

1

t2.1

2

t2.1

3

t2.1

4

t2.1

5

t2.1

6

t2.1

7

t2.1

8

Table 5.3 List of provenances used in the FAO/Silva Mediterranea international network of field trials

Code Provenance name Country Longitude Latitude

Pinus halepensisA 1 Albania Albania 19°25¢ E 40°37¢A 2 Elea Greece 21°32¢ E 37°46¢A 3 Euboea Greece 23°18¢ E 38°58¢A 4 Chalkidiki 1 Greece 23°21¢ E 40°11¢A 5 Chalkidiki 2 Greece 23°44¢ E 40°03¢A 6 Shaharia Israel 34°50¢ E 31°36¢A 7 Elkosh Israel 35°18¢ E 33°01¢A 8 Sakiet Sidi Y. Tunisia 08°25¢ E 36°15¢A 9 Qoum Djeddur Tunisia 08°57¢ E 35°38¢A10 Djebel Selloum Tunisia 08°40¢ E 35°05¢A12 Zaouia Ifrane Morocco 05°23¢ W 33°15¢A13 J. Afra Selm.te Morocco 07°55¢ W 30°44¢A14 Quardane B. Morocco 05°08¢ W 35°03¢A15 Tamga Zaonia Morocco 06°07¢ W 32°02¢A17 Guadameldina Spain 02°15¢ W 37°02¢A24 Gemenos France 05°40¢ E 43°25¢A25 Imperia Italy 08°03¢ E 43°54¢A26 Otricoli Italy 12°38¢ E 42°24¢A27 Vico del Garg. Italy 16°00¢ E 41°54¢A28 Patemisco Italy 17°20¢ E 40°39¢A29 Aures Beni M. Algeria 06°50¢ E 35°10¢A30 Senalba Algeria – –A31 Telagh Algeria – –A32 Ourasensis Algeria 05°04¢ E 35°05¢A33 Lebanon Lebanon – –

Pinus brutiaB 1 Chania Greece 23°57¢ E 35°17¢B 2 Kavala Greece 24°42¢ E 40°48¢B 3 Lassithiou Greece 25°32¢ E 35°06¢B 5 Cyprus Cyprus 33°17¢ E 35°08¢B 6 Marmaris Turkey 28°18¢ E 37°00¢B 7 Isparta Turkey 29°32¢ E 38°04¢B 8 Duzlerçani Turkey 30°25¢ E 37°03¢B 9 Pamuçak Turkey 30°41¢ E 37°40¢B10 Bozburun Turkey 30°45¢ E 37°21¢B11 Bakara Turkey 32°43¢ E 36°09¢B12 Silifke Turkey 33°43¢ E 36°13¢B13 Gamgolu Turkey 35°20¢ E 41°50¢B14 Baspinar Turkey 35°15¢ E 37°48¢B15 Kisildag Turkey 35°58¢ E 36°21¢B16 Zawita Iraq 44°20¢ E 36°35¢B17 – Lebanon – –B25 UnknownB27 UnknownB29 Unknown

Pinus brutia subsp. eldaricaE 1 Karaj Iran 51°00¢ E 35°56¢The longitude is given as the first coordinate, given its importance in the range distribution

t3.1

t3.2

t3.3

t3.4

t3.5

t3.6

t3.7

t3.8

t3.9

t3.10

t3.11

t3.12

t3.13

t3.14

t3.15

t3.16

t3.17

t3.18

t3.19

t3.20

t3.21

t3.22

t3.23

t3.24

t3.25

t3.26

t3.27

t3.28

t3.29

t3.30

t3.31

t3.32

t3.33

t3.34

t3.35

t3.36

t3.37

t3.38

t3.39

t3.40

t3.41

t3.42

t3.43

t3.44

t3.45

t3.46

t3.47

t3.48

t3.49

t3.50

t3.51


According to results shown by Eccher et al. (1987), Bariteau (1992) and Ducci and Guidi (1998), P. brutia showed higher homogeneity of performances, allowing the definition of wider provenance regions which can be used as good seed sources: predominantly the southern Turkey coastal range between 30° and 36° longitude East. The easternmost provenances (B13 – Gamgolu to B16 – Zawita and E1) are the most suitable for the warm deciduous oak range areas, whilst the western basic material (B8 – Duzlerçani to B10 – Bozburun) performs better in the wet-cold evergreen/deciduous oak transition range. The best stem quality can be found in the oriental group of provenances. The Eldar pine performances could be interesting in ecological areas on lime stones, higher elevations and a more continental climate, whilst it must be avoided in lower sites and/or where there are larger amounts of clay.

In the two French sites, the lowest survival rates were observed for some P. halepensis provenances from Greece (A2-Elea), Italy (A27-Vico del Gargano and A26-Otricoli) and Morocco (A14-Ouardane Bouksane), while provenances from France (A24-Gémenos), Spain and north-eastern Greece exhibited good sur-vival rates, with no clear geographic pattern (Pichot and Vauthier 2007). The highest

Fig. 5.6 Field trial belonging to the FAO/Silva Mediterranea network at Ovile, Rome (E. Fusaro)

273

274

275

276

277

278

279

280

281

282

283

284

285

286

287

288

M.R. Chambel et al.

survival rates in this trial were those of the Pinus brutia provenances originating from eastern Taurus (B15-Kisildag and B14-Baspinar).

5.4.1.2 Other Provenance Trial Networks

More recent experimental networks were implemented in Spain and France for assessing respectively the variability of Aleppo pine national resources and evaluating the adaptation of Brutia pine in less dry but colder Mediterranean (deeper soils or higher elevation) conditions than those previously assessed.

In Spain, the network includes a six-site provenance trial with Spanish and Mediterranean-wide provenances and a three-site trial with a progeny within prov-enance structure, including only Spanish materials. The provenance trials were planted in 1998 with 42 populations from Spain, one from France, three from Greece, three from Italy and two from Tunisia. Additionally, three highly productive plantations of unknown origin located in north continental Spain, a commercial seed lot from eastern Spain and a seed lot from a Greek seed orchard (Fig. 5.7) were also included in these trials.

Phenotypic plasticity (site effect) was by far the main source of variation for growth variables in this trial, while provenance and g × e interaction had significant

Fig. 5.7 Location of provenances and experimental sites included in the Spanish Aleppo pine network of genetic trials. Black flags represent provenance trial sites (6); blue flags represent provenance/progeny trial sites (2); dots represent sampled populations, 24 of them included in both trials

289

290

291

292

293

294

295

296

297

298

299

300

301

302

303

304

305


but reduced effects. No clear geographic patterns were found for growth or phenotypic plasticity (Chambel et al. 2007). The higher rankings across sites for growth were obtained by the provenances from Greece, but their survival was generally low.

The provenance/progeny trial includes two sites with highly contrasting environ-ments (Fig. 5.8), planted in 1995 with 148 open-pollinated families from 32 popula-tions that cover the entire natural and planted range of peninsular Spain and Balearic islands (Santos-del-Blanco et al. 2010); 24 of these populations are also included in the provenance trial. Although also in this trial, site is still the most significant factor influencing growth and survival; there were also significant differences among prov-enances, while the effect of family within provenance was scarce. The estimated dif-ference in biomass production between the best and the worst performing provenance at age ten was as high as 145 % for the harsher site, highlighting the importance of an adequate selection of the seed source used for afforestation (Climent et al. 2009).

In France, a “new” five-site comparative trial was settled between 1995 and 1997 within the Mediterranean Pine and Cedar (MPC) European project and the FORADAPT INCO project (Plomion and Pichot 2001; Fig. 5.9; Table 5.4). Eighteen Pinus brutia provenances from Turkey were planted in this trial along with four P. halepensis populations to be used as controls (Table 5.5). An additional set of seven P. brutia provenances commonly used by the French forest service were also included in Bousquet d’Orb and Naucadery plantations. These plantations share some provenances with trials established within the same project by Moroccan and Tunisian partners.

Fig. 5.8 Provenance/progeny trial with 9 years of age at a marginal site for the species (low winter temperatures, low rainfall and gypsum marls soil) in Central Spain, Valladolid province (R. López)

306

307

308

309

310

311

312

313

314

315

316

317

318

319

320

321

322

323

324

325

326

327

M.R. Chambel et al.

Fig. 5.9 Location of provenances and experimental sites included in the ‘new’ French brutia pine provenance trial. Black triangles represent trial sites (5); orange dots (18) represent Pinus brutia provenances and red dots represent P. halepensis provenances used as control (4)

Table 5.4 List of the “new” French Pinus brutia provenance plantations

Site Year plant Altitude Surface (ha) SoilNumber of provenances

Bousquet d’Orb F 1995 580 2 Limestone 28Naucadery (Laure) S 1996 150 1.5 Limestone 27Toulourenc forest F 1996 600 2 Limestone 22Bedoin forest S 1997 600 2 Limestone 25Laquina S 1997 600 1.8 Schist 22

Table 5.5 List of the provenances tested in the “new” 5-site trial

Species Provenance Country Species Provenance Country

P. brutia 050 Karsanti Turkey P. brutia 116 Eskibag TurkeyP. brutia 071 Kiyra Turkey P. brutia 120 Karabucak TurkeyP. brutia 073 Suçati Turkey P. brutia 121 Güzelcluk TurkeyP. brutia 076 Guzelbag Turkey P. brutia 122 Bigadic TurkeyP. brutia 079 Pinargözü Turkey P. brutia 124 Camkonagi TurkeyP. brutia 100 Karadag Turkey P. brutia 125 Göktepe TurkeyP. brutia 109 Gökçesu Turkey P. brutia 129 Koças TurkeyP. brutia 110 Findikpinari Turkey P. halepensis 001 St Etienne du Grès FranceP. brutia 113 Melli Turkey P. halepensis 002 Port-Cros FranceP. brutia 114 Merkez Turkey P. halepensis 003 Mejjou MoroccoP. brutia 115 Karaçay Turkey P. halepensis 011 Vilmorin France

t4.1

t4.2

t4.3

t4.4

t4.5

t4.6

t4.7

t4.8

t5.1

t5.2

t5.3

t5.4

t5.5

t5.6

t5.7

t5.8

t5.9

t5.10

t5.11

t5.12

t5.13


5.4.2 Provenance Research in Other Countries

In Greece, a network of provenance trials has been established for both P. halepensis (Fig. 5.10a) and P. brutia (Fig. 5.10b), to explore the levels of the existing variation for adaptive traits. This network consisted of five trial sites for Pinus halepensis and seven trial sites for Pinus brutia including both Greek and foreign materials. These were, for P. halepensis: three Italian, one from Spain, three Moroccan, five Tunisian and six other provenances form North African countries and, for Pinus brutia: two Turkish provenances and one from Cyprus.

In Morocco, provenance research on the species of the halepensis group was very active during the second half of the twentieth century, when there were a large number of trials. A five-site Pinus brutia provenance test was planted in 1969; another trial was installed in only one location in 1973–1974 with seven provenances of P. halepen-sis and P. brutia; and in 1976–1977 a four-site provenance trial within the FAO/Silva Mediterranea network including 51 provenances was installed. Unfortunately, most of these plantations do not exist anymore and little information on the results is easily available (Pichot 1995). In 1992 a new two-site provenance test was planted with 58 provenances (1 Eldarica pine, 8 Brutia pine and 49 Aleppo pine). In 1999, 18 P. brutia provenances and 1 P. halepensis were planted in a two-site trial located in the Moroccan dry area, implemented within the EU Mediterranean INCO-FORADAPT project.

In Turkey, a wide programme of provenance research was carried out by the Turkish Forest Research Institute, with a set of 26 trial sites of Brutia pine planted in 1988 and 1989 throughout Turkey and Northern Cyprus, with 47 provenances from Turkey and 3 from Cyprus, mostly from selected stands.

In Tunisia, Aleppo pine populations cover 200,000 ha and P. halepensis is the first species for afforestation (80,000 ha). The studies on genetic diversity of both Aleppo and Brutia pines are based on two sets of provenance trials, a seven-site trial estab-lished in the 1960s with 49 provenances, and a four-site trial planted in 1998, focusing on drought tolerance, implemented within the EU Mediterranean INCO-FORADAPT project and including 13 provenances of Aleppo pine and 27 of Brutia pine.

5.4.3 Geographic Patterns of Genetic Diversity

5.4.3.1 Biochemical and Molecular Variation

Discrimination between species and their hybrids was carried out by terpene and spSSR analyses (Gallis et al. 1997; Korol et al. 1995; Tognetti et al. 1997; Bucci et al. 1998).

Studies on Aleppo and Brutia pine genetic diversity and population genetics based on isozymes (Schiller et al. 1986; Conkle et al. 1988; Teisseire et al. 1995, Agúndez et al. 1997, 1999; Korol et al. 2002; Wahid et al. 2010), RAPDs and cpSSR (Bucci et al. 1998; Gomez et al. 2001; Alrababah et al. 2011) generally indicate lower levels of intra-population diversity for Aleppo pine and the existence of genetic barriers for introgression between both species.

328

329

330

331

332

333

334

335

336

337

338

339

340

341

342

343

344

345

346

347

348

349

350

351

352

353

354

355

356

357

358

359

360

361

362

363

364

365

366

367

M.R. Chambel et al.

Fig. 5.10 Location of provenances and trial sites of the Pinus halepensis network (a) and of the Pinus brutia network (b) in Greece. Yellow stars represent trial sites and trees represent prove-nances (P. Alizoti)


The genetic variability of Pinus halepensis has a geographic structure that reflects migration pathways from glacial remnants after the last glaciations (Gómez et al. 2005). Recent results confirmed a marked loss of genetic diversity from Greek populations towards the western range of the species, as well as molecular sig-natures of intense bottlenecks (Grivet et al. 2009). Genetic analyses on P. brutia indicated a high differentiation between populations but with no clear geographic structure (Bucci et al. 1998).

5.4.3.2 Quantitative and Adaptive Variation

The previously mentioned trial networks and smaller-scale eco-physiological research give evidence of both an adaptive divergence between the two species and differentia-tion among provenances within species. A joint analysis of all extant data from different trial sets is still lacking, nevertheless, there are several published works regarding growth, mortality and frost resistance (Calamassi et al. 1984); needle morphology and anatomy (Calamassi et al. 1980, 1987); drought tolerance (Calamassi et al. 1980; Falusi et al. 1983; Grunwald and Schiller 1988; Atzmon et al. 2004; Voltas et al. 2008), seed and cone morphology (Boulii et al. 2001, 2003) and pest resistance, stem form and forking (Eccher et al. 1987; Weinstein 1989; Bariteau 1992; Ducci and Guidi 1998). The results obtained confirm that Pinus halepensis and Pinus brutia have different ecological optima. Pinus brutia is less tolerant to dry environments, being out-performed by most P. halepensis provenances at drier sites, but on the other hand, it is less frost sensitive, therefore better adapted to continental areas and high altitudes. Pinus brutia generally presents better form and it has also been proven to be more resistant to the insect Pissodes notatus (Ducci and Guidi 1998).

In general, Aleppo pine tends to show a higher differentiation between prove-nances than Brutia pine, but this varies considerably among studies due to prove-nance sampling and test environments.

In Aleppo pine, the north-east–south-west cline for adaptive characters seems very well established experimentally, with north-eastern provenances (Greece) showing higher early growth rates and better forms than south Iberian and north African provenances. Recently, a similar geographic trend for early reproductive allocation has also been confirmed, with eastern and northern populations (particu-larly Greece, Catalonia and Balearic Islands) far less precocious for cone bearing than South Iberian and North African provenances (Climent et al. 2008).

For Brutia pine, despite its smaller range of distribution, there are also experi-mental evidences of differences between populations and altitudinal gradients for adaptive traits (i.e.: different resistance to cold and developmental rates, Isik 1986; Kaya and Isik 1997; Pichot and Vauthier 2007). Provenances from the middle eleva-tion of the Mediterranean region generally perform the best while provenances from the peripheral areas of the distribution range of the species or isolated populations generally grow more slowly. However, other reports postulate scarcely distinct geographic patterns of diversity in this species as a consequence of human influence and recurrent fires (Gülcü and Çelik 2009).

368

369

370

371

372

373

374

375

376

377

378

379

380

381

382

383

384

385

386

387

388

389

390

391

392

393

394

395

396

397

398

399

400

401

402

403

404

405

406

407

408

409

M.R. Chambel et al.

Besides this between-population variation, high intra-population diversity (both through heritability and/or additive co-efficient of variation) has been recorded in Aleppo pine for growth (Matziris 2000) and particularly for reproductive traits (Matziris 1997; Santos-del-Blanco et al. 2010). This is also true for Brutia pine, with very similar heritability values for the different sets of traits (Gülcü and Çelik 2009; Isik et al. 2002; Kaya and Isik 1997).

In general, the formerly mentioned trial networks and smaller-scale eco-physio-logical research (Grunwald and Schiller 1988) give evidence for both an adaptive divergence between the two species and differentiation among provenances within. The results obtained in different countries confirm that Pinus halepensis and Pinus brutia have different ecological optima species (Bariteau 1992; Ducci and Guidi 1998; Boulii et al. 2001, 2003). Pinus brutia is less tolerant to dry environments, being out-performed by most P. halepensis provenances at more Mediterranean sites, but on the other hand, it is less frost sensitive, presenting higher adaptability to continental areas and high altitudes. Pinus brutia generally presents best stem form and tree architecture and it has also proven to have better resistance to the insect Pissodes notatus (Ducci and Guidi op. cit.).

Aleppo pine tends to show a high inter-provenance variability, while Brutia pine tends to show lower differentiation between provenances, but the latter varies con-siderably among studies (provenance sampling and environments). A joint analysis of all extant data from different trial sets is still lacking. Therefore, the following sections will quote the most consistent results and then some more details on the results obtained in each country up until now.

5.4.4 Seed and Plant Transfer: History and Current Trends

Today, in Italy, France and Spain, seed imports from Turkey are allowed for Pinus brutia. In Italy, imports have been programmed with reference to indications issued from FAO Silva Mediterranean field tests. At present, most needs can be covered by a seed orchard derived from these programmes.

By contrast, Aleppo pine seeds must come from the national provenance regions and seed stands in the three countries (Fig. 5.11). In France, however, significant quantities of Aleppo pine seeds were imported in the past from some foreign countries for historical afforestation programmes in the South of France. This reproduc-tive material often came from stands exhibiting warmer climatic conditions than those observed in France and its susceptibility to frost led to high mortality during the coldest winters (1956 and 1985) within forest stands planted respectively with Algerian and Italian provenances (Tabeaud and Simon 1993; Bedel 1986).

The situation was different in Italy and Spain, where the importation of foreign Aleppo pine seeds, if any, can be considered negligible due to the high production of seeds from local basic materials. Traceability of former seed transfer between regions within Spain is unrealistic, so both successful and unsuccessful plantations outwith the natural range are difficult to trace in terms of seed origin.

[AU3]

410

411

412

413

414

415

416

417

418

419

420

421

422

423

424

425

426

427

428

429

430

431

432

433

434

435

436

437

438

439

440

441

442

443

444

445

446

447

448

449

450


In Italy, wide forestation programmes were carried out in Tuscany for coastal dunes and abandoned interior and Mediterranean areas. This work was initiated by the Dukes of Tuscany before 1860 and then continued by the Kingdom of Italy after the Unification. The amount of Aleppo pine seed distributed documented by the Italian Forest Service was nearly 2,000 kg in the early 1960s, reaching about 20,000 kg in the early 1970s with a peak in 1989 when about 40,000 kg were pro-duced. These amounts were estimated to cover 60 % of the Aleppo pine seeds used at national level. Thereafter, production decreased progressively and today only a few hundred kilos are being produced and only a proportion are sold. That is due to the end of forestation programmes in the early 1990s (Mencaccini 2010).

5.4.5 Approved Forest Reproductive Material Within the TreeBreedex Countries

A total of 54 seed stands registered according to rules under the European Directive 1999/105/CE are officially approved in the three TreeBreedex countries for the pro-duction of Pinus halepensis seeds (Table 5.6).

In Spain there are 14 selected stands, belonging to 7 out of the 20 provenance regions defined for the species (Fig. 5.12). These selected stands cover a total of 495 ha. All except one were homologated in February 2002 (the remaining one was homologated in July 2009). There is also one clonal seed orchard which was homologated in July 2007 with material selected from three provenance regions. The selection of parent material for this seed orchard was based on growth and form. The seed from this seed orchard is currently catalogued as qualified FRM, but

Fig. 5.11 Aleppo pine seeds clean and ready for the market (1966, courtesy of the Italian Forest Service)

451

452

453

454

455

456

457

458

459

460

461

462

463

464

465

466

467

468

469

470

471

472

M.R. Chambel et al.

there are already comparative trials which have been planted with the aim of raising the category up to tested FRM.

In France, there are 25 selected stands for the production of Pinus halepensis seed, totalling 228 ha, and all of them are included in the only provenance region defined for the species (Fig. 5.13) with all having been homologated in November 2003.

In Italy, there are 15 selected seed stands, broadly covering the national range of the species (Fig. 5.14). All of these seed stands were homologated before 1975 (Morandini and Magini 1975; Ducci et al. 2010).

Table 5.6 Homologated FRMs in TreeBreedex countries

Country Selected Qualified Tested

Spain Number 14 selected seed stands 1 seed orchard (P. halepensis) –Area 495 ha 4.31 ha –

France Number 25 selected seed stands – –Area 228 ha – –

Italy Number 15 selected seed stands 1 seed orchard (P. brutia) –Area About 2,600 ha 2.5 ha –

Fig. 5.12 Regions of provenance defined in Spain for Aleppo pine: 1. Alta Cataluña, 2. Cataluña Litoral, 3. Cataluña Interior, 4. Bárdenas-Ribagorza, 5. Ibérico Aragonés, 6. Monearos-Depresión del Ebro, 7. Alcarria, 8. La Mancha, 9. Maestrazgo-Los Serranos, 10. Levante Interior, 11. Litoral Levantino, 12. Pitiusas, 13. Sudeste, 14. Bética, 15. Bética Meridional, 16. Cazorla, 17. Sur, 18. Mallorca y Menorca, 19. Repoblaciones de la Meseta Norte. Region 19, in blue, consists of planta-tions outwith the natural range of the species

473

474

475

476

477

478

479

480

481

t6.1

t6.2

t6.3

t6.4

t6.5

t6.6

t6.7

t6.8


Fig. 5.13 Regions of provenance defined in France for Aleppo pine: a single provenance region named PHA 700: Region Méditerranéenne

Fig. 5.14 (a) The range of Pinus halepensis in Italy and the location of selected seed stands: 125 – Vallecrosia (Imperia-Liguria), 110 – Chiavari-Le Grazie (Genova-Liguria), 15 –Cerchiara (Terni-Umbria), 18 – Arrone (Terni-Umbria), 19 – Otricoli (Terni-Umbria), 60 – Gargano-Marzini (Foggia-Puglia), 78 – Gargano-Monte Pucci (Foggia-Puglia), 59 – Patemisco (Taranto-Puglia), 52, 53, 54, 55, 56, 57, 58 – Litorale Tarantino (Taranto-Puglia). Capital letters represent FAO trial sites (see Table 5.2). (b) The provisional official map of provenance regions in Italy (Partitionist method 2010) agreed by all the Regional and State Forest Services. A – Alpine Region, B – Planitial Po valley Region, C – Central–Northern Mediterranean Region, D – Southern Mediterranean region, E – Sardinian (Mediterranean) Region, F – Sicily (Mediterranean) Region

M.R. Chambel et al.

There are no specific provenance regions defined for P. brutia within TreeBreedex countries, but a seed orchard of this species was recently registered in Italy (Ducci et al. 2010).

1. Among all the TreeBreedex countries, 70 Aleppo pine and 77 of Brutia pine selected seed stands were inventoried by FAO Silva Mediterranea (Topak 1997). Seed orchards recorded were 4 for Pinus halepensis, 55 for Brutia pine and 2 for Pinus eldarica, The majority are located in Cyprus (6) and Turkey (49) and only one in Egypt.

5.4.6 Web Links to Approved FRMs in TreeBreedex Countries

Spainhttp://www.mma.es/portal/secciones/biodiversidad/montes_politica_forestal/recur-sos_geneticos_forestal/programas_mejora_genetica/catalogo_materiales_base/index.htmhttps://sgw.mma.es/ (this page can only be access by authorized users)

Francehttp://agriculture.gouv.fr/graines-et-plants-forestiershttp://agriculture.gouv.fr/IMG/pdf/pin_alep-2.pdf

Italyhttp://www.infc.ithttp://www.ricercaforestale.it/http://www.ricercaforestale.it/modules.php?op=modload&name=BoschiDaSeme&file=index

5.5 Current Breeding Programmes, Description and Results

5.5.1 Breeding Within TreeBreedex Countries

In Spain, a seed orchard programme was carried out in the 1980s and 1990s, from a selection of plus trees in Eastern continental Spain (three regions of provenance). Progeny trials were established, both with open-pollinated seeds from the selected mother trees and, more recently, with the seeds collected from seed orchards. A more recent set of comparative trials was planted to promote the FRM of the most successful seed orchard to the tested category (Nieto et al. 2009), even though the demand for improved seed of this species in Spain (justifying a higher price) is very low. Thanks to the precocious and prolific cone-bearing in seedlings, progeny tests can easily be converted into seedling seed orchards of 1.5 generations.

In France, controlled pollinations inter- and intra-species and provenances were done in order to study the relationships and genetic control of main adaptive traits (survival, growth, phenology, architecture and fertility) within the halepensis group.

482

483

484

485

486

487

488

489

490

491

492

493

494

495

496

497

498

499

500

501

502

503

504

505

506

507

508

509

510

511

512

513

514

515

516

http://www.mma.es/portal/secciones/biodiversidad/montes_politica_forestal/recursos_geneticos_forestal/programas_mejora_genetica/catalogo_materiales_base/index.htm



https://sgw.mma.es/

http://agriculture.gouv.fr/graines-et-plants-forestiers

http://agriculture.gouv.fr/IMG/pdf/pin_alep-2.pdf

http://www.infc.it

http://www.ricercaforestale.it/

http://www.ricercaforestale.it/modules.php?op=modload&name=BoschiDaSeme&file=index

http://www.ricercaforestale.it/modules.php?op=modload&name=BoschiDaSeme&file=index


Five seed trees within each of four provenances (two of P. brutia: Baspinar and Marmaris, and two of P. halepensis: Cehegin and Gémenos) were pollinated with mixtures of pollen collected from ten pollen trees within each of the four prove-nances. The seeds produced made it possible to plant an experimental trial in 2000, in the South of France, near Perpignan. Preliminary results reveal significant differ-ences among families for survival but no species effect. Pinus halepensis families grow faster than P. brutia and P. brutia × P. halepensis hybrids are significantly more vigorous than pure P. brutia trees. In order to estimate genetic parameters in P. brutia, one provenance/progeny test with 90 half-sib families (3 stands × 30 families) was planted in southern France in 2000.

In Italy, two progeny trials with half-sib families of the best performing prove-nances from Greece and Italy, selected by stem form, were established in 1985, covering 4.5 ha.

5.5.2 Breeding in Other Countries

Long-lasting breeding programmes of both Aleppo and Brutia pines have been car-ried out in other countries outwith the TreeBreedex consortium.

In Greece, the initial breeding effort that was launched in 1962 mainly involved hybridization efforts between Pinus brutia and Pinus halepensis. Organized breed-ing programmes were initiated in 1970, with selection and collection of material on a provenance and mother-tree basis. Both P. halepensis and P. brutia provenances exhibited significant genetic variation for all the adaptive traits studied, while their performances across sites indicated the high potential for effective across-site selec-tion (Matziris 1997; Alizoti et al. 2000). P. brutia provenances of Greek origin out-grew the provenances originating from Turkey and Cyprus when tested in two different sites (Alizoti et al. 2001). The outstanding performance of P. brutia × P. halepensis hybrids is worth mentioning, as when tested together with their parental species in harsh environmental conditions they significantly outperformed the only parent that was able to survive (P. brutia). The results indicate the existence of ample genetic variation among and within the natural populations of both species for adaptive traits (Matziris 2000), the potential for selection and breeding and the need for conservation of their unique genetic resources in the face of global change. A first-generation seed orchard exists for Pinus halepensis covering 20 ha and including 76 genotypes (2,630 grafted individuals in total). There are two progeny trials associated with this seed orchard, established in 1987, with 70 open-pollinated families that correspond to 70 out of the 76 clones from the seed orchard.

In Israel, a three-site provenance trial with provenances of Pinus halepensis Mill. and related species, P. brutia Ten. and P. eldarica Medw. was established within the framework of the FAO/Silva Mediterranea programme FAO/SCM/CRFM/4 bis, as for the other partners of Silva Mediterranea. The results obtained 10 years after planting were discussed by Weinstein (1989). According to these results, the most promising provenances of P. halepensis are from the lower altitudes of Greece and

517

518

519

520

521

522

523

524

525

526

527

528

529

530

531

532

533

534

535

536

537

538

539

540

541

542

543

544

545

546

547

548

549

550

551

552

553

554

555

556

557

M.R. Chambel et al.

some native stands within Israel, while the most promising seed sources of P. brutia are from the lower elevation on the Mediterranean coast of Turkey. In addition, P. eldarica from Iran showed good growth potential for Israel. Research projects continued within the Aleppo pine forest plantations and new provenance trials at Yatir, near the northern edge of the Israeli Negev desert (Schiller and Atzmon 2009), were established, with the aim of improving the sustainability of plantations with this species (Fig. 5.15).

These aims imply that seed sources to be used (provenances, ecotypes) must be selected according to their genetic diversity, drought resistance and water use efficiency. Harsh environmental site conditions at the edge of the desert provide the opportunity of exerting heavy selection pressure among and within provenances of a priori drought-tolerant species. Survivors should be used as a seed source for new forest plantations aiming to combat desertification.

In Morocco, as already mentioned, intense provenance research for both Aleppo and Brutia pine was undertaken in the second half of the twentieth century, includ-ing several trials based on the Silva Mediterranea initiative, and a P. halepensis seed orchard was established in 1978.

In Tunisia, as well as the provenance trial sets of Aleppo and Brutia pines planted since the 1960s, a P. brutia provenance/progeny test was planted in 2000 with 90 half-sib families (three stands × 30 families).

In Turkey, again in parallel with wide provenance research formerly described for Brutia and Aleppo pines, a wide programme of seed orchard plantation has been carried out. As many as 67 seed orchards of Brutia pine covering 472 ha and 2 seed

Fig. 5.15 The Yatir (edges of Negev desert) plantations (G. Schiller)

558

559

560

561

562

563

564

565

566

567

568

569

570

571

572

573

574

575

576

577

578

579

580


orchards of Aleppo pine totalling 8.2 ha were established. Progeny testing has also been regularly undertaken and has been the object of intense research (Kaya and Isik 1997; Isik et al. 2002; Fig. 5.16).

5.6 Additional Factors Affecting Future Breeding of Aleppo and Brutia Pines

5.6.1 Aleppo and Brutia Pines and Climate Change

Past predictions of enhanced growth in Mediterranean countries due to the fertilizing effect of increased CO2 (Sabaté et al. 2002) are largely surpassed today because of the crucial role of water availability (Scarascia-Mugnozza 1986; Schroter et al. 2005). Nevertheless, both experimental evidences and predictions of the effects of climate change on Aleppo and Brutia pine growth and distribution are highly diver-gent, depending on the particular region.

In northern Mediterranean regions, such as southern France and north-eastern Spain, growth increase and/or enlargement of the potential area for the species have been postulated, following temperature rising and increased CO2 (Rathgeber et al. 2000; Thuiller 2003). A putative decrease of total and summer rainfall (Fig. 5.17) would favour Aleppo pine’s spreading and colonizing potential at the expense of other more mesic species, such as evergreen oaks (Zavala and Zea 2004).

By contrast, a marked decline in growth has been demonstrated by dendro-chronological methods on the edge of distributions, such as very dry coastal areas of south-eastern Spain for Aleppo pine (De Luis et al. 2007) or some Greek Islands for Brutia pine (Körner et al. 2005; Sarris et al. 2007). In these cases, extreme drought will put some populations at risk of extinction. One main aim of future

[AU4]

Fig. 5.16 Dr. Murat Alan follows Brutia pine progeny tests in Turkey (Ducci et al. 2010)

581

582

583

584

585

586

587

588

589

590

591

592

593

594

595

596

597

598

599

600

601

602

603

M.R. Chambel et al.

breeding programmes is to evaluate and select provenances and genotypes adapted to drought and efficient for water and nutrients use. In recent decades, innovative methods and new indices have been introduced to quantify the impact of stress on carbon gas exchanges between plants and the atmosphere and the efficiency of uti-lization of light and water resources (Farquhar et al. 1989; Tognetti et al. 1997; Scarascia-Mugnozza et al. 2011; Table 5.7). Good examples of physiologically-based selection of drought-tolerant genetic materials of Aleppo pine have been made in recent decades in the Negev desert in Israel (Schiller 2011).

A less obvious outcome of climate change is the higher risk of frost damage in mountain or continental areas, derived from a lack of hardening and more sudden than normal temperature extremes accompanying global warming (Kozlowski and Pallardy 2002; Fernández et al. 2003). This effect can be more relevant for Aleppo pine, given its lower general tolerance to cold temperatures (Calamassi et al. 2001; Climent et al. 2009).

The increase in fire recurrence and virulence as experienced in recent decades over southern France (Fig. 5.18), Spain and Greece has been shown to favour pyro-phites like Aleppo and Brutia pines at the expense of less adapted species such as P. nigra or even P. pinaster (Tapias et al. 2001, 2004). However, even when some degree of fast adaptation through reducing the age or size needed for reproduction can be expected according to recent research (Santos-del-Blanco et al. 2010) where there is a short recurrence interval between forest fires the resilience of these species can be challenged (Rigolot 2004).

Fig. 5.17 Annual (a) and summer (b) changes in precipitation, expressed in % as simulated by ENSEMBLES regional climate models for the IPCC SRES A1B emission scenario (Source: DMI ensembles-eu.metoffice.com, www.nature.com. http://ensemblesrt3.dmi.dk/ENSEMBLE-MEAN_A1B_GCM_ MM_25km-CRU_pr.nc)

604

605

606

607

608

609

610

611

612

613

614

615

616

617

618

619

620

621

622

623

624

625

http://ensembles-eu.metoffice.com

http://www.nature.com

http://ensemblesrt3.dmi.dk/ENSEMBLE-MEAN_A1B_GCM_MM_25km-CRU_pr.nc

http://ensemblesrt3.dmi.dk/ENSEMBLE-MEAN_A1B_GCM_MM_25km-CRU_pr.nc

Table 5.7 Possible relationships between some physiological and growth-related traits and drought resistance among Mediterranean and related conifers (Scarascia-Mugnozza et al. 2011)

Traits Xeric ecotypes Mesic ecotypes References

Photosynthetic capacity

High Low P. taeda: Teskey et al. (1986)

Stomatal conductance

High Low P. halepensis: Tognetti et al. (1997)

Stomatal sensitivity

Low High P. halepensis: Tognetti et al. (1997)

Mesophyll conductance

High Low ?

Transpiration rate High Low P. halepensis: Tognetti et al. (1997)

Intrinsic water-use efficiency

Low High P. pinaster: Tognetti et al. (2000)

Hydraulic conductance

High Low P. halepensis: Tognetti et al. (1997)

Above ground juvenile growth

Slow Fast ?

Above ground adult growth

Fast Slow ?

Adult tree shape Vigorous Tall P. pinaster: Guehl et al. (1995)

Carbon isotope discrimination

High Low P. pinaster: Tognetti et al. (2000)

Drought adaptive strategy

Find water and use it

Save water

[AU5]

[AU6]

Fig. 5.18 Number of fires and burnt areas from 1980 to 2002 in France, Greece, Italy, Portugal and Spain (Source: European Commission, Topic Centre for Terrestrial Environment (ETC./TE). Forest fires in Europe – 2002 fire campaign)

t7.1

t7.2

t7.3

t7.4

t7.5

t7.6

t7.7

t7.8

t7.9

t7.10

t7.11

t7.12

t7.13

t7.14

t7.15

t7.16

t7.17

t7.18

t7.19

t7.20

t7.21

t7.22

t7.23

t7.24

t7.25

t7.26

t7.27

M.R. Chambel et al.

5.6.2 Vegetative Propagation and Somatic Embryogenesis

Somatic embryogenesis of both Aleppo pine and Brutia pine has been achieved suc-cessfully over the last decades (Diamantoglou and Banilas 1996; Lambardi et al. 1993; Scaltsoyiannes et al. 1994). However, this technique has been applied only at an experimental level, and not for breeding purposes.

Operational propagation has been carried out using cuttings and grafting, with a previous emphasis on the multiplication of P. brutia × P. halepensis hybrids (Panetsos et al. 1994). Interestingly, Aleppo and Brutia pines have been used as dwarfing rootstocks for enhancing cone production in heteroblastic grafts with scions of other species, like Pinus pinea or even Pinus nigra and P. sylvestris (Climent et al. 1997; Gil et al. 1990; Parra 1980). Both Aleppo and Brutia pine rootstocks induce a dwarfing effect and a neat increase of reproductive allocation, and can be specially indicated for ex-situ conservation clonal banks or seed orchards (Fig. 5.19).

5.6.3 Knowledge Gaps and Research Needs

Aleppo pine provides a very fine example of colonization from eastern to western Mediterranean and, as such, population genetics of this species are still under-inves-tigated. It is surprising how fast adaptive radiation at the western extreme of the

Fig. 5.19 Experimental heteroblastic grafting of black pine (Pinus nigra subsp. salzmannii) on Aleppo pine rootstocks (left) and Scots pine on Pinus brutia rootstocks at Alaquas and Puerta de Hierro breeding centres, Spain (Ministry of Environment), completely outwith the climatic range of these species (J. Climent)

626

627

628

629

630

631

632

633

634

635

636

637

638

639

640

641

642


natural distribution range has occurred despite a westward reduction in neutral genetic variation. Separating founder effects from local adaptation processes driven by climate, soil and perturbation regimes (mostly fires) is the object of ongoing research that should be reinforced in the near future.

A joint analysis of all extant data from different trial sets would be extremely useful, provided that advanced, suitably powerful statistical tools are used.

The knowledge on the vegetative and reproductive phenology of this species and the effects of climate change would be extremely enhanced if the genetic variation (both between and within provenances) and g × e interaction were considered (Girard et al. 2011); there are interesting data sets still under-exploited for this purpose. In all these aspects, integrating the fast progresses of molecular genetics will foster high-quality research with the potential application for any level of genetic and breeding activities.

5.6.4 European-Wide Breeding Perspectives

A collaborative, joint analysis of the extant field trial networks would form the basis for a putative European-wide breeding programme for Aleppo and Brutia pines. However, the interested countries would be different for the two species. From this review, it seems that breeding of Aleppo pine for wood production would be more rewarding for North African countries than for western European countries, whose wood production is more focused on other species. However, some interest could be found in increasing growth for maximizing biomass production (hence carbon capture) under limiting edapho-climatic conditions. Brutia pine could be a valuable resource for plantation programmes in areas where both moderate drought and frosts are combined limiting factors. Mixed plantations with broadleaves could enhance resilience and maximize carbon fixation in durable wood products for mitigation purposes.

Maintaining the adaptive capacity of Mediterranean forests through optimal deployment of the available genetic resources of these species, or enhancing some particular traits through artificial selection, already counts with a wide knowledge base which could be used in a collaborative way through actions such as FAO’s Silva Mediterranea or other support tools.

References

Agúndez D, Degen B, von Wuehlisch G, Alía R (1997) Genetic variation of Aleppo pine (Pinus halepensis Mill.) in Spain. For Genet 4:201–209

Agúndez D, Degen B, von Wuehlisch G, Alía R (1999) Multilocus analysis of Pinus halepensis Mill. from Spain: genetic diversity and clinical variation. Silvae Genetica 48:173–178

Alizoti PG, Panetsos KP, Moulalis DCh (2000) Genetic parameters and performance of Pinus halepensis Mill. genetic material. In: Proceedings of the 8th conference of the Hellenic Scientific Society of Plant Breeding, Arta, 23–25 Oct 2000, pp 37–49

643

644

645

646

647

648

649

650

651

652

653

654

655

656

657

658

659

660

661

662

663

664

665

666

667

668

669

670

671

672

673

674

675

676

677

678

679

680

681

M.R. Chambel et al.

Alizoti PG, Panetsos KP, Moulalis DC (2001) Genetic variation of Pinus brutia Ten. provenances and their evaluation for growth and adaptation. Sci Ann Dep For Nat Environ AUTh (M/1) 40:59–74

Alrababah M, Al-Horani A, Alhamad M, Migdadi H (2011) Genetic diversity of the easternmost fragmented Mediterranean Pinus halepensis Mill. populations. Plant Ecol 212(5):843–851. doi:10.1007/s11258-010-9872-0

Anonymous (2010) La Forêt Française (French forests). © Inventaire forestier national, 2010, 58pp

Atzmon N, Moshe Y, Schiller G (2004) Ecophysiological response to severe drought in Pinus halepensis Mill. trees of two provenances. Plant Ecol 171:15–22

Bacandritsos N, Saitanis C, Papanastasiou I (2004) Morphology and life cycle of Marchalina hel-lenica (Gennadius) (Hemiptera: Margarodidae) on pine (Parnis Mt.) and fir (Helmos Mt.) for-ests of Greece. Annals de la Société Entomologique de France 40:169–176

Bariteau M (1992) Geographic variation and stress adaptation of Pinus halepensis-Pinus brutia complex in Mediterranean conditions. Preliminary results of a provenance test in France. Annales des Sciences Forestières 49:261–276

Bedel J (1986) Aménagement et gestion des peuplements de pin d’Alep dans la zone Méditerranéenne Française (Management of Aleppo pine forests in the French Mediterranean area). In: Morandini R (ed) Le pin d’Alep et le pin Brutia dans la sylviculture méditerranéenne, vol 86/1. CIHEAM-IAMM, Montpellier, pp 109–125

Bellot J, Maestre F, Chirino E, Hernández N, de Urbina J (2004) Afforestation with Pinus halepensis reduces native shrub performance in a Mediterranean semiarid area. Acta Oecologica 25:7–15

Boulii A, Baazis M, M’Hirit O (2001) Polymorphism of natural populations of Pinus halepensis Mill. in Morocco as revealed by morphological characters. Euphytica 119:309–316

Boulii A, Pichot C, Baaziz M, M’Hirit O (2003) Variabilité morphologique des graines de cinq populations marocaines de pin d’Alep (Pinus halepensis Mill) (Seed morphologic variability in five Moroccan populations of Aleppo pine (Pinus halepensis Mill) from Morocco). Naturalia Maroccana 1:161–163

Bucci G, Anzidei M, Madaghiele A, Vendramin GG (1998) Detection of haplotypic variation and natural hybridization in halepensis-complex pine species using chloroplast simple sequence repeat (SSR) markers. Mol Ecol 7:1633–1643

Calamassi R, Falusi M, Tocci A (1980) Variazione geografica e resistenza a stress idrici in semi di Pinus halepensis Mill., Pinus brutia Ten. e Pinus eldarica Medw. (Geographic variation and resistance to water stress in seeds of Pinus halepensis Mill., Pinus brutia Ten. and Pinus elder-ica Medw.). Annali dell’Istituto Sperimentale per la Selvicoltura 11:193–230

Calamassi R, Falusi M, Tocci A (1984) Effets de la température de germination et de la stratification sur la germination des semences de Pinus halepensis Mill (Effects of germination temperature and stratification on the germination of Pinus halepensis seeds). Silvae Genetica 33:133–139

Calamassi R, Falusi M, Mugnai L (1987) Shoot morphology and growth pattern in seedlings of Pinus brutia provenances. Can J For Res 18:188–194

Calamassi R, Paoletti E, Strati S (2001) Frost hardening and resistance in three Aleppo pine (Pinus halepensis Mill.) provenances. Isr J Plant Sci 49:179–186

Chambel MR, Climent J, Voltas J, Sanchez de Ron D, Calama R, Alia R (2007) Aleppo pine prov-enance trial in Spain. FAO-Silva Mediterranea and IUFRO WG 20213 expert joint meeting: promotion and use of results from the international trials of Mediterranean Conifers, Arezzo and Rome, 21–23 June 2007

Climent J, Prada MA, Gil L, Pardos JA (1997) Increase of flowering in Pinus nigra Arn. subsp. salzmanni (Dunal) Franco by means of heteroplastic grafts. Annales des Sciences Forestières 54:145–153

Climent J, Prada MA, Calama R, Sánchez de Ron D, Chambel MR, Alía R (2008) To grow or to seed: ecotypic variation in reproductive allocation and cone production by young female Aleppo pine (Pinus halepensis, Pinaceae). Am J Bot 94:1316–1320

Climent MJ, Chambel MR, del Caño Prieto F, López Rodríguez R, Alia Miranda R (2009) Adaptación de procedencias de Pinus halepensis en las margas del Valle del Duero: crecimiento,

[AU7]

682

683

684

685

686

687

688

689

690

691

692

693

694

695

696

697

698

699

700

701

702

703

704

705

706

707

708

709

710

711

712

713

714

715

716

717

718

719

720

721

722

723

724

725

726

727

728

729

730

731

732

733

734

735

http://dx.doi.org/10.1007/s11258-010-9872-0


supervivencia y caracteres reproductivos (Adaptation of Pinus halepensis provenances to the Duero valley margins: growth, survival and reproductive characters). In: Montes y sociedad: saber que hacer. Actas del 5º Congreso Forestal Español, Avila, 21–25 Sept 2009

Climent J, San-Martín R, Chambel MR, Mutke S (2011) Ontogenetic differentiation between Mediterranean and Eurasian pines (sect. Pinus) at the seedling stage. Tree Struct Funct 25:175–186

Conkle M, Schiller G, Grunwald C (1988) Electrophoretic analysis of diversity and phylogeny of Pinus brutia and closely related taxa. Syst Bot 13:411–424

Daskalakou EN, Thanos CA (1996) Aleppo pine (Pinus halepensis) postfire regeneration: the role of canopy and soil seed banks. Int J Wildland Fire 6:59–66

De Luis M, Gricar J, Cufar K, Raventós J (2007) Seasonal dynamics of wood formation in Pinus halepensis from dry and semi-arid ecosystems in Spain. IAWA J 28:389–404

Diamantoglou S, Banilas GP (1996) II.4 Pinus pinea L. (Stone Pine) and Pinus halepensis Mill. (Aleppo Pine). In: Bajaj YPS (ed) Biotechnology in agriculture and forestry, trees IV. Springer, Berlin/Heidelberg/New York, pp 389–406

Ducci F, Guidi G (1998) I pini della sez. halepensis, selezione e possibilità di impiego di specie e provenienze per l’Italia (The Pines of the Aleppo pine section: selection and use of species and provenances in Italy). Annali dell’Istituto Sperimentale per la Selvicoltura 29:58–67

Ducci F, Vannuccini M, Germani A, Carone G (2010) Cap. 3.3, Regione Toscana (Tuscany Region). In: Ducci F, De Rogatis A (eds) Risorse Genetiche Forestali in Italia (Forest genetic resources in Italy). CRA SEL, Arezzo, pp 32–34

Eccher A, Fusaro E, Pelleri F (1987) Résultats de l’expérimentation italienne sur les principales provenances de pins de la section halepensis dix ans après la plantation (Results from the Italian research on the main provenances of pines of the halepensis section ten years after plant-ing). Forêt Méditerranéenne 9:5–14

Fady B, Semerci H, Vendramin G (2003) EUFORGEN technical guidelines for genetic conserva-tion and use for Aleppo pine (Pinus halepensis) and Brutia pine (Pinus brutia). International Plant Genetic Resources Institute, Rome

Falusi M, Calamassi R, Tocci A (1983) Sensitivity of seed germination and seedling root growth to moisture stress in four provenances of Pinus halepensis Mill. Silvae Genetica 32:4–9

Farquhar GD, Ehleringer JR, Hubick KT (1989) Carbon isotope discrimination and photosynthe-sis. Annu Rev Plant Physiol Plant Mol Biol 40:503–537

Fernandez M, Royo A, Gil L, Pardos JA (2003) Effects of temperature on growth and stress hard-ening development of phytotron-grown seedlings of Aleppo pine (Pinus halepensis Mill.). Ann For Sci 60:277–284

Fisher J, Neumann R, Mexal J (1986) Performance of Pinus halepensis/brutia group pines in southern New Mexico. For Ecol Manage 16:403–410

Frankis M (1999) Pinus brutia (Pinaceae). Curtis’s Bot Mag 16:173–184Gallis AT, Lang KJ, Panetsos KP (1997) Bud monoterpene composition in Pinus brutia (Ten.),

Pinus halepensis (Mill.) and their hybrids. Silvae Genetica 47:71–74Gernandt DS, Lopez GG, Garcia SO, Liston A (2005) Phylogeny and classification of Pinus.

Taxon 54:29–42Gernandt DS, Magallon S, Lopez GG, Flores OZ, Willyard A, Liston A (2008) Use of simultane-

ous analyses to guide fossil-based calibrations of Pinaceae phylogeny. Int J Plant Sci 169:1086–1099

Gil L, Gordo J, Alia R, Catalán G, Pardos JA (1990) La mejora genética del pino piñonero (Breeding of Mediterranean Stone Pine). Montes 21:4–12

Gil L, Díaz-Fernández P, Jiménez MP, Roldán M, Alía R, Agúndez D, De Miguel J, Martín S, De Tuero M (1996) Las regiones de procedencia de Pinus halepensis Mill. en España (Regions of Provenance for Pinus halepensis Mill. in Spain). Organismo Autónomo Parques Nacionales, Madrid, 31pp

Giordano G (1976) Tecnologia del Legno. III. Le prove ed i legnami di più frequente impiego (Wood Tecchnology. III. Tests and the most frequently used woods). Unione Tipografico. Editrice Torinese, Turin, 1654pp

736

737

738

739

740

741

742

743

744

745

746

747

748

749

750

751

752

753

754

755

756

757

758

759

760

761

762

763

764

765

766

767

768

769

770

771

772

773

774

775

776

777

778

779

780

781

782

783

784

785

786

787

788

789

M.R. Chambel et al.

Girard F, Vennetier M, Ouarmim S, Caraglio Y, Misson L (2011) Polycyclism, a fundamental tree growth process, decline with recent climate change: the example of Pinus halepensis Mill. in Mediterranean France. Trees Struct Funct 25(2):311–322. doi:10.1007/s00468-010-0507-9

Gomez A, Alia R, Bueno MA (2001) Genetic diversity of Pinus halepensis Mill. populations detected by RAPD loci. Ann For Sci 58:869–875

Gómez A, Vendramin GG, González-Martínez SC, Alía R (2005) Genetic diversity and differentia-tion of two Mediterranean pines (Piuns halepensis Mill. and Pinus pinaster Ait.) along a latitu-dinal cline using chloroplast microsatelitte markers. Biodivers Res 11:257–263

Grivet D, Sebastiani F, González Martínez S, Vendramin G (2009) Patterns of polymorphism resulting from long range colonization in the Mediterranean conifer Aleppo pine. New Phytol 184:1016–1028

Grunwald C, Schiller G (1988) Needle xylem water potential and water saturation deficit in prov-enances of Pinus halepensis Mill. and P. brutia Ten. Forêt Méditerranéenne 10:407–414

Gülcü S, Çelik S (2009) Genetic variation in Pinus brutia Ten. seed stands and seed orchards for growth, stem form and crown characteristics. Afr J Biotechnol 8:4387–4394

Isik K (1986) Altitudinal variation in Pinus brutia Ten.: seed and seedling characteristics. Silvae Genetica 35:58–67

Isik F, Isik K, Yildrim T, Li B (2002) Annual shoot growth components related to growth of Pinus brutia. Tree Physiol 22:51–58

Kaya Z, Isik F (1997) The pattern of genetic variation in shoot growth of Pinus brutia Ten. popula-tions sampled from the Toros mountains in Turkey. Silvae Genetica 46:73–81

Keeley J, Zedler P (1998) Life history evolution in pines. In: Richardson DM (ed) Ecology and biogeography of Pinus. Cambridge University Press, Cambridge, pp 219–242

Körner C, Sarris D, Christodoulakis D (2005) Long-term increase in climatic dryness in the East-Mediterranean as evidenced for the island of Samos. Reg Environ Change 5:27–36

Korol L, Madmony A, Riov Y, Schiller G (1995) Pinus halepensis × Pinus brutia subsp. brutia hybrids? Identification using morphological and biochemical traits. Sivae Genetica 46:155–161

Korol L, Shklar E, Schiller G (2002) Diversity among circum-Mediterranean populations of Aleppo pine and differentiation from Brutia pine in their isoenzymes: additional results. Silvae Genetica 51:35–41

Kozlowski TT, Pallardy SG (2002) Acclimation and adaptive responses of woody plants to envi-ronmental stresses. Bot Rev 68:270–334

Lambardi M, Sharma K, Thorpe T (1993) Optimization of in vitro bud induction and plantlet for-mation from mature embryos of Aleppo pine (Pinus halepensis Mill.). In Vitro Cell Dev Biol Plant 29:189–199

López GG, Kamiya K, Harada K (2002) Phylogenetic relationships of Diploxylon pines (Subgenus Pinus) based on plastid sequence data. Int J Plant Sci 163:737–747

Maestre F, Cortina J, Bautista S, Bellot J (2003) Does Pinus halepensis facilitate the establishment of shrubs in Mediterranean semi-arid afforestations? For Ecol Manage 176:147–160

Magini E (1955) Pinete di Pino d’Aleppo (Pinewoods of Aleppo Pines). In: Atti del Congresso Nazionale di Selvicoltura, Firenze, 14–18 Mar 1954, vol 1, pp 49–69

Matziris D (1997) Variation in growth, flowering and cone production in a clonal seed orchard of Aleppo pine grown in Greece. Silvae Genetica 46:224–228

Matziris DI (2000) Genetic variation and realized genetic gain from Aleppo pine tree improve-ment. Silvae Genetica 49:5–10

Mencaccini S (2010) L’attivita’ dello stabilimento semiforestalie del vivaio forestale “Alto Tevere” nell’ambito della legge sulla montagna (The activity of the Seed Centre and Forest nursery “Alto Tevere” in the framework of the law for the mountains). In: Ducci F, De Rogatis A (eds) Risorse Genetiche Forestali in Italia. I. Breve storia delle attività sulle Risorse Genetiche Forestali (Forest genetic resources in Italy. I. Short history of activities on forest genetic resources). CRA SEL, Arezzo, pp 32–36

Morandini R, Magini E (1975) Il materiale forestale di propagazione in Italia (Forest reproductive materials in Italy). Ministero dell’ Agricoltura e Foreste, Roma, 295pp

790

791

792

793

794

795

796

797

798

799

800

801

802

803

804

805

806

807

808

809

810

811

812

813

814

815

816

817

818

819

820

821

822

823

824

825

826

827

828

829

830

831

832

833

834

835

836

837

838

839

840

841

842

843

http://dx.doi.org/10.1007/s00468-010-0507-9


Nahal I (1986) Taxonomie et aire géographique des pins du groupe halepensis (Taxonomy and distribution areas of the pines belonging to the halepensis group). Options Méditerranéennes 1:1–9

Ne’eman G, Lahav H, Izhaki I (1992) Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest. Oecologia 91:365–370

Ne’eman G, Goubitz S, Nathan R (2004) Reproductive traits of Pinus halepensis in the light of fire – a critical review. Plant Ecol 171:69–79

Nieto P, Aguado A, Ruano JR, Climent J, Chambel R, Argilés MA (2009) Ensayo de evaluación genética para la admisión de la Unidad HS-24/46/001 como material de base para la produc-ción de material forestal de reproducción de la categoría ‘CONTROLADO’ (Genetic evalua-tion trial for the admission of unit HS-24/46/001 as a basic material for the production of forest reproductive materials of CONTROLLED quality). In: Montes y sociedad: saber que hacer. Actas del 5º Congreso Forestal Español, Avila, 21–25 Sept 2009

Panetsos K (1975) Natural hybridisation between Pinus halepensis and Pinus brutia in Greece. Silvae Genetica 24:163–168

Panetsos KP (1981) Monograph of Pinus halepensis Mill. and P. brutia Ten. Ann For 9:39–77Panetsos K, Scaltsoyiannes A, Alizoti P (1994) Effect of genotype and cutting type on the vegeta-

tive propagation of the pine hybrid (Pinus brutia (Ten.) × Pinus halepensis (Mill.)). Annales des Sciences Forestières 51:447–454

Panetsos KP, Scaltsoyiannes A, Aravanopoulos FA, Dounavi K, Demetrakopoulos A (1997) Identification of Pinus brutia Ten., P. halepensis Mill., and their putative hybrids. Silvae Genetica 46:253–257

Pardos M, Climent J, Gil L, Pardos JA (2003) Shoot growth components and flowering phenology in grafted Pinus halepensis Mill. Tree Struct Funct 17:442–450

Pardos M, Calama R, Climent J (2009) Difference in cuticular transpiration and sclerophylly in juvenile and adult pine needles relates to the species-specific rates of development. Tree Struct Funct 23:501–508

Parra JL (1980) Creación de huertos semilleros de pino carrasco e injertos de pino piñonero sobre carrasco, en la provincia de Murcia (Creation of seed orchards of Pinus halepensis and grafting of Pinus pinea on Pinus halepensis in Murcia province (Spain)). Boletin de la Estacion Central de Ecologia 18:15–23

Pausas J, Ribeiro E, Vallejo R (2004) Post-fire regeneration variability of Pinus halepensis in the eastern Iberian Peninsula. For Ecol Manage 203:251–259

Pausas J, Llovet J, Rodrigo A, Vallejo R (2008) Are wildfires a disaster in the Mediterranean basin? – a review. Int J Wildland Fire 17:713–723

Pichot C (1995) Conservation et amélioration du Cyprès de l’Atlas et du pin d’Alep au Maroc (Conservation and breeding of Moroccan Cypress and Aleppo pine in Morocco). Mission report F.A.O.-UTF/MOR/011. 27/11-3/12/1994 and 27/3-8/4/1995, 28pp

Pichot C, Vauthier D (2007) Pinus halepensis – Pinus brutia: French comparative provenance tests. FAO-Silva Mediterranea and IUFRO WG 20213 expert joint meeting: promotion and use of results from the international trials of Mediterranean Conifers, Arezzo/Rome, 21–23 June 2007

Plomion C, Pichot C (2001) Global, physiological and molecular responses to climatic stresses of three mediterranean conifers. Final report. INCO-DC: international cooperation with develop-ing countries. Contract number: ERBIC 18CT 970 200. 277

Price RA, Liston A, Strauss SH (1998) Phylogeny and systematic of Pinus. In: Richardson DM (ed) Ecology and biogeography of Pinus. Cambridge University Press, Cambridge, pp 49–68

Rathgeber C, Nicault A, Guiot J, Keller T, Guibal F, Roche P (2000) Simulated responses of Pinus halepensis forest productivity to climatic change and CO

2 increase using a statistical model.

Glob Planet Change 26:405–421Richardson D (1998) Forestry trees as invasive aliens. Conserv Biol 12:18–26Richardson D, Cowling R, Le Maitre D (1990) Assessing the risk of invasive success in Pinus and

Banksia in South African mountain fynbos. J Veg Sci 1:629–642Rigolot E (2004) Predicting postfire mortality of Pinus halepensis Mill. and Pinus pinea L. Plant

Ecol 171:139–151

844

845

846

847

848

849

850

851

852

853

854

855

856

857

858

859

860

861

862

863

864

865

866

867

868

869

870

871

872

873

874

875

876

877

878

879

880

881

882

883

884

885

886

887

888

889

890

891

892

893

894

895

896

897

M.R. Chambel et al.

Sabaté S, Gracia C, Sánchez A (2002) Likely effects of climate change on growth of Quercus ilex, Pinus halepensis, Pinus pinaster, Pinus sylvestris and Fagus sylvatica forests in the Mediterranean region. For Ecol Manage 162:23–37

Santas LA (1983) Insects producing honeydew exploited by bees in Greece. Apidologie 14:93–103Santos-del-Blanco L, Notivol E, Zas R, Chambel MR, Majada J, Climent J (2010) Variation of

early reproductive allocation in multi-site genetic trials of Maritime pine and Aleppo pine. For Syst 19:381–392

Sarris D, Christodoulakis D, Koerner C (2007) Recent decline in precipitation and tree growth in the eastern Mediterranean. Glob Chang Biol 13:1187–1200

Scaltsoyiannes A, Panetsos K, Economou A, Tsoulpha P (1994) Micropropagation of the pine hybrid Pinus-brutia (Ten.) × Pinus-halepensis (Mill.) by culturing fascicle shoots. Annales des Sciences Forestières 51:175–182

Scarascia-Mugnozza G (1986) Recherches sur l’ecophysiologie de Pinus halepensis Mill. (Research on the ecophysiology of Pinus halepensis Mill.). Options Méditerranéennes 1:89–97

Scarascia-Mugnozza G, Riguzzi F, Lauteri M (2011) Effects of the climate change on the Mediterranean conifer ecosystems. In: Besacier C, Ducci F, Malagnoux M, Souvannavong O (eds) Status of the experimental network of Mediterranean forest genetic resources. CRA SEL, Arezzo and FAO Silva Mediterranea, Rome, pp 185–191

Schiller G (2011) The case of Yatir forest. In: Bredemeier M, Cohen S, Godbold DL, Lode E, Pichler V, Schleppi P (eds) Forest management and the water cycle – an ecosystem-based approach, Ecological studies 212. Springer, Dordrecht/Heidelberg/London/New York, pp 163–186

Schiller G, Atzmon N (2009) Performance of Aleppo pine (Pinus halepensis) provenances grown at the edge of the Negev desert: a review. J Arid Environ 73:1051–1057

Schiller G, Conkle MT, Grunwald C (1986) Local differentiation among Mediterranean popula-tions of Aleppo pine in their isoenzymes. Silvae Genetica 35:11–19

Schroter D, Cramer W, Leemans R, Prentice I, Araujo M, Arnell N, Bondeau A, Bugmann H, Carter T, Gracia C (2005) Ecosystem service supply and vulnerability to global change in Europe. Science 310:1333–1337

Shmida A, Lev-Yadun S, Goubitz S, Ne’eman G (2000) Sex allocation and gender segregation in Pinus halepensis, P. brutia and P. pinea. In: Ne’eman G, Trabaud L (eds) Ecology, biogeogra-phy and management of Pinus halepensis and P. brutia forest ecosystems in the Mediterranean Basin. Backhuys Publishers, Leiden, pp 91–104

Spanos I, Daskalakou E, Thanos C (2000) Postfire, natural regeneration of Pinus brutia forests in Thasos island, Greece. Acta Oecologica 21:13–20

Spencer D (1985) Dry country pines: provenance evaluation of the Pinus halepensis-P. brutia complex in the semi-arid region of south-east Australia. Aust For Res 15:263–279

Tabeaud M, Simon L (1993) Dommages et dégâts dans les forêts françaises: éléments pour un bilan (Damages and injuries to French forests: elements for an assessment). Annales de Géographie 102:339–358

Tapias R, Gil L, Fuentes-Utrilla P, Pardos JA (2001) Canopy seed banks in Mediterranean pines of southeastern Spain: a comparison between Pinus halepensis Mill., P. pinaster Ait., P. nigra Arn. and P. pinea L. J Ecol 89:629–638

Tapias R, Climent J, Pardos JA, Gil L (2004) Life histories of Mediterranean pines. Plant Ecol 171:53–68

Teisseire H, Fady B, Pichot C (1995) Allozyme variation in five French populations of Aleppo pine (Pinus halepensis Miller). For Genet 2:225–236

Thanos C, Daskalakou E, Nikolaidou S (1996) Early post fire regeneration of a Pinus halepensis forest on Mount Párnis, Greece. J Veg Sci 7:273–280

Thibault B, Loup C, Chanson B, Dilem A (1992) La valorisation du pin d’Alep en zone méditer-ranéenne française (Promotion of Aleppo pine in the Mediterranean French area). Forêt Méditerranéenne 13:226–237

Thuiller W (2003) BIOMOD–optimizing predictions of species distributions and projecting potential future shifts under global change. Glob Chang Biol 9:1353–1362

898

899

900

901

902

903

904

905

906

907

908

909

910

911

912

913

914

915

916

917

918

919

920

921

922

923

924

925

926

927

928

929

930

931

932

933

934

935

936

937

938

939

940

941

942

943

944

945

946

947

948

949

950

951


Tognetti R, Michelozzi M, Giovannelli A (1997) Geographical variation in water relations, hydraulic architecture and terpene composition of Aleppo pine seedlings from Italian prov-enances. Tree Physiol 17:241–250

Topak M (1997) Directory of seed sources of the Mediterranean conifers. Fao, Rome, 118ppTozkar CO, Önde S, Kaya Z (2009) The phylogenetic relationship between populations of margin-

ally and sympatrically located Pinus halepensis Mill. and Pinus brutia Ten. in Turkey, based on the ITS-2 region. Turk J Agric For 33:363–373

Vidakovic M (1991) Conifers: morphology and variation. Graficki Zavod, Hrvatske, 750ppVoltas J, Chambel MR, Prada MA, Ferrio JP (2008) Climate-related variability in carbon and

oxygen stable isotopes among populations of Aleppo pine grown in common-garden tests. Tree Struct Funct 22:759–769

Wahid N, Jouidre H, Lamhamedi MS, El Abidine AZ, Boulli A (2010) Évaluation de la structure et de la variabilité génétiques des populations naturelles du pin d’Alep (Pinus halepensis Mill.) au Maroc à l’aide de marqueurs isoenzymatiques (Evaluation of the genetic variability and structure of natural populations of Aleppo pine (Pinus halepensis Mill.) in Morocco using allozyme markers). Acta Botanica Gallica 157:419–431

Weinstein A (1989) Provenance evaluation of Pinus halepensis, P. brutia and P. eldarica in Israel. For Ecol Manage 26:215–225

Zavala M, Zea E (2004) Mechanisms maintaining biodiversity in Mediterranean pine-oak forests: insights from a spatial simulation model. Plant Ecol 171:197–207

952

953

954

955

956

957

958

959

960

961

962

963

964

965

966

967

968

969

970

971

Author QueriesChapter No.: 5 0001864952

Queries Details Required Author’s Response

AU1 Please provide department name for all the authors.

AU2 Thibault (1992), Bariteau et al. (1992), Agúndez (1997, 1999), Schiller et al. (2011) have been changed to Thibault et al. (1992), Bariteau (1992), Agúndez et al. (1997, 1999), Schiller (2011), respectively as per the reference list. Please check.

AU3 Please confirm the inserted citation for Fig. 5.11.

AU4 CO2 it should be changed to CO2. Please confirm.

AU5 Please provide significance for “?” in Table 5.7.

AU6 Partitionist Method (2010), Teskey et al. (1986), Tognetti et al. (2000), Guehl et al. (1995) are cited in text but not given in the reference list. Please check.

AU7 Please provide complete details in Anonymous (2010), Pichot (1995), Plomion and Pichot (2001).

Species breeding monograph: Aleppo pine (Pinus halepensis Mill.) and Brutia pine (Pinus brutia Ten.)

Documents