02.02.2013 Views

Pinus halepensis - Euforgen

Pinus halepensis - Euforgen

Pinus halepensis - Euforgen

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

EUFORGEN<br />

Technical guidelines for genetic conservation and use<br />

Aleppo and Brutia pines<br />

<strong>Pinus</strong> <strong>halepensis</strong>/<strong>Pinus</strong> brutia<br />

Bruno Fady1 , Hacer Semerci2 and Giovanni G. Vendramin3 1 INRA, Mediterranean Forest Research Unit, Avignon, France<br />

2 Forest Tree Seeds and Tree Breeding Research Directorate, Gazi-<br />

Ankara, Turkey<br />

3 CNR, Institute of Plant Genetics, Firenze, Italy<br />

These Technical Guidelines are intended to assist those who cherish the valuable Aleppo pine<br />

and Brutia pine genepools and wish to ensure their sustainability, through conserving important<br />

seed sources or use in practical forestry. The focus is on conserving the genetic diversity of the<br />

species at the European scale. The recommendations provided in this module should be<br />

regarded as a commonly agreed basis to be complemented and further developed in local,<br />

national or regional conditions. The Guidelines are based on the available knowledge of the<br />

species and on widely accepted methods for the conservation of forest genetic resources.<br />

Biology and ecology<br />

Aleppo pine (<strong>Pinus</strong> <strong>halepensis</strong><br />

Mill.) has a pedunculate cone,<br />

and fine, flexible, light green needles<br />

5–10 cm long. Brutia pine<br />

(<strong>Pinus</strong> brutia Ten.) has a sessile<br />

cone and strong, dark green<br />

needles 10–18 cm long. Both<br />

species are wind-pollinated and<br />

allogamous. Male and female<br />

flowers are located on different<br />

parts of a tree (monoecy). Both<br />

species are extremely prolific<br />

seed dispersers and can colonize<br />

open and disturbed<br />

areas easily.<br />

Aleppo and<br />

Brutia pine<br />

forests can grow<br />

on all substrates and<br />

almost all bioclimates<br />

of the Mediterranean<br />

region. They can be found<br />

at altitudes of 0–600 m in the<br />

northern Mediterranean and<br />

0–1400 m in the southern<br />

Mediterranean (thermo- and<br />

meso-Mediterranean levels).<br />

Locally, they can reach higher<br />

altitudes, e.g. 2600 m for


inus <strong>halepensis</strong> P<br />

ppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong><br />

P. <strong>halepensis</strong> in the Higher Atlas<br />

of Morocco and 1650 m for<br />

P. brutia in the Taurus Mountains<br />

of Turkey. At the upper limit of<br />

their distribution, they often constitute<br />

a pre-forest colonizing<br />

stage or are part of a mixed pineoak<br />

forest.<br />

Optimal development of<br />

P. <strong>halepensis</strong> forests occurs at<br />

annual rainfalls of 350–700 mm<br />

and absolute mean minimum<br />

temperatures between –2 and<br />

+10°C (semi-arid and sub-humid<br />

bioclimates). Optimal development<br />

of P. brutia forests requires<br />

higher rainfalls but accepts a<br />

wider range of temperatures<br />

(absolute mean minimum temperatures<br />

between –5 and +10°C,<br />

sub-humid and humid bioclimates).<br />

Distribution Importance and use<br />

<strong>Pinus</strong> <strong>halepensis</strong> and P. brutia<br />

form a group of related species<br />

that can intercross, but occupy<br />

different geographical ranges<br />

and bioclimates.<br />

Aleppo pine forests cover<br />

extensive areas in the western<br />

Mediterranean: Spain, France,<br />

Italy, Croatia, Albania, Greece,<br />

Morocco, Algeria, Tunisia, Libya<br />

and Malta. A few natural and artificial<br />

populations can be found in<br />

the eastern Mediterranean in<br />

Turkey, Syria, Israel, Jordan and<br />

Lebanon. Total forest cover is<br />

estimated to be approximately<br />

3.5 million hectares.<br />

Brutia pine forests cover<br />

extensive areas in the Eastern<br />

Mediterranean: Greece, Turkey,<br />

Cyprus, Syria and Lebanon. A<br />

few small populations can be<br />

found in Iraq and Iran.<br />

Other related taxonomic<br />

groups are<br />

present in Ukraine<br />

(Crimea, P. stankewiczii<br />

Sukaczew), around the<br />

Black Sea (Georgia, Russian<br />

Federation, Ukraine, P. pithyusa<br />

Stevenson) and in the Caucasus<br />

(Azerbaijan, Georgia, Iran,<br />

Turkey, P. eldarica Medw.). Total<br />

forest cover is estimated to be<br />

over 4 million hectares, of which<br />

3.8 million hectares are in Turkey.<br />

Aleppo and Brutia pines represent<br />

the only or main source of<br />

wood and forest cover in many<br />

Mediterranean countries. Economically,<br />

P. brutia is the most<br />

important conifer species in<br />

Turkey; P. <strong>halepensis</strong> is the most<br />

important forest species of North<br />

Africa, and has high ecological<br />

importance in southern France<br />

and Italy, especially at the urbanforest<br />

interface. Mean productivity<br />

is approximately 1–2 m 3<br />

ha -1 year -1 for Aleppo pine, and<br />

2–3 m 3 ha -1 year -1 for Brutia pine.<br />

Maximum yield can reach<br />

12–15 m 3 for both species. The<br />

wood of these Mediterranean<br />

pines is used for many purposes:<br />

construction, industry, carpentry,<br />

firewood and pulp. Seeds are<br />

also used for making pastry.


inus brutia <strong>Pinus</strong><br />

<strong>Pinus</strong> brutiaAleppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and Brutia pinesP<br />

Genetic knowledge<br />

Genetic inventories using biochemical<br />

and DNA markers have<br />

demonstrated that genetic diversity<br />

is geographically structured.<br />

Most P. <strong>halepensis</strong> diversity was<br />

found in Greek and Spanish populations<br />

although other populations<br />

had lower diversity than<br />

other conifer species. This is<br />

consistent with the hypothesis of<br />

a recent expansion of the<br />

species (in the last 10 000 years)<br />

from these two refugial areas,<br />

with colonizing populations<br />

establishing by migration of a<br />

limited number of individuals<br />

(founder effect) and/or population<br />

dynamics regulated by fire<br />

(population bottlenecks). Genetic<br />

diversity is higher for P. brutia<br />

and roughly separates<br />

Western and Eastern populations.<br />

Controlled pollination<br />

experiments and monitoring<br />

of sympatric east Mediterranean<br />

populations using<br />

molecular markers have<br />

demonstrated that unidirectional<br />

gene flow is possible from<br />

P. <strong>halepensis</strong> to P. brutia resulting<br />

in the emergence of hybrids.<br />

Hybridization is not possible<br />

using P. brutia as pollen parent.<br />

Provenance and laboratory<br />

tests have revealed significant<br />

geographic patterns in adaptive<br />

trait variability. Although both<br />

species demonstrate polycyclic<br />

annual growth patterns, initial<br />

shoot units are smaller in<br />

P. <strong>halepensis</strong>. The easternmost<br />

P. <strong>halepensis</strong> provenances tend<br />

to have higher juvenile growth.<br />

<strong>Pinus</strong> <strong>halepensis</strong> is better adapted<br />

to drought but less adapted<br />

to cold than P. brutia. However,<br />

under severe water-stress conditions,<br />

P. <strong>halepensis</strong> has an<br />

increased sensitivity to the fungus<br />

Sphaeropsis sapinea. Both<br />

species are sensitive to the pine<br />

procecionnary moth Thaumetopoea<br />

pityocampa which can<br />

cause severe defoliation. <strong>Pinus</strong><br />

<strong>halepensis</strong> is sensitive to the pine<br />

bast scale Matsucoccus josephii,<br />

although P. brutia is resistant to<br />

it.<br />

These, along with ecological<br />

studies, have been used to<br />

define seed-collection zones and<br />

seed stands in several countries<br />

(e.g. 29 P. <strong>halepensis</strong><br />

seed stands<br />

in France).<br />

Threats to<br />

genetic diversity<br />

Aleppo and Brutia pines are not<br />

considered ecologically threatened<br />

as a whole. However P.<br />

pityusa is considered vulnerable<br />

because of population size<br />

reduction linked to habitat<br />

decline (IUCN red list). <strong>Pinus</strong><br />

eldarica has a patchy distribution<br />

and its genetic diversity is lowest<br />

among the taxa of this group.<br />

Insects such as Matsucoccus<br />

josephii represent a major threat<br />

in the Eastern Mediterranean.<br />

Thaumatopea pityocampa can<br />

also induce severe defoliation<br />

throughout the distribution area<br />

of both pines although it does<br />

not often lead to mortality.<br />

Recently, the canker Crumenulopsis<br />

sororia has started to<br />

cause severe defoliation<br />

and dieback on<br />

P. <strong>halepensis</strong> in<br />

France. The impact of<br />

forest fires is ambivalent.<br />

Although they actually promote<br />

regeneration, they could be<br />

responsible for rare allele<br />

changes over generations,<br />

explain the very low diversity<br />

found in P. <strong>halepensis</strong> and promote<br />

the spreading of<br />

P. <strong>halepensis</strong> genes into P. brutia<br />

forests. Among-region seed<br />

transfers have led to significant<br />

frost or water-stress damage<br />

after planting when ill-adapted<br />

material was used. Reducing<br />

local population adaptability<br />

through gene flow from plantations<br />

is also a risk. Finally,


<strong>halepensis</strong> Pin<br />

<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and<br />

because these species (and<br />

especially P. <strong>halepensis</strong>) are<br />

highly resistant to drought, they<br />

are often the last forest species<br />

to be found at desert or steppe<br />

margins. Global warming and its<br />

collateral modification of rainfall<br />

regimes may dramatically modify<br />

their distribution ranges.<br />

Guidelines for genetic<br />

conservation and use<br />

Current conservation measures<br />

undertaken at national levels<br />

most commonly include in situ<br />

gene conservation networks<br />

specifically designed for the target<br />

species (e.g. in Turkey, 52<br />

P. brutia conservation units) and<br />

forest reserves or national parks<br />

which include the target species.<br />

Ex situ measures include clonal<br />

archives, cold storage seed<br />

banks and DNA banks.<br />

To increase the efficiency<br />

of in situ genetic resource<br />

conservation, a concerted<br />

management effort should<br />

be carried out range-wide.<br />

Although transfer of seed<br />

material is often legally possible,<br />

it should be avoided<br />

across zones and countries<br />

with different ecological<br />

requirements, notably<br />

because of cold, drought and<br />

insect damage risks.<br />

Locally, some populations<br />

require specific attention and<br />

appropriate forestry practice.<br />

Marginal populations. As populations<br />

at high altitudes, in desert<br />

margins and mixed forests may<br />

contain valuable genes (resistance<br />

to drought, cold, pests) for<br />

adaptation under global warming,<br />

efforts such as gene<br />

reserves should be made<br />

to conserve them.<br />

Population under recurrent<br />

forest fires. Because they are<br />

adapted to forest fires, both<br />

pines usually regenerate well<br />

after fire, using the seed bank<br />

released from serotinous cones.<br />

If regeneration happens to be<br />

poor in the first 2 years after fire,<br />

and if only a few isolated seed<br />

trees remain in the burnt area,<br />

artificial regeneration should be<br />

used to counteract the risk of<br />

genetic erosion in the juveniles.<br />

In this case, seed lots collected<br />

from large genepools should be<br />

used (e.g. at least 30 trees per<br />

population from at least three<br />

populations from a single seed<br />

zone).<br />

Populations where hybridization<br />

may occur. Planting Aleppo<br />

pine where Brutia pine is present<br />

should be avoided in areas<br />

where frost and potential pest<br />

damage are limiting factors, or<br />

strictly monitored in areas where<br />

drought is the limiting factor.<br />

Owing to the anisotropy of<br />

between-species gene flow, the<br />

impact should be reduced when<br />

planting Brutia pine in the vicinity<br />

of Aleppo pine forests.


<strong>Pinus</strong> <strong>halepensis</strong><br />

Aleppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and Brutia pines<strong>Pinus</strong> h<br />

Distribution range of Aleppo pine<br />

Distribution range of Brutia pine


<strong>Pinus</strong> brutia Pi<br />

alepensis <strong>Pinus</strong> brutiaAleppo and Brutia pines<strong>Pinus</strong> <strong>halepensis</strong> <strong>Pinus</strong> brutiaAleppo and Bruti<br />

EUFORGEN<br />

These Technical Guidelines were<br />

produced by members of the<br />

EUFORGEN Conifers Network.<br />

The objective of the Network is to<br />

identify minimum genetic conservation<br />

requirements in the long<br />

term in Europe, in order to reduce<br />

the overall conservation cost and<br />

to improve the quality of standards<br />

in each country.<br />

Citation: Fady, B., H. Semerci and<br />

G.G. Vendramin. 2003. EUFOR-<br />

GEN Technical Guidelines for<br />

genetic conservation and use for<br />

Aleppo pine (<strong>Pinus</strong> <strong>halepensis</strong>)<br />

and Brutia pine (<strong>Pinus</strong> brutia).<br />

International Plant Genetic<br />

Resources Institute, Rome, Italy.<br />

6 pages.<br />

Drawings: <strong>Pinus</strong> <strong>halepensis</strong>, Claudio<br />

Giordano. © IPGRI, 2003.<br />

ISBN 92-9043-571-2<br />

EUFORGEN Secretariat c/o IPGRI<br />

Via dei Tre Denari, 472/a<br />

00057 Maccarese (Fiumicino)<br />

Rome, Italy<br />

Tel. (+39)066118251<br />

Fax: (+39)0661979661<br />

euf_secretariat@cgiar.org<br />

Selected bibliography<br />

Bariteau, M., R. Huc and G.G. Vendramin (coordinators). 2001 Adaptation and<br />

selection of Mediterranean <strong>Pinus</strong> and Cedrus for sustainable afforestation<br />

of marginal lands. Final report of EU Project FAIR CT95-0097, 173 p.<br />

Bucci, G., M. Anzidei, A. Madaghiele and G.G. Vendramin. 1998. Detection of<br />

haplotypic variation and natural hybridization in <strong>halepensis</strong>-complex pine<br />

species using chloroplast simple sequence repeat (SSR) markers. Molecular<br />

Ecology 7(12):1633-1643.<br />

Conkle, M.T., G. Schiller and C. Grunwald. 1988. Electrophoretic analysis of<br />

diversity and phylogeny of <strong>Pinus</strong> brutia and closely related taxa. Systematic<br />

Botany 13(3):411-424.<br />

Kaundun, S.S., B. Fady and Ph. Lebreton. 1997. Genetic differences between<br />

<strong>Pinus</strong> <strong>halepensis</strong>, <strong>Pinus</strong> brutia and <strong>Pinus</strong> eldarica based on needle<br />

flavonoids. Biochemical Systematics and Ecology 25(6):553-562.<br />

Ne’eman, G. and L. Trabaud (eds.). 2000. Ecology, biogeography and management<br />

of <strong>Pinus</strong> <strong>halepensis</strong> and P. brutia forest ecosystems in the Mediterranean<br />

basin. Backhuys Publishers, Leiden, The Netherlands. 407 pages.<br />

The distribution map was compiled by members of the EUFORGEN<br />

Conifers Network based on an earlier map published by W.B.Critchfield &<br />

E.L.Little, Jr. in 1966 (Geographic Distibution, of the Pines of the World,<br />

USDA Forest Service Misc. Publication, 991 pages).<br />

More information<br />

www.euforgen.org

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!