Establishing Quercus ilex under Mediterranean dry conditions: sowing recalcitrant acorns versus planting seedlings at different depths and tube shelter light transmissions

Article New Forests November 2015, Volume 46, Issue 5, pp 869-883 First online: 17 June 2015 Establishing Quercus ilex under Mediterranean dry conditions: sowing recalcitrant acorns versus planting seedlings at different depths and tube shelter light transmissionsJuan A. OlietAffiliated withDepartamento de Sistemas y Recursos Naturales, E.T.S. Ingenieros de Montes, Universidad Politécnica de Madrid Email author View author's OrcID profile , Alberto Vázquez de CastroAffiliated withDepartamento de Sistemas y Recursos Naturales, E.T.S. Ingenieros de Montes, Universidad Politécnica de Madrid, Jaime PuértolasAffiliated withLancaster Environment Centre, Lancaster University $39.95 / €34.95 / £29.95 * Rent the article at a discount Rent now * Final gross prices may vary according to local VAT. Get Access AbstractSuccess of Mediterranean dry areas restoration with oaks is a challenging goal. Testing eco-techniques that mimic beneficial effects of natural structures and ameliorate stress contributes to positive solutions to overcoming establishment barriers. We ran a factorial experiment in a dry area, testing two levels of solid wall transmission of tube shelters (60 and 80 %) plus a control mesh, and two depths (shallow and 15 cm depth) of placing either planted seedlings or acorns of Quercus ilex. Microclimate of the planting or sowing spots was characterized by measuring photosynthetically active radiation, temperature and relative humidity. Plant response was evaluated in terms of survival, phenology, acorn emergence and photochemical efficiency (measured through chlorophyll fluorescence). We hypothesize that tube shelters and deep planting improve Q. ilex post-planting and sowing performance because of the combined effects of reducing excessive radiation and improving access to moist soil horizons. Results show that temperature and PAR was reduced, and relative humidity increased, in deep spots. Midsummer photochemical efficiency indicates highest level of stress for oaks in 80 % light transmission shelter. Optimum acorn emergence in spring was registered within solid wall tree shelters, and maximum summer survival of germinants and of planted seedlings occurred when acorns or seedlings were placed at 15 cm depth irrespectively of light transmission of shelter. Survival of germinants was similar to that of planted seedlings. The importance of techniques to keep high levels of viability after sowing recalcitrant seeds in the field is emphasized in the study

The influence of commercial film-forming polymers on reducing salt spray injury in evergreen oak (Quercus ilex L.) and laurel (Prunus laurocerasus L.)

A field trial was undertaken to determine the influence of four commercially available film-forming polymers (Bond [alkyl phenyl hydroxyl polyoxyethylene], Newman Crop Spray 11E™ [paraffinic oil], Nu-Film P [poly-1-p menthene], and Spray Gard [di-1-p menthene]) on reducing salt spray injury on two woody species, evergreen oak (Quercus ilex L.) and laurel (Prunus laurocerasus L.). Irrespective of species, the film-forming polymers Nu-Film-P and Spay Gard did not provide any significant degree of protection against salt spray damage irrespective of concentration (1% or 2%) applied as measured by leaf chlorophyll concentrations, photosynthetic efficiency, visual leaf necrosis, foliar sodium and chloride content, and growth (height, leaf area). The film-forming polymer Newman Crop Spray 11E™ provided only 1-week protection against salt spray injury. The film-forming polymer Bond provided a significant (P < 0.05) degree of protection against salt spray injury 3 months after application as manifest by higher leaf chlorophyll content, photosynthetic efficiency, height and leaf area, and lower visual leaf necrosis and foliar Na and Cl content compared with nontreated controls. In conclusion, results indicate that application of a suitable film-forming polymer can provide a significant degree of protection of up to 3 months against salt spray injury in evergreen oak and laurel. Results also indicate that when applied at 1% or 2% solutions, no problems associated with phytotoxicity and rapid degradation on the leaf surface exist.

Holm Oak (Quercus ilex L.) canopy as interceptor of airborne trace elements and their accumulation in the litter and topsoil

We investigated the role of urban Holm Oak (Quercus ilex L.) trees as airborne metal accumulators and metals' environmental fate. Analyses confirmed Pb, Cd, Cu and Zn as main contaminants in Siena's urban environment; only Pb concentrations decreased significantly compared to earlier surveys. Additionally, we determined chemical composition of tree leaves, litter and topsoil (underneath/outside tree crown) in urban and extra-urban oak stands. Most notably, litter in urban samples collected outside the canopy had significantly lower concentrations of organic matter and higher concentrations of Pb, Cu, Cd and Zn than litter collected underneath the canopy. There was a greater metals' accumulation in topsoil, in samples collected under the tree canopy and especially near the trunk ('stemflow area'). Thus, in urban ecosystems the Holm Oak stands likely increase the soil capability to bind metals.

Sulfate and Nitrate Assimilation in Leaves of Quercus ilex and Quercus pubescens Grown Near Natural CO2 Springs in Central Italy

The effect of long-term exposure to elevated pCO2 concentrations on sulfate and nitrate assimilation was studied under field conditions using leaves from Quercus ilex and Quercus pubescens trees growing with ambient or elevated CO2 concentrations in the vicinity of three natural CO2 springs, Bossoleto, Laiatico and Sulfatara, in Tuscany, Italy. The activity of the key enzymes of sulfate assimilation, adenosine 5′-phosphosulfate reductase (APR) and nitrate assimilation, nitrate reductase (NR), were measured together with the levels of acid soluble thiols, and soluble non-proteinogenic nitrogen compounds. Whereas NR activity remained unaffected in Q. ilex or increased Q. pubescence, APR activity decreased in the area of CO2 springs. The latter changes were often accompanied by increased GSH concentrations, apparently synthesized from H2S and SO2 present in the gas mixture emitted from the CO2 springs. Thus, the diminished APR activity in leaves of Q. ilex and Q. pubescence from spring areas can best be explained by the exposure to gaseous sulfur compounds. Although the concentrations of H2S and SO2 in the gas mixture emitted from the vents at the CO2 springs were low at the Bossoleto and Laiatico spring, these sulfur gases pose physiological effects, which may override consequences of elevated pCO2.

Life-long growth of Quercus ilex L. at natural CO2 springs acclimates sulphur, nitrogen and carbohydrate metabolism of the progeny to elevated pCO2

The aim of the present study was to analyse whether offspring of mature Quercus ilex trees grown under life-long elevated pCO2 show alterations in the physiological response to elevated pCO2 in comparison with those originating from mature trees grown at current ambient pCO2. To investigate changes in C- (for changes in photosynthesis, biomass and lignin see Polle, McKee & Blaschke Plant, Cell and Environment 24, 1075–1083, 2001), N-, and S-metabolism soluble sugar, soluble non-proteinogenic nitrogen compounds (TSNN), nitrate reductase (NR), thiols, adenosine 5′-phosphosulphate (APS) reductase, and anions were analysed. For this purpose Q. ilex seedlings were grown from acorns of mother tree stands at a natural spring site (elevated pCO2) and a control site (ambient pCO2) of the Laiatico spring, Central Italy. Short-term elevated pCO2 exposure of the offspring of control oaks lead to higher sugar contents in stem tissues, to a reduced TSNN content in leaves, and basipetal stem tissues, to diminished thiol contents in all tissues analysed, and to reduced APS reductase activity in both, leaves and roots. Most of the components of C-, N- and S-metabolism including APS reductase activity which were reduced due to short-term elevated pCO2 exposure were recovered by life-long growth under elevated pCO2 in the offspring of spring oaks. Still TSNN contents in phloem exudates increased, nitrate contents in lateral roots and glutathione in leaves and phloem exudates remained reduced in these plants. The present results demonstrated that metabolic adaptations of Q. ilex mother trees to elevated pCO2 can be passed to the next generation. Short- and long-term effects on source-to-sink relation and physiological and genetic acclimation to elevated pCO2 are discussed.

Wettability, polarity and water absorption of Quercus ilex leaves: effect of leaf side and age

Plant trichomes play important protective functions and may have a major influence on leaf surface wettability. With the aim of gaining insight into trichome structure, composition and function in relation to water-plant surface interactions, we analyzed the adaxial and abaxial leaf surface of Quercus ilex L. (holm oak) as model. By measuring the leaf water potential 24 h after the deposition of water drops on to abaxial and adaxial surfaces, evidence for water penetration through the upper leaf side was gained in young and mature leaves. The structure and chemical composition of the abaxial (always present) and adaxial (occurring only in young leaves) trichomes were analyzed by various microscopic and analytical procedures. The adaxial surfaces were wettable and had a high degree of water drop adhesion in contrast to the highly unwettable and water repellent abaxial holm oak leaf sides. The surface free energy, polarity and solubility parameter decreased with leaf age, with generally higher values determined for the abaxial sides. All holm oak leaf trichomes were covered with a cuticle. The abaxial trichomes were composed of 8% soluble waxes, 49% cutin, and 43% polysaccharides. For the adaxial side, it is concluded that trichomes and the scars after trichome shedding contribute to water uptake, while the abaxial leaf side is highly hydrophobic due to its high degree of pubescence and different trichome structure, composition and density. Results are interpreted in terms of water-plant surface interactions, plant surface physical-chemistry, and plant ecophysiology.

Development of somatic embryogenesis for cloning and conservation of mature holm oak trees (Quercus ilex L.)

La encina (Quercus ilex L.) es una de las especies forestales mediterráneas más importantes. Constituye gran parte del estrato arbóreo de dehesas o montados, produce bellota como alimento del ganado y establece simbiosis con hongos micorrizógenos de gran valor económico. La encina está considerada como una especie recalcitrante en términos de conservación de semillas y capacidad morfogénica, lo que dificulta los programas de conservación de recursos genéticos y la mejora de la especie. La propagación vegetativa es una potente herramienta de los programas de mejora, por lo que es preciso desarrollar protocolos de regeneración somática en encina. La embriogénesis somática está considerada como la modalidad más adecuada de regeneración basada en técnicas de cultivo de tejidos vegetales utilizada en biotecnología forestal. Este trabajo se centra en el estudio de determinados aspectos de la embriogénesis somática para la regeneración clonal de encinas adultas. La memoria de esta tesis se ha dividido en capítulos que se corresponden con diferentes aspectos del sistema embriogénico. La embriogénesis somática se indujo en tegumentos maternos de óvulos en desarrollo procedentes de bellotas inmaduras de encinas adultas. A pesar de las bajas frecuencias de inducción, las líneas embriogénicas generadas se amplificaron mediante embriogénesis secundaria observándose cierta pérdida de la capacidad de diferenciación con el tiempo. Tanto el genotipo como la formulación del medio de cultivo influyeron en la respuesta embriogénica, concluyendo que la formulación de macronutrientes de Schenk y Hildebrant del medio sin reguladores de crecimiento fue la combinación más efectiva en la inducción. Los resultados sugirieron la existencia de una ventana en el desarrollo del óvulo más sensible a la inducción. El genotipo in[luyó en la capacidad proliferativa de los cultivos y en la conversión de los embriones somáticos, que se incrementó suplementando el medio con ácido indol-3-butírico y 6-benciladenina. El cultivo en medio líquido de líneas embriogénicas en condiciones de inmersión transitoria incrementó el crecimiento, dependiendo del genotipo, con respecto al cultivo en medio semisólido. Sin embargo, no mejoró la capacidad de diferenciar embriones cotiledonares aislados. Se estableció un protocolo de inicio y mantenimiento de cultivos en suspensión para varias líneas embriogénicas mediante inoculación en alta densidad de agregados embrionarios procedentes del medio semisólido. Para evitar la pérdida de vigor y la capacidad morfogénica debida al cultivo prolongado se desarrolló un protocolo de crioconservación de líneas embriogénicas mediante vitrificación. Al determinar la influencia de los agentes crioprotectores antes y después de su inmersión en nitrógeno líquido se concluyó que las respuestas de capacidad de crecimiento y de diferenciación del material embriogénico son independientes, además de estar bajo influencia del genotipo y el tipo de material crioconservado. La combinación de sacarosa y PVS2 previa a la inmersión en nitrógeno líquido proporcionó la mayor tasa de recuperación. Cuando las líneas fueron crioconservadas 30 días la capacidad de diferenciación se perdió en todas ellas. El análisis de SSR detectó variación somaclonal en el material crioconservado a corto plazo. SSR y RAPD mostraron importantes diferencias genéticas entre los árboles donantes y el material embriogénico que dependieron del genotipo. El grado de detección dependió del marcador empleado. Ambos marcadores revelaron baja inestabilidad intraclonal. Los RAPD revelaron variación genética intra-individuo en las encinas donantes. Se discuten la variación genética pre-existente en encina, su aparición durante las primeras fases de la inducción de embriogénesis, y la presencia de tejidos provenientes de la fertilización en el explanto materno. Esto hace preciso definir la identidad genética del material donante y acometer ensayos de detección precoz de variación somaclonal. ABSTRACT Holm oak (Quercus ilex L.) is one of the most important Mediterranean forest species. It conforms the tree layer of dehesas or montados, it produces acorns to feed the livestock and it establishes symbiosis with profitable mycorrhizal fungi. Holm oak is considered as recalcitrant species in terms of seed conservation and morphogenic capacities, which complicates the development of genetic conservation and improvement programs. Vegetative propagation is one of the mightiest tools for breeding programs therefore; developing protocols for clonal regeneration of holm oak is essential. Somatic embryogenesis is considered the best tissue culture-based way of plant regeneration in forest biotechnology. The present study is focused on the study of certain aspects of somatic embryogenesis for clonal regeneration of mature holm oak. This thesis manuscript is divided into several chapters that match with different aspects of the embryogenic system. Somatic embryogenesis induction was achieved on maternal teguments of developing ovules from immature acorns of adult holm oak trees. Despite the low induction frequencies, the generated embryogenic lines were amplified by secondary embryogenesis. A decline in the differentiation capacity over time was also observed. It was concluded that both genotype and culture media formulation influenced the embryogenic response, being the Schenk and Hildebrandt´s macronutrients formulation from culture medium and the lack of plant growth regulators the most effective combination for the induction of the embryogenic response. It has been suggested the existence of a developmental window in which ovules are prone to induction. Genotype influenced the proliferation capacity and the plant conversion of somatic embryos, which was also favoured by the presence of indol-3-butyric acid and 6-bencyladenine. The use of temporary immersion systems as proliferation in liquid culture of the embryogenic lines increased the growth depending on genotype, when compared to semisolid cultures. However, it did not improve the differentiation of single cotyledonary embryos. A protocol for the initiation and maintenance of embryogenic suspension cultures was established for several embryogenic lines with highly dense inoculi of embryogenic clusters from proliferating semisolid cultures. In order to avoid the loss of vigour and morphogenic ability of embryogenic lines due to prolonged cultures, a cryopreservation protocol for embryogenic lines of holm oak has been developed. During the determination of the influence of cryoprotective agents on the growth and differentiation capacities before and after liquid nitrogen immersion, it was concluded that both responses were independent from each other and also under the influence of genotype and the type of cryopreserved material. The combination of sucrose and PVS2 prior liquid nitrogen immersion provided higher recovery rates. When the same embryogenic lines were cryopreserved for 30 days, none was able to differentiate. The SSRs analysis of the short-term cryopreserved material detected somaclonal variation. Both SSR and RAPD markers showed high sensitivity to detect genetic differences between the donor trees and the generated embryogenic material. Nevertheless, the degree of instability detection depended on the marker. The SSR analysis indicated a relationship between genotype, the studied loci and the located polymorphisms. Also, both markers revealed low intraclonal genetic variation. The RAPD detected genetic variation within the donor trees. The presence of pre-existent genetic variation within mature trees, in addition to its occurrence during the early stages of the embryogenic induction, and the presence of tissues of fertilisation origin within the maternal explants are all discussed. Nonetheless, the determination of the genetic identity of donor material is required, in addition to early detection methods of somaclonal variation.

Development of somatic embryogenesis for cloning and conservation of mature holm oak trees (Quercus ilex L.)

La encina (Quercus ilex L.) es una de las especies forestales mediterráneas más importantes. Constituye gran parte del estrato arbóreo de dehesas o montados, produce bellota como alimento del ganado y establece simbiosis con hongos micorrizógenos de gran valor económico. La encina está considerada como una especie recalcitrante en términos de conservación de semillas y capacidad morfogénica, lo que dificulta los programas de conservación de recursos genéticos y la mejora de la especie. La propagación vegetativa es una potente herramienta de los programas de mejora, por lo que es preciso desarrollar protocolos de regeneración somática en encina. La embriogénesis somática está considerada como la modalidad más adecuada de regeneración basada en técnicas de cultivo de tejidos vegetales utilizada en biotecnología forestal. Este trabajo se centra en el estudio de determinados aspectos de la embriogénesis somática para la regeneración clonal de encinas adultas. La memoria de esta tesis se ha dividido en capítulos que se corresponden con diferentes aspectos del sistema embriogénico. La embriogénesis somática se indujo en tegumentos maternos de óvulos en desarrollo procedentes de bellotas inmaduras de encinas adultas. A pesar de las bajas frecuencias de inducción, las líneas embriogénicas generadas se amplificaron mediante embriogénesis secundaria observándose cierta pérdida de la capacidad de diferenciación con el tiempo. Tanto el genotipo como la formulación del medio de cultivo influyeron en la respuesta embriogénica, concluyendo que la formulación de macronutrientes de Schenk y Hildebrant del medio sin reguladores de crecimiento fue la combinación más efectiva en la inducción. Los resultados sugirieron la existencia de una ventana en el desarrollo del óvulo más sensible a la inducción. El genotipo in[luyó en la capacidad proliferativa de los cultivos y en la conversión de los embriones somáticos, que se incrementó suplementando el medio con ácido indol-3-butírico y 6-benciladenina. El cultivo en medio líquido de líneas embriogénicas en condiciones de inmersión transitoria incrementó el crecimiento, dependiendo del genotipo, con respecto al cultivo en medio semisólido. Sin embargo, no mejoró la capacidad de diferenciar embriones cotiledonares aislados. Se estableció un protocolo de inicio y mantenimiento de cultivos en suspensión para varias líneas embriogénicas mediante inoculación en alta densidad de agregados embrionarios procedentes del medio semisólido. Para evitar la pérdida de vigor y la capacidad morfogénica debida al cultivo prolongado se desarrolló un protocolo de crioconservación de líneas embriogénicas mediante vitrificación. Al determinar la influencia de los agentes crioprotectores antes y después de su inmersión en nitrógeno líquido se concluyó que las respuestas de capacidad de crecimiento y de diferenciación del material embriogénico son independientes, además de estar bajo influencia del genotipo y el tipo de material crioconservado. La combinación de sacarosa y PVS2 previa a la inmersión en nitrógeno líquido proporcionó la mayor tasa de recuperación. Cuando las líneas fueron crioconservadas 30 días la capacidad de diferenciación se perdió en todas ellas. El análisis de SSR detectó variación somaclonal en el material crioconservado a corto plazo. SSR y RAPD mostraron importantes diferencias genéticas entre los árboles donantes y el material embriogénico que dependieron del genotipo. El grado de detección dependió del marcador empleado. Ambos marcadores revelaron baja inestabilidad intraclonal. Los RAPD revelaron variación genética intra-individuo en las encinas donantes. Se discuten la variación genética pre-existente en encina, su aparición durante las primeras fases de la inducción de embriogénesis, y la presencia de tejidos provenientes de la fertilización en el explanto materno. Esto hace preciso definir la identidad genética del material donante y acometer ensayos de detección precoz de variación somaclonal. ABSTRACT Holm oak (Quercus ilex L.) is one of the most important Mediterranean forest species. It conforms the tree layer of dehesas or montados, it produces acorns to feed the livestock and it establishes symbiosis with profitable mycorrhizal fungi. Holm oak is considered as recalcitrant species in terms of seed conservation and morphogenic capacities, which complicates the development of genetic conservation and improvement programs. Vegetative propagation is one of the mightiest tools for breeding programs therefore; developing protocols for clonal regeneration of holm oak is essential. Somatic embryogenesis is considered the best tissue culture-based way of plant regeneration in forest biotechnology. The present study is focused on the study of certain aspects of somatic embryogenesis for clonal regeneration of mature holm oak. This thesis manuscript is divided into several chapters that match with different aspects of the embryogenic system. Somatic embryogenesis induction was achieved on maternal teguments of developing ovules from immature acorns of adult holm oak trees. Despite the low induction frequencies, the generated embryogenic lines were amplified by secondary embryogenesis. A decline in the differentiation capacity over time was also observed. It was concluded that both genotype and culture media formulation influenced the embryogenic response, being the Schenk and Hildebrandt´s macronutrients formulation from culture medium and the lack of plant growth regulators the most effective combination for the induction of the embryogenic response. It has been suggested the existence of a developmental window in which ovules are prone to induction. Genotype influenced the proliferation capacity and the plant conversion of somatic embryos, which was also favoured by the presence of indol-3-butyric acid and 6-bencyladenine. The use of temporary immersion systems as proliferation in liquid culture of the embryogenic lines increased the growth depending on genotype, when compared to semisolid cultures. However, it did not improve the differentiation of single cotyledonary embryos. A protocol for the initiation and maintenance of embryogenic suspension cultures was established for several embryogenic lines with highly dense inoculi of embryogenic clusters from proliferating semisolid cultures. In order to avoid the loss of vigour and morphogenic ability of embryogenic lines due to prolonged cultures, a cryopreservation protocol for embryogenic lines of holm oak has been developed. During the determination of the influence of cryoprotective agents on the growth and differentiation capacities before and after liquid nitrogen immersion, it was concluded that both responses were independent from each other and also under the influence of genotype and the type of cryopreserved material. The combination of sucrose and PVS2 prior liquid nitrogen immersion provided higher recovery rates. When the same embryogenic lines were cryopreserved for 30 days, none was able to differentiate. The SSRs analysis of the short-term cryopreserved material detected somaclonal variation. Both SSR and RAPD markers showed high sensitivity to detect genetic differences between the donor trees and the generated embryogenic material. Nevertheless, the degree of instability detection depended on the marker. The SSR analysis indicated a relationship between genotype, the studied loci and the located polymorphisms. Also, both markers revealed low intraclonal genetic variation. The RAPD detected genetic variation within the donor trees. The presence of pre-existent genetic variation within mature trees, in addition to its occurrence during the early stages of the embryogenic induction, and the presence of tissues of fertilisation origin within the maternal explants are all discussed. Nonetheless, the determination of the genetic identity of donor material is required, in addition to early detection methods of somaclonal variation.

Different sources of reduced carbon contribute to form three classes of terpenoid emitted by Quercus ilex L. leaves.

Quercus ilex L. leaves emit terpenes but do not have specialized structures for terpene storage. We exploited this unique feature to investigate terpene biosynthesis in intact leaves of Q. ilex. Light induction allowed us to distinguish three classes of terpenes: (i) a rapidly induced class including alpha-pinene; (ii) a more slowly induced class, including cis-beta-ocimene; and (iii) the most slowly induced class, including 3-methyl-3-buten-1-ol. Using 13C, we found that alpha-pinene and cis-beta-ocimene were labeled quickly and almost completely while there was a delay before label appeared in linalool and 3-methyl-3-buten-1-ol. The acetyl group of 3-methyl-3-buten-1-yl acetate was labeled quickly but label was limited to 20% of the moiety. It is suggested that the ocimene class of monoterpenes is made from one or more terpenes of the alpha-pinene class and that both classes are made entirely from reduced carbon pools inside the chloroplasts. Linalool and 3-methyl-3-buten-1-ol are made from a different pool of reduced carbon, possibly in nonphotosynthetic plastids. The acetyl group of the 3-methyl-3-buten-1-yl acetate is derived mostly from carbon that does not participate in photosynthetic reactions. Low humidity and prolonged exposure to light favored ocimenes emission and induced linalool emission. This may indicate conversion between terpene classes.

意大利威尼托（Veneto）的刺叶栎林-植物社会学生物多样性研究

刺叶栎( Quercus ilex L．)是地中海的常绿树种，属于古老的第三纪植物区系成份。本文根据苏黎世——蒙特利尔学派的植被学基本原理，全面地研究了分布在整个威尼托(Veneto)大区的剌叶栎林，详细研究刺叶栎林的群落组成、结构及类型划分。同时，对威尼托的刺叶栎林同北京山区的栎林进行了比较研究。最后，着重研究了威尼托刺叶栎林的生物多样性。 威尼托的刺叶栎林主要分布在亚得里亚海沿岸（包括Foci del Tagtiamento和Bosco Nordio e Rosolina Mare两地）、Lago di Garda和Colli Euganei。 在气候上，亚得里亚海沿岸属于半地中海气候。加尔达湖区域(Laqo di Garda)则是接近欧洲中部的大陆性气候，且维持半地中海的气候特点。Co t l i Euganei是这两地气候的过渡类型，且更接近亚得里亚海沿岸的类型。 在亚得里亚海沿岸的刺叶栎林可以分为三类，第一类(I)是一些矮树丛，这是Fraxinus ornus和Quercus ilex混交林的前身，较干旱。第二类群落(II)缺少乔木层，灌木层是由一些盖度不大的刺叶栎代替，更干旱。第三类群落(Ⅲ)是一群在外貌上相同的成熟群落．Fraxinus ornus和Quercus ilex得到充分发展，较中生。 在加尔达湖区的刺叶栎林可划分为三类。第一类群落(I)代表一组耐旱、开敞的矮树丛，含有Sesterio Variae-Ostryelum群丛的特征种，这一类可划分为SesLerio Variae-Ostryetum群 丛，土壤贫脊和干旱。第二类群落(Ⅱ)代表一类较郁闭的矮树丛，含有较多的Prunetalia群落目的成份，土壤较贫脊。第三类群(Ⅲ)代表一类郁闭的群落，乔木具有很大的密度因而林下灌木不能充分发展。SesLerio Variae-Ostryetum群丛和Prunetalia群落目的成份均不多。 在Colli Euganei的刺叶栎林可划分为二类。第一类群落(I)是一些不郁闲的矮树林组成。大体上分为地中海旱生栎林和地中海假灌丛。第二类群落(II)代表了较中生状态的植被，刺叶标种群绝对郁闭。 北京地处华北大平原的西北部。北京山地的气候为温带陆地性季风气候，其地带性的落叶阔叶林是以栎林为典型。虽然这些栎林同意大利威尼托刺叶栎林是两种不同的森林类型，但两者之间是存在着一定的联系。其共有的科有20个，共有属有11个。他们在植物组成中，以禾本科，蔷薇科和豆科的植物种类为最多。在乔木层中，他们都是以壳斗科的栎属(Quercus)为优势，其中木犀科的白蜡属(Fraxinus)和槭树属(Acer)较多。 本文对威尼托大区刺叶栎林的物种多度分布格局进行了全面探讨，计算出刺叶栎林的几何分布模型、Broken-stick分布模型、Log分布模型、Log-normat分布模型等四种物种多度分布的理论模型，并将这些理论分布模型用“序列／多度”图解和“多度／频度”图解表示出来。其中，几何分布模型．Broken-stick1分布模型用“序列／多度”图解表示。Broken-stick2分布模型、Log分布模型、Log-normal分布模型用“多度／频度”图解表示。 对上述四个物种多度的理论分布同实际现察的物种多度分布进行X2分析，在5％的显著性水平上，对整个威托大区的刺叶 栎林，几何分布模型最能代表其物种多度分布，显著性最大;Log-normal分布模型也可以用来代表威尼托刺叶标妹的物种多度分布，其显著性次于几何分布模型的显著性。这表明威尼托的刺叶栎林尚处于演替的早期阶段，这些刺叶栎林曾受到严重破坏，现正在恢复。 通过比较Foci del Tagtiamento和Bosco Nordio加尔达湖区、Colli Euganei四个地方刺叶栎林的物种多度的几何分布模型和Log-normal分布模型，显示出Lago di Garda（加尔达湖区）的刺叶栎林生物多样性最好、Foci del Tagtiamento刺叶栎林生物多样性较好．Bosco Nordio的剌叶栎林生物多样性较差．Colli Euganei的刺叶栎林生物多样性最差。 再利用多样性指数计算全部威尼托大区剌叶栎林的生物多样性。计算的多样性指数有丰富度指数（包括Margalef指数、Men-hinick指数、Monk指数）、多样性指数(Shannon信息指数、Bri llouin个息指数、Gini指数、PIE指数、Mcintosh指数)、优势度指数(Berger-Parker指数、Simpson指数)、均一度指数(Pielou均一度指数、Brillouin均一度指数，PIE的V’均一度指数．PIE的V均一度指数，N2的V’均一度指数，N2的V均一度指数，Mclntosh均一度指数，Hill的F10均一度指数，Hill的E21均一度指数，Hill的F21的一度指数）。通过比较丰富度指数，多样性指数、均一度指数与优势度之间的关系，结果，Simpson优势度指数同Men-hinick物种丰富度指数、Shannon信息指数、Bril-louin信息指数、Pielou均一度指数，Brillouin均一度指数，Mcintosh均一度指数、PIE的V’均一度指数呈负相关关系，因此，上述生物多样性指数可以较好地反映威尼托大区刺叶栎林的生物多样性。反映出的结果是：加尔达湖区刺叶栎林生物多样性最好，Foci del Tagtiamento刺叶栎林生物多样性较好，Bosco Nordio的剌叶栎林生物多样性较差．Colli Euganei的刺叶栎林生物多样性最差。 生物多样性的研究显示出生物多样性同生境状况的密切联系。往往受人为干扰严重的群落生物多样性低、如Coli Euganei和亚得里亚海岸刺叶栋林;而受人为破坏较轻的群落其生物多样性高，如加尔达湖区刺叶栎。 生物多样性的研究还显示出生物多样性同群落演替的发展阶段密切相关，在群落演替初期，由于缺乏优势种，而又有大量物种侵入，物多样性相对较高。在群落演替中期，由于形成了一个或几个优势种，优势度的增加导致了生物多样性相对减低。如Foci del Tagtiamento刺叶栎林生物多样性高于Bosco Nordio刺叶栎林的生物多样性。到演替后期，随着更多物种的侵入，群落结构的复杂化、生物多样性又将逐步提高。

The past ecology of Abies alba provides new perspectives on future responses of silver fir forests to global warming

Paleoecology can provide valuable insights into the ecology of species that complement observation and experiment-based assessments of climate impact dynamics. New paleoecological records (e.g., pollen, macrofossils) from the Italian Peninsula suggest a much wider climatic niche of the important European tree species Abies alba (silver fir) than observed in its present spatial range. To explore this discrepancy between current and past distribution of the species, we analyzed climatic data (temperature, precipitation, frost, humidity, sunshine) and vegetation-independent paleoclimatic reconstructions (e.g., lake levels, chironomids) and use global coupled carbon-cycle climate (NCAR CSM1.4) and dynamic vegetation (LandClim) modeling. The combined evidence suggests that during the mid-Holocene (6000 years ago), prior to humanization of vegetation, A. alba formed forests under conditions that exceeded the modern (1961-1990) upper temperature limit of the species by 5-7°C (July means). Annual precipitation during this natural period was comparable to today (>700-800 mm), with drier summers and wetter winters. In the meso-Mediterranean to sub-Mediterranean forests A. alba co-occurred with thermophilous taxa such as Quercus ilex, Q. pubescens, Olea europaea, Phillyrea, Arbutus, Cistus, Tilia, Ulmus, Acer, Hedera helix, Ilex aquifolium, Taxus, and Vitis. Results from the last interglacial (ca. 130 000-115 000 BP), when human impact was negligible, corroborate the Holocene evidence. Thermophilous Mediterranean A. alba stands became extinct during the last 5000 years when land-use pressure and specifically excessive anthropogenic fire and browsing disturbance increased. Our results imply that the ecology of this key European tree species is not yet well understood. On the basis of the reconstructed realized climatic niche of the species, we anticipate that the future geographic range of A. alba may not contract regardless of migration success, even if climate should become significantly warmer than today with summer temperatures increasing by up to 5-7°C, as long as precipitation does not fall below 700-800 mm/yr, and anthropogenic disturbance (e.g., fire, browsing) does not become excessive. Our finding contradicts recent studies that projected range contractions under global-warming scenarios, but did not factor how millennia of human impacts reduced the realized climatic niche of A. alba.

Natural regeneration in mediterranean mixed forests: effect of environmental conditions on the physiological optimum niche of the species = Regeneración natural en masas mixtas mediterráneas: efecto de las condiciones ambientales en el nicho óptimo fisiológico de las especies

Los montes Mediterráneos han experimentado múltiples cambios en las últimas décadas (tanto en clima como en usos), lo que ha conducido a variaciones en la distribución de especies. El aumento previsto de las temperaturas medias junto con la mayor variabilidad intra e inter anual en cuanto a la ocurrencia de eventos extremos o disturbios naturales (como periodos prolongados de sequía, olas de frío o calor, incendios forestales o vendavales) pueden dañar significativamente al regenerado, llevándolo hasta la muerte, y jugando un papel decisivo en la composición de especies y en la dinámica del monte. La amplitud ecológica de muchas especies forestales puede verse afectada, de forma que se esperan cambios en sus nichos actuales de regeneración. Sin embargo, la migración latitudinal de las especies en busca de mejores condiciones, podría ser una explicación demasiado simplista de un proceso mucho más complejo de interacción entre la temperatura y la precipitación, que afectaría a cada especie de un modo distinto. En este sentido tanto la capacidad de adaptación al estrés ambiental de una determinada especie, así como su habilidad para competir por los recursos limitados, podría significar variaciones dentro de una comunidad. Las características fisiológicas y morfológicas propias de cada especie se encuentran fuertemente relacionadas con el lugar donde cada una puede surgir, qué especies pueden convivir y como éstas responden a las condiciones ambientales. En este sentido, el conocimiento sobre las distintas respuestas ecofisiológicas observadas ante cambios ambientales puede ser fundamentales para la predicción de variaciones en la distribución de especies, composición de la comunidad y productividad del monte ante el cambio global. En esta tesis investigamos el grado de tolerancia y sensibilidad que cada una de las tres especies de estudio, coexistentes en el interior peninsular ibérico (Pinus pinea, Quercus ilex y Juniperus oxycedrus), muestra ante los factores abióticos de estrés típicos de la región Mediterránea. Nuestro trabajo se ha basado en la definición del nicho óptimo fisiológico para el regenerado de cada especie a través de la investigación en profundidad del efecto de la sequía, la temperatura y el ambiente lumínico. Para ello, hemos desarrollado un modelo de predicción de la tasa de asimilación de carbono que nos ha permitido identificar las condiciones óptimas ambientales donde el regenerado de cada especie podría establecerse con mayor facilidad. En apoyo a este trabajo y con la idea de estudiar el efecto de la sequía a nivel de toda la planta hemos desarrollado un experimento paralelo en invernadero. Aquí se han aplicado dos regímenes hídricos para estudiar las características fisiológicas y morfológicas de cada especie, sobre todo a nivel de raíz y crecimiento del tallo, y relacionarlas con las diferentes estrategias en el uso del agua de las especies. Por último, hemos estudiado los patrones de aclimatación y desaclimatación al frio de cada especie, identificando los periodos de sensibilidad a heladas, así como cuellos de botella donde la competencia entre especies podría surgir. A pesar de que el pino piñonero ha sido la especie objeto de la gestión de estas masas durante siglos, actualmente se encuentra en la posición más desfavorable para combatir el cambio global, presentado el nicho fisiológico más estrecho de las tres especies. La encina sin embargo, ha resultado ser la especie mejor cualificada para afrontar este cambio, seguida muy de cerca por el enebro. Nuestros resultados sugieren una posible expansión en el rango de distribución de la encina, un aumento en la presencia del enebro y una disminución progresiva del pino piñonero a medio plazo en estas masas. ABSTRACT Mediterranean forests have undergone multiple changes over the last decades (in both climate and land use), which have lead to variations in the distribution of species. The expected increase in mean annual temperature together with the greater inter and intra-annual variability in extreme events and disturbances occurrence (such as prolonged drought periods, cold or heat waves, wildfires or strong winds) can significantly damage natural regeneration, up to causing death, playing a decisive role on species composition and forest dynamics. The ecological amplitude for adaptation of many species can be affected in such a way that changes in the current regeneration niches of many species are expected. However, the forecasted poleward migration of species seeking better conditions could be an oversimplification of what is a more complex phenomenon of interactions among temperature and precipitation, that would affect different species in different ways. In this regard, either the ability to adapt to environmental stresses or to compete for limited resources of a single species in a mixed forest could lead to variations within a community. The ecophysiological and morphological traits specific to each species are strongly related to the place where each species can emerge, which species can coexist, and how they respond to environmental conditions. In this regard, the understanding of the ecophysiological responses observed against changes in environmental conditions can be essential for predicting variations in species distribution, community composition, and forest productivity in the context of global change. In this thesis we investigated the degree of tolerance and sensitivity that each of the three studied species, co-occurring in central of the Iberian Peninsula (Pinus pinea, Quercus ilex and Juniperus oxycedrus), show against the typical abiotic stress factors in the Mediterranean region. Our work is based on the optimal physiological niche for regeneration of each species through in-depth research on the effect of drought, temperature and light environment. For this purpose, we developed a model to predict the carbon assimilation rate which allows us to identify the optimal environmental conditions where regeneration from each species could establish itself more easily. To obtain a better understanding about the effect of low temperature on regeneration, we studied the acclimation and deacclimation patterns to cold of each species, identifying period of frost sensitivity, as well as bottlenecks where competition between species can arise. Finally, to support our results about the effect of water availabilty, we conducted a greenhouse experiment with a view of studying the drought effect at the whole plant level. Here, two watering regimes were applied in order to study the physiological and morphological traits of each species, mainly at the level of the root system and stem growth, and so relate them to the different water use strategies of the species. Despite the fact that stone pine has been the target species for centuries, nowadays this species is in the most unfavorable position to cope with climate change. Holm oak, however, resulted the species that is best adapted to tolerate the predicted changes, followed closely by prickly juniper. Our results suggest a feasible expansion of the distribution range in holm oak, an increase in the prickly juniper presence and a progressive decreasing of stone pine presence in the medium term in these stone pine-holm oak-prickly juniper mixed forests.

Paleobotánica del epipaleolítico y mesolítico vascos

Homenaje a Ignacio Barandiarán Maestu / coord. por Javier Fernández Eraso, Juan Santos Yanguas

Impacts of changing climate and land use on vegetation dynamics in a Mediterranean ecosystem: insights from paleoecology and dynamic modeling

Forests near the Mediterranean coast have been shaped by millennia of human disturbance. Consequently, ecological studies relying on modern observations or historical records may have difficulty assessing natural vegetation dynamics under current and future climate. We combined a sedimentary pollen record from Lago di Massacciucoli, Tuscany, Italy with simulations from the LandClim dynamic vegetation model to determine what vegetation preceded intense human disturbance, how past changes in vegetation relate to fire and browsing, and the potential of an extinct vegetation type under present climate. We simulated vegetation dynamics near Lago di Massaciucoli for the last 7,000 years using a local chironomid-inferred temperature reconstruction with combinations of three fire regimes (small infrequent, large infrequent, small frequent) and three browsing intensities (no browsing, light browsing, and moderate browsing), and compared model output to pollen data. Simulations with low disturbance support pollen-inferred evidence for a mixed forest dominated by Quercus ilex (a Mediterranean species) and Abies alba (a montane species). Whereas pollen data record the collapse of A. alba after 6000 cal yr bp, simulated populations expanded with declining summer temperatures during the late Holocene. Simulations with increased fire and browsing are consistent with evidence for expansion by deciduous species after A. alba collapsed. According to our combined paleo-environmental and modeling evidence, mixed Q. ilex and A. alba forests remain possible with current climate and limited disturbance, and provide a viable management objective for ecosystems near the Mediterranean coast and in regions that are expected to experience a mediterranean-type climate in the future.

Vegetation and fire history of coastal north-eastern Sardinia (Italy) under changing Holocene climates and land use

Little is known about the vegetation and fire history of Sardinia, and especially the long-term history of the thermo-Mediterranean belt that encompasses its entire coastal lowlands. A new sedimentary record from a coastal lake based on pollen, spores, macrofossils and microscopic charcoal analysis is used to reconstruct the vegetation and fire history in north-eastern Sardinia. During the mid-Holocene (c. 8,100–5,300 cal bp), the vegetation around Stagno di Sa Curcurica was characterised by dense Erica scoparia and E. arborea stands, which were favoured by high fire activity. Fire incidence declined and evergreen broadleaved forests of Quercus ilex expanded at the beginning of the late Holocene. We relate the observed vegetation and fire dynamics to climatic change, specifically moister and cooler summers and drier and milder winters after 5,300 cal bp. Agricultural activities occurred since the Neolithic and intensified after c. 7,000 cal bp. Around 2,750 cal bp, a further decline of fire incidence and Erica communities occurred, while Quercus ilex expanded and open-land communities became more abundant. This vegetation shift coincided with the historically documented beginning of Phoenician period, which was followed by Punic and Roman civilizations in Sardinia. The vegetational change at around 2,750 cal bp was possibly advantaged by a further shift to moister and cooler summers and drier and milder winters. Triggers for climate changes at 5,300 and 2,750 cal bp may have been gradual, orbitally-induced changes in summer and winter insolation, as well as centennial-scale atmospheric reorganizations. Open evergreen broadleaved forests persisted until the twentieth century, when they were partly substituted by widespread artificial pine plantations. Our results imply that highly flammable Erica vegetation, as reconstructed for the mid-Holocene, could re-emerge as a dominant vegetation type due to increasing drought and fire, as anticipated under global change conditions.