Flora Palinologica Italiana,Sezione Aeropalinologica — Specie esotiche - S 236:Cedrus libani A. Rich

Riassunto Nell'ambito del programma della Flora Palinologica Italiana, Sezione Aerobiologica, viene presentata la scheda palinologica diCedrus libani A. Rich. (specie esotica). Sono puntualizzate le differenze tra i parametri morfologici del polline «fresco» e acetolizzato.

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Summary According to the program «Palynological Italian Flora, Aeropalynological Section» the palynological card ofCedrus libani A. Rich. (exotic species) relevant to «fresh» and acetolyzed pollen grains is presented. The differences between the morphological parameters of fresh and acetolyzed pollen are pointed out.

     

Cedar forest communities in Western Antalya (Taurus Mountains,Turkey)

Abstract

The most extensive and magnificent forests of Cedrus libani A. Rich. in the world are on the Taurus Mountains in Turkey. They cover an area of about 400,000 hectares. C. libani forms either pure stands or mixed stands with other tree species, such as Abies cilicica, Pinus nigra, P. brutia, Quercus species (Q. cerris, Q. infectoria, Q. libani etc.), Juniperus excelsa and J. foetidissima. In this work, the floristic composition of the cedar forests in Western Antalya has been investigated and the ecological factors have been measured. The main forest types have been established and correlated with the ecological factors. In total, 201 relevés were analyzed by numerical methods and classified into five groups of C. libani forests and one group of Q. coccifera scrublands. It was observed that the topographical gradient mainly determines the differentiation patterns of cedar forests in Western Antalya.     

Gene flow among different taxonomic units: evidence from nuclear and cytoplasmic markers in Cedrus plantation forests

Hybridization and introgression are important natural evolutionary processes that can be successfully investigated using molecular markers and open- and controlled-pollinated progeny. In this study, we collected open-pollinated seeds from Cedrus atlantica, Cedrus libani and C. libani × C. atlantica hybrids from three French-plantation forests. We also used pollen from C. libani and Cedrus brevifolia to pollinate C. atlantica trees. The progeny were analyzed using three different types of molecular markers: RAPDs, AFLPs and cpSSRs. Chloroplast DNA was found to be paternally inherited in Cedrus from the progeny of controlled-crosses. Heteroplasmy, although possible, could not be undoubtedly detected. There was no indication of strong reproductive isolating barriers among the three Mediterranean Cedrus taxa. Gene flow between C. atlantica and C. libani accounted for 67 to 81% of viable open-pollinated seedlings in two plantation forests. We propose that Mediterranean Cedrus taxa should be considered as units of a single collective species comprising two regional groups, North Africa and the Middle East. We recommend the use of cpSSRs for monitoring gene flow between taxa in plantation forests, especially in areas where garden specimens of one species are planted in the vicinity of selected seed-stands and gene-conservation reserves of another species.Communicated by D.B. Neale     

Scale‐dependent intraspecific competition of Taurus cedar (Cedrus libani A. Rich.) saplings in the Southern Turkey

Better understanding of the competitive interaction at the early development stages of the stand is crucial to help schedule silvicultural treatments for young stands and for the better management of the future stands. We used scale‐dependent analysis to improve our understanding of sapling dynamics in the pure Taurus cedar (Cedrus libani A. Rich.) stands in Southern Turkey. Using data from nine plots established at the western Taurus Mountains, diameter, height, and crown radii of saplings were compared, and spatial point pattern analyses were performed. We found significant differences for the mean diameter and height, and crown radii of saplings among the plots. Univariate pair correlation function showed that sapling pattern was regular only at small scales (r < 0.4 m) but was predominantly random. Bivariate pair correlation function revealed no evidence of spatial interaction between tall saplings and short saplings. Univariate mark correlation function revealed that strong intraspecific competition was detected at small scales (up to 0.55 m). This distance is reasonable for the juvenile age tending of Taurus cedar saplings and should be under consideration during silvicultural treatments to use the site productivity more efficiently.     

Genetic consequences of past climate and human impact on eastern Mediterranean <Emphasis Type="Italic">Cedrus libani</Emphasis> forests. Implications for their conservation

Summary This study demonstrates the impact of natural factors and human activities on biodiversity at gene level on a keystone Mediterranean forest ecosystem species. We monitored the within and among population gene diversity of Cedrus libani, a forest tree species of the Eastern Mediterranean mountains. We used paternally inherited chloroplast microsatellites (57 haplotypes) and bi-parentally inherited isozymes (12 loci) to estimate allelic richness, heterozygosity, and differentiation in 18 natural and 1 planted populations from Turkey and Lebanon. We showed that there is a phylogeographic structure in C. libani, and that forests from Lebanon and Turkey constitute two genetically isolated groups which probably arose from distinct refugia after the last Quaternary glacial cycle. We found extensive gene flow and relatively low differentiation in Turkey, as well as little evidence of genetic drift within populations. However, one population we analyzed, which was planted more than 20 centuries ago, and is isolated from core populations in Turkey, demonstrated extremely low genetic diversity and deserves high conservation priority. In contrast, we found low gene flow, high differentiation and severe cases of genetic drift in Lebanon. As forests there are the remnants of millennia-long extensive deforestation, all deserve high conservation priority.     

A new fossil Cedrus species from the early Miocene of northwestern Turkey and its possible affinities

A new fossil species, Cedrus anatolica n. sp., is described from the early Miocene Hançili Formation of Turkey. All analyses were performed on the thin sections housed at Istanbul University – Cerrahpasa. The new species was interpreted as having the closest affinity with the modern Mediterranean species Cedrus atlantica (Endlicher) Manetti ex Carriere and Cedrus libani Richard. The evolutionary line shows some changes in wood anatomy. From the early Cretaceous to the early Miocene, the pits on the tangential walls of the tracheids gradually decreased, the height of rays increased and the number of epithelial cells in the traumatic resin canals increased slightly. These features are similar in three modern species; other wood anatomical features are also quite stable among the new fossil and modern species.     

In vitro storage of cedar shoot cultures under minimal growth conditions

We developed procedures for slow-growth storage of Cedrus atlantica and Cedrus libani microcuttings of juvenile and adult origin, noting factors favouring the extension of subculture intervals. Microcuttings could be stored effectively up to 6 months at 4°C and reduced light intensity, provided that they were grown on a diluted modified MS medium. The addition of 6% mannitol to the storage media affected negatively survival and multiplication capacity of the cultures. The slow-growth storage conditions used in our experiments did not induce remarkable effects on both RAPD variability and average DNA methylation in the species.     

Geographical diversity and genetic relationships among Cedrus species estimated by AFLP

Genetic diversity was described in 17 cedar populations covering the geographical range of the four species of the genus Cedrus. The study was conducted using amplified fragment length polymorphism (AFLP) on haploid tissues (megagametophytes). Eleven selective AFLP primer pairs generated a total of 107 polymorphic amplification products. Correspondence and genetic distance analyses indicated that Cedrus deodara constitutes a separate gene pool from the Mediterranean cedars. Within Mediterranean cedars, we distinguished two groups: the first one is made of Cedrus atlantica, while the second one is made of Cedrus libani and Cedrus brevifolia, these latter two species being genetically similar despite important divergence previously observed for morphological and physiological traits. The lowest intrapopulation variability was found in the two C. deodara populations analyzed. Surprisingly, C. brevifolia, the endemic taxon from the island of Cyprus that is found in small and fragmented populations, showed one of the highest levels of diversity. This unexpected pattern of diversity and differentiation observed for C. brevifolia suggests a recent divergence rather than a relictual, declining population. Patterns of diversity within- and among-populations were used to test divergence and fragmentation hypotheses and to draw conclusions for the conservation of Cedrus gene pools.     

Assessments of somaclonal variation in micropropagated shoots of <Emphasis Type="Italic">Cedrus</Emphasis>: consequences of axillary bud breaking

Reversed-phase HPLC analysis and random amplified polymorphic DNA (RAPD) markers were used to monitor DNA methylation status and genetic stability of C. atlantica and C. libani shoots generated through axillary bud proliferation. Average DNA methylation in C. atlantica or C. libani seedlings and mature 200-year-old trees of C. libani was 19.8, 19.5 and 22.3%, respectively. These global amounts showed no significant variation after the in vitro establishment of seedling-originated shoot stocks. In contrast, in vitro culture caused a significant decrease in the amount of 5-methylcytosine in genomic DNA of the tissue culture (TC) progenies of one of the adult C. libani genotypes. This DNA demethylation event accompanied an enhancement of the regrowth capacity of this genotype. Detected RAPD variation between mother plants and their TC progenies was species-related, with C. libani TC progenies being genetically more stable than those of C. atlantica. Nevertheless, similarity indices ranged from 0.97 to 1 among mother plants and their TC progenies. Furthermore, the analyses of molecular variance (AMOVA) suggest that RAPD variation among the mother plants and their TC progenies might be considered as not significant. The application of various statistical approaches, including cluster-based genetic distance methods and AMOVA, demonstrates that RAPD markers discriminate C. atlantica and C. libani appropriately.     

Karyotype analysis reveals interspecific differentiation in the genus Cedrus despite genome size and base composition constancy

The DNA content and GC% of the four true cedar (Cedrus) species, C. atlantica, C. brevifolia, C. deodara and C. libani, were assessed. Genome size was homogeneous among representative populations of the four species with an average of 32.6±0.6 pg per 2 C or 15.7×10⁹ base pairs per 1 C. The composition in GC was calculated to be 40.7%. A simple monosomatic haploid level was found in the megagametophyte, as compared to the diploid level of the corresponding embryo. Cytogenetic studies showed a diploid chromosome number of 2n=2x=24 in 11 populations sampled over the four species. The chromosome complements have similar morphology and symmetry. However, fluorochromes revealed specific banding patterns in each of the four cedar species. Eight GC-rich chromomycin A₃ bands were observed in Cedrus deodara chromosomes, six in both Cedrus libani and Cedrus brevifolia, and four bands in Cedrus atlantica chromosomes. Moreover, Hoechst 33258 fluorochrome revealed AT-rich sequences specifically located in the centromeric regions while the GC-rich sequences appeared negatively stained. These investigations provide a systematic characterisation of the Cedrus genus and should contribute towards clarification of the phylogenetic relationships among the four species. Received: 10 October 2000 / Accepted: 20 March 2001     

Modeling above‐ground litterfall in eastern Mediterranean conifer forests using fractional tree cover,and remotely sensed and ground data

Question: How can we model above‐ground litterfall in Mediterranean conifer forests using remotely sensed and ground data, and geographic information systems (GIS)? Location: Eastern Mediterranean conifer forest of Turkey. Methods: Above‐ground litterfall from Mediterranean forest stands of Pinus nigra, Cedrus libani, Pinus brutia and Juniperus excelsa and mixed Abies cilicica, C. libani and P. nigra was modeled as a function of fractional tree cover using a regression tree algorithm, based on IKONOS and Landsat TM/ETM+data. Landsat TM/ETM+images for the study area were used to map actual stand patterns, based on a land‐cover map of species stands using a supervised classification. Results: Total amount of annual above‐ground litterfall for the entire study area (12 260 km²) was estimated at 417.2 Mg ha^?1 for P. brutia, 291.1 Mg ha^?1 for the mixed stand, 115.5 Mg ha^?1 for P. nigra, 54.6 Mg ha^?1 for J. excelsa and 45.9 Mg ha^?1 for C. libani. The maps generated indicate the distribution of the seasonal amount of total above‐ground litterfall for different species and the distribution of species stands in the study area. There was an increase in the amount of above‐ground litterfall for P. brutia stand in summer, for J. excelsa in autumn and for C. libani, P. nigra and the mixed stand of A. cilicica, P. nigra and C. libani in winter. Conclusion: Application of this model helps to improve the accuracy of estimated litterfall input to soil organic carbon pools in the Mediterranean conifer forests.     

Competition between Exotic and Native Insects for Seed Resources in Trees of a Mediterranean Forest Ecosystem

The seeds of both cedar-of-Lebanon (Cedrus libani) and Cyprus cedar (Cedrus brevifolia) are attacked in their natural range by a specialised chalcid, Megastigmus schimitscheki. From 1995 to 1999, seeds were screened for insect damage in the main cedar plantations of southern France, as well as in the stands where cedar is mixed with firs (Abies spp.). X-rays were used to identify chalcid-infested seeds from which the insects were then reared. The surveys revealed the presence of M. schimitscheki in all the stands of Atlas cedar, Cedrus atlantica, planted at Mt Ventoux, southeastern France. The chalcid also infested seeds of an exotic fir, Abies pinsapo, planted in the same area. However, it has not yet reached the cedar plantations in southwestern France, where the seeds are colonised by a related exotic insect, Megastigmus pinsapinis, originating from North Africa. The latter species was common in cedar seeds at Mt Ventoux in the early 1990s but seems to have been supplanted by M. schimitscheki in the invasion zone. A native chalcid species, Megastigmus suspectus, was also shown to have shifted to a slight extent from a native fir, A. alba, onto cedar. The presence of three chalcid species competing for cedar seed resources may result in a substantial decline of the regeneration potential of that tree species. At Mt Ventoux, up to 92.6% of the cedar seeds were attacked, with 86.8% due to M. schimitscheki. The survey also revealed the widespread presence of another North American chalcid, Megastigmus rafni, in the fir stands of southern France. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cedrus libani (A. Rich) distribution in Lebanon: Past,present and future

Long-term vegetation studies are needed to better predict the impact of future climate change on vegetation structure and distribution. According to the IPCC scenario, the Mediterranean region is expected to undergo significant climatic variability over the course of this century. Cedrus libani (A. Rich), in particular, is currently distributed in limited areas in the Eastern Mediterranean region, which are expected to be affected by such climate change. In order to predict the impact of future global warming, we have used fossil pollen data and model simulations. Palaeobotanical data show that C. libani has been affected by both climate change and human activities. Populations of C. libani survived in refugial zones when climatic conditions were less favourable and its range extended during periods of more suitable climate conditions. Simulations of its future geographical distribution for the year 2100 using a dynamic vegetation model show that only three areas from Mount Lebanon may allow its survival. These results extrapolated for cedar forests for the entire Eastern Mediterranean region show that forests in Syria are also threatened by future global warming. In southern Turkey, cedar forests seem to be less threatened. These results are expected to help in the long-term conservation of cedar forests in the Near East.     

Antimicrobial Activity of Resins Obtained from the Roots and Stems of Cedrus libani and Abies cilicia

The antimicrobial activity of the ethanol extract of resins obtained from the roots and stems of Cedrus libani and Abies cilicia has been studied, using the disc diffusion method. The results obtained indicate that crude extracts of the resins of both plants are highly efficient in preventing the growth of microorganisms.     

Genetic variability and structure of the remnant natural populations of <Emphasis Type="Italic">Cedrus libani</Emphasis> (Pinaceae) of Lebanon

Cedrus libani of Lebanon is a valuable natural resource and the dominant species in its natural ecosystem. Intense and diverse anthropogenic pressures over historical times raised concerns about its genetic vigor and continued survival. Our investigation of the genetic diversity included samples from all remnant natural populations. Assessment of the genetic diversity using random amplified polymorphic DNA markers revealed the persistence of considerable variation distributed within populations with low population differentiation corroborated by Bayesian and analysis of molecular variance estimates (G _ST = 0.07, Φ _ST = 0.09). Individual assignment tests were carried out to investigate measures of gene flow. Inferences concluded that this natural heritage is not currently threatened by inbreeding or by random genetic drift. Correlation studies investigated possible effects of spatial distribution and environmental conditions on genetic structure. A climatic trend corresponding to a temperature–humidity gradient correlated significantly with the level of genetic diversity, while the edaphic variation did not.     

Conservation of the Cedrus libani populations in Lebanon: history, current status and experimental application of somatic embryogenesis

Cedrus libani, the cedar of Lebanon, is a threatened conifer native to the Levant. Over 4000 years of exploitation have resulted in the fragmentation and degradation of the Lebanese cedar populations. Continued urban and agricultural development in Lebanon adds to the difficulty of effective conservation. Two protected areas have recently been established which contain two of the more important forests: a cedar dominated forest in the Shouf region and a mixed forest at Ehden. A number of other populations are protected by ministerial decrees, and there is a need for rigorous management of all the remaining populations. The application of in vitro techniques such as somatic embryogenesis may assist in the conservation of this species. We have produced somatic pro-embryos using immature zygotic tissue as explants cultured on half-strength MS medium containing an auxin and a cytokinin (10 M 2,4-D and 5 M BAP). The application of somatic embryogenesis to the Lebanese cedar would be in the propagation and preservation of selected genotypes, either those from old growth provenance for use in restoration, or those with desirable commercial or horticultural characteristics.     

Traditional Knowledge and Management of <Emphasis Type="Italic">Feijoa (Acca sellowiana)</Emphasis> in Southern Brazil

Traditional Knowledge and Management of Feijoa (Acca sellowiana) in Southern Brazil. This paper investigates traditional knowledge of the use and management of Acca sellowiana in southern Brazil. Fifty-six informants from three rural communities were assigned to one of four subgroups (“maintainers,” “managers,” “cultivators,” or “users”) based on their responses regarding management and use of A. sellowiana. Traditional knowledge related to use of this species is widespread among rural residents, but traditional knowledge related to management is fragmented depending on whether one uses, manages, or cultivates the species. Knowledge held in rural communities suggests that A. sellowiana could play an expanded role in local economies as well as biodiversity conservation. We suggest that participatory research could stimulate greater local use as well as on-farm conservation of A. sellowiana.     

Reclamation of abandoned magnesite site using amendments and plantation

This study aimed to determine the suitable reclamation ways of post-mining magnesite. It was hypothesized that adding amendments and planting would decrease pH, increasing the nutrient availability and enhancing the soil microbial activity. Treatments included 200 g gypsum, 2 kg goat manure, and 200 g sulfur per square meter. Cedar (Cedrus libani A. Rich.), black locust (Robinia pseudoacacia L.), black pine (Pinus nigra J.F. Arnold), and sainfoin (Onobrychis sativa L.) were planted. Before and after the treatments, samples were taken from each experimental unit and analyzed. Analysis of the data indicated that the pH had decreased by 0.6 and 0.5 units on the gypsum + sulfur- and gypsum + manure-treated experimental units, respectively, compared to the other treated plots. In addition, using manure significantly increased the organic matter N, P, and K content, by 3-fold, 2.5-fold, 7.5-fold, and 2.5-fold, respectively, and enhanced microorganism activity. The sulfur + manure treatment increased microbial C and N approximately twofold. Because the manure, sulfur, and gypsum treatments improved the tailings conditions for plant growth, the use of these amendments for mine tailing reclamation was strongly recommended. Across all treatments, cedar and black pine demonstrated 83 and 77% survival rates for the first 3 years of afforestation, respectively. For the same period, the black locust survival rate was 35% lower than that of the other species. The gypsum + manure application increased seedling height growth by about 20% for all three species compared to the other treatments.     

Effects of soil and air drought on growth,plant water status and leaf gas exchange in three Mediterranean cedar species: <Emphasis Type="Italic">Cedrus atlantica</Emphasis>, <Emphasis Type="Italic">C. brevifolia</Emphasis> and <Emphasis Type="Italic">C. libani</Emphasis>

Three- and four-year-old potted, greenhouse-grown cedar seedlings were subjected to two different watering regimes: half received full water supply and the other half was submitted to moderate drought (50% of the full water supply). Height growth was the greatest for C. atlantica and the most-limited for C. brevifolia in the well-watered set. However, in the dry set, height growth was less affected by drought conditions for C. brevifolia than for C. atlantica. Cedrus libani gave intermediate results for both watering regimes. Moderate drought provoked a decrease in osmotic potential at full leaf turgor and a long-lasting osmotic adjustment. When irrigation was withheld completely to induce severe soil drying, gas exchange decreased and then stopped at predawn water potentials of −3.0 MPa for C. brevifolia, between −2.6 and −2.8 MPa for C. libani, and at −2.4 MPa for C. atlantica, irrespective of watering regime. For all species, the dry set showed lower net photosynthesis (A) and stomatal conductance (g _s) than the plants in the well-watered set. A and g _s responded to variations in atmospheric water-vapour pressure deficit (VPD). As VPD increased, A and g _s decreased, and this trend was proportionate to initial values at low VPD, but remained independent of previous watering treatments, plant water status or species. To conclude, C. brevifolia appears to be a species with limited growth potential but strong soil drought tolerance whereas C. atlantica has strong growth potential when an adequate water supply is available but is more sensitive to soil drought. C. libani shows an intermediate behaviour for growth and drought tolerance.     

Flavonoids in plant nuclei: detection by laser microdissection and pressure catapulting (LMPC), in vivo staining, and uv–visible spectroscopic titration

Previous studies in our laboratory have indicated that the nuclei of a number of trees are associated with flavonoids, especially flavan‐3‐ols. In the present study, three techniques were applied to verify that flavonoids are naturally incorporated into nuclei. These were histochemistry, UV–visible (UV‐VIS) titration and laser microdissection. Nuclei from intact seed wings of Tsuga canadensis were isolated from their cells using laser microdissection and pressure catapulting (LMPC). Thereafter, the excised nuclei were stained with p‐dimethylamino‐cinnamaldehyde (DMACA), which resulted in a blue coloration due to the presence of flavanols. Thus, there is no doubt that the nuclei were, prior to staining, associated with flavanols. The nuclei of the coniferous species Abies lasiocarpa, Cedrus deodara, Cedrus libani, Juniperus communis, Picea abies, Picea orientalis and Pseudotsuga menziessii(Douglas fir) showed a yellow fluorescence typical for flavonols from the beginning of bud break over the entire growing season. However, after the bud‐breaking period, the nuclei of all species, except for Cedrus deodara, showed additionally a blue reaction for flavanols. Rather late, in midsummer, blue‐stained flavanols in nuclei were found in Picea orientalis. Generally, zeatin intensified the flavanol association with the nuclei. The main components of nucleosomes are DNA and the histone proteins. The nature of their association with the flavonols quercetin and rutin was investigated by UV‐VIS spectroscopic titration. The data were evaluated by means of the Mauser (A and AD) diagrams. The results indicate that DNA shows largely no spectroscopically detectable association equilibria under the experimental conditions chosen. However, association (aggregation) equilibria can be observed with rutin or quercetin and histone sulphate in Tris buffer (pH 8.0, 7.4 and 7.0). In phosphate buffer, rutin shows spectroscopically no or only weak association with histone sulphate, in contrast to its behaviour towards quercetin.