Modelling Mediterranean oak palaeolandscapes using the MaxEnt model algorithm: The case of the NE Iberia under the Middle Holocene climatic scenario

The Mediterranean Basin is a global biodiversity hotspot, and oak tree species play an important role in it. Since the beginning of the Holocene (∼11.4 kyr BP), the distribution of forests has not occurred uniformly, resulting in diverse types of vegetation landscapes. In this study, we used a maximum entropy algorithm (MaxEnt) to obtain the ecological niche model (ENM) of two sub-Mediterranean oak species, Quercus pubescens Willd. (pubescent oak) and Quercus ilex subsp. ilex (holm oak), both in the present day in the Iberian Peninsula and within a Middle Holocene (8.2–4.2 kyr BP) climatic scenario in the NE Iberian Peninsula. Moreover, we used the locations of Neolithic archaeological sites containing anthracological data to analyze the relationship between human occupations and oak habitats. Our results suggest that the two oaks have responded differently to the climatic conditions that have occurred, and show changes in both potential distributions. The palaeolandscape vegetation map shows a denser vegetation cover in the lowlands and a more open landscape in the highlands, with a higher dominance of Quercus pubescens in the septentrional areas, while Quercus ilex was more restricted to certain coastal areas. Temperature and precipitation factors, mainly seasonal climatic conditions, have had a greater impact on the distribution of vegetation than other factors. We found a good overlap between the ENM of the two oaks and the locations of the Neolithic sites analysed, and determined that the distribution of Neolithic archaeological sites is not random. The Neolithic populations in the study area depended heavily on the resources of the deciduous and evergreen sub- Mediterranean forest, although they also exploited the resources of the mountain pine forest. Neolithic sites distribution suggests that Neolithic human groups were aware of the potential of forests and probably gathered woody resources in their surroundings.     

Natural hybridisation between kermes (Quercus coccifera L.) and holm oaks (Q. ilex L.) revealed by microsatellite markers

Hybridisation between species of the genus Quercus is a common phenomenon as a result of weak reproductive isolation mechanisms between phylogenetically close species that frequently co-occur in mixed stands. In this study, we use microsatellite markers to analyse introgression between kermes ( Quercus coccifera L.) and holm ( Q. ilex L.) oak, two closely related taxa that frequently dominate the landscape in extensive areas in the Mediterranean region. All tested microsatellites amplified and were polymorphic in both kermes and holm oaks. Bayesian admixture analyses showed a good correspondence between each species and one of the two inferred genetic clusters. Five sampled individuals were a priori tentatively identified as hybrids on the basis of intermediate morphological characteristics, and it was confirmed that they also presented mixed genotypes. However, we also detected different levels of genetic introgression among morphologically pure individuals, suggesting that successful backcrossing and/or reduced phenotypic expression of genetic variance in certain individuals may have resulted in strong convergence towards a single species phenotype.     

Frequent cytoplasmic exchanges between oak species that are not closely related: Quercus suber and Q. ilex in Morocco

Chloroplast (cp) and mitochondrial (mt) DNA variation were studied in 97 populations of cork oak (Quercus suber) in Morocco; in 31 of these populations, holm oak (Quercus ilex), a clearly distinct species, also occurred and was compared with Q. suber. Three cpDNA and one mtDNA primer pairs were used in the survey, each in combination with one restriction enzyme. Six haplotypes belonging to two very divergent lineages were detected; one lineage predominates in each species, and is probably ancestral, as inferred from comparisons with other oak species. In the mixed-species populations, cytoplasmic genomes were frequently shared across species, as indicated by an introgression ratio of 0.63. This index is a new measure of the propensity of species to share locally genetic markers, varying from zero (complete differentiation) to one (no differentiation). By contrast, more closely related deciduous oak species (Q. robur, Q. petraea and Q. pubescens) have introgression ratios varying from 0.82 to 0.97. The introgression events appear to have been more frequent in the direction Q. ilex (female) x Q. suber (male), a finding which seems attributable to the flowering phenology of these two species. This asymmetry may have favoured immigration of Q. suber beyond its main range, in regions already colonized by Q. ilex. There, rare hybridization and further introgression through long distance pollen flow have established populations that are morphologically indistinguishable from Q. suber but that have cytoplasmic genomes originating from the local Q. ilex populations.     

High variability of chloroplast DNA in three Mediterranean evergreen oaks indicates complex evolutionary history

Chloroplast DNA variation was studied in three evergreen Quercus species (Q. suber L., Q. ilex L. and Q. coccifera L.) from the Western Mediterranean Basin using PCR-RFLP. We studied five primer pair/enzyme combinations, four of them previously used in other European Quercus, obtaining a large number of haplotypes (81) grouped in three main types (suber type, ilex-coccifera I type and ilex-coccifera II type). Such level of haplotype diversity is higher than previously reported for the genus. Remarkable differences in haplotype richness between species have been found. Q. ilex and Q. coccifera usually share the same haplotypes, while a number of Q. suber populations possesses variants of the ilex-coccifera I type. This fact is interpreted as a result of genetic introgression between Q. suber and Q. ilex. Reproductive factors that could determine this exchange are discussed, as well as the influence of different species histories on the present structure of evergreen Quercus in the Western Mediterranean Basin.     

Soil persistence and biodiversity of ericoid mycorrhizal fungi in the absence of the host plant in a Mediterranean ecosystem

The occurrence of suitable mycorrhizal inocula may be an important factor affecting the dynamics of plant communities. We investigated the persistence and diversity of ericoid mycorrhizal fungi in the soil of a mature Quercus ilex forest where ericaceous hosts were absent. Erica arborea was used as a bait plant and results were compared to soil samples from experimental plots where cuttings had allowed reappearance of this ericaceous species. Fungal endophytes were isolated and tested in mycorrhiza resynthesis trials. Sterile mycorrhizal endophytes were assigned to morphotypes whose consistency was confirmed by ITS-RFLP. The ITS region of a representative of each morphotype was sequenced. BLAST searches and Neighbour-Joining analysis indicated taxonomic affinities with different classes within Ascomycota. Our results indicate that ericoid mycorrhizal fungi persist and maintain mycorrhizal ability in habitats lacking the ericaceous host. Their persistence could favour the establishment of E. arborea seedlings in pure Q. ilex forests after disturbance phenomena.     

Variation in the herb species response and the humus quality across a 200-year chronosequence of beech and oak plantations in Belgium

The present study aimed at exploring the long-term impact of pure and mixed beech Fagus sylvatica and oak Quercus robur stands on the forest floor by documenting changes in the herb species' behaviour and in humus index across a 200-yr chronosequence of forest stands. The research was conducted in central Belgium, in a 4383 ha beech-dominated forest. Analyses were carried out in stands which are replicated, of the same age, managed in the same way, and growing on the same soil type with the same land-use history. The results of this study indicate that stand aging is an important determinant of herb species occurrence in the studied area. Most of the species studied show a different response to stand age in pure compared to mixed stands. Our results clearly show a decrease of the humus quality with age in pure stands (beech as well as oak). On the other hand, we found that mixing beech and oak maintained or improved the humus status along the chronosequence according to the proportion of each tree. So the addition of some oak to the beech made it possible to keep a constant quality of the humus. We found that, even if the understory tree species is very scarce, it may be sufficient to maintain the humus status on the long term. In the present study, a cover of 1% oak in a beech stand was sufficient to show an effect of the minor species on these soils. This pattern contrasts with the widespread idea that substantial effects of the minor tree species on soils might not develop if the ratio of major/minor species is low.     

Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

Tree mortality of five major species on Hokkaido Island, northern Japan

The mortality rates of five major tree species (Abies sachalinensis, Acer mono, Magnolia obovata, Quercus crispula, Tilia japonica) on Hokkaido Island, northern Japan were modeled using logistic regressions based on the long-term observation of 8929 individuals in 65 permanent plots. Individual size, recent growth, one-sided and two-sided interactions with neighbors, climatic and topographic factors were used as independent variables in the logistic regressions, and relevant variables were selected using the stepwise method. For all species analyzed, the final models significantly explained tree mortality rates. The regression analyses indicated that individual size and/or recent growth had effects on tree mortality. A significant effect of one-sided interaction on tree mortality was detected for three species, and a significant effect of two-sided interaction was detected for two species. The interactions with neighbors were not necessarily competitive. For all species analyzed, climatic and topographic factors affected tree mortality. The mortality models obtained can be used for forest dynamics simulations. One ecological question was examined using these models. Species that can grow fast in forest stands with smaller basal areas tended to have high mortality rates in forest stands with larger basal areas. Some ecological characteristics of the analyzed species are described based on the results of the regressions.     

高山栎叶的形态结构及其与生态环境的关系

 对分布在不同生态环境条件下的硬叶常绿阔叶林建群种高山栎组植物叶的形态解剖特征进行了比较研究。结果表明,高山栎组植物叶的形态结构兼有非肉质旱生植物和高山植物的性状,这是它们对生态环境广泛适应的基础。每一个种除气孔器类型及腹面二层表皮细胞性状比较稳定外,其他性状,如气孔的大小和密度、表皮细胞的形状、垂周壁的类型、叶片的厚度、角质膜的厚度均受环境条件修饰,不是稳定性状,不具分类学意义。对同一个种的不同生活型,如乔木、小乔木和灌木,在解剖结构土没有明显的区别特征。地中海的冬青栎(Quercusilex)的表皮特征和高山栎组植物有很大的相似性。     

Contrasting nuclear and cytoplasmic exchanges between phylogenetically distant oak species (Quercus suber L. and Q. ilex L.) in Southern France: inferring crosses and dynamics

Gene flow is particularly frequent in the genus Quercus (oaks), especially between closely related species. We focus here on Quercus ilex and the cork-producing Quercus suber , which occasionally hybridize although they are phylogenetically markedly separated. Morphological observations were combined with both allozymic and chloroplastic diagnostic markers to characterize hybridization and introgression and to infer their dynamics in two French regions (French Catalonia and Provence), which are separated by several hundred kilometres. Some hybrids were found in both regions, indicating recent hybridization events. As expected from previous studies, most hybrids resulted from ♀ Q. ilex  × ♂ Q. suber crosses, but our data showed that the reciprocal cross is also possible. Partial independence between nuclear and chloroplastic introgression was observed in the two species. Nuclear introgression was limited in both species and both regions, with no preferred direction. In Provence, chloroplastic introgression was very rare in both species. Conversely, all Q. suber individuals from French Catalonia were introgressed by Q. ilex chlorotypes. This might be explained by introgression in the Iberian Peninsula antedating the first occurrence of the two species in French Catalonia. We also observed a new chlorotype that was created locally, and was exchanged between the two species. However, the two species still remain genetically differentiated. The dynamics and complexity of exchanges and the factors determining them (including human management of Q. suber ) are discussed.     

Implications of foliar terpene content and hydration on leaf flammability of Quercus ilex and Pinus halepensis

We investigated the implications of foliar hydration and terpene content on leaf flammability in two widely distributed forest species of the Mediterranean basin, Quercus ilex, which does not store terpenes, and Pinus halepensis, a terpene-storing species. The experiments were carried out in plants grown under different water regimes that generated a wide range of foliar hydration and terpene contents. We monitored the temperatures and time elapsed to reach the smoke, pyrolysis and flame phases. Smoke appeared much earlier (37 versus 101 s) and at lower temperatures (96 versus 139 degrees C) in Quercus ilex than in Pinus halepensis. Quercus ilex reached pyrolysis earlier than Pinus halepensis (278 versus 338 s) but at the same temperature (365-371 degrees C). There were no significant differences in time elapsed nor in temperature for flammability (386-422 s; 505-487 degrees C in both species). Quercus ilex had lower water hydration than Pinus halepensis (41 versus 100%) and the leaf content of terpenes in Quercus was three orders of magnitude lower. The results of this study show no differences in the flame phase between the two species and the absence of a significant relationship between temperature and elapsed time of the different flammability phases in relation to monoterpene content; thus indicating that the role of monoterpenes in flammability phases is smaller than that of the water content. This, however, does not exclude the effects of terpene content on plant combustibility and fire propagation once fires start.     

Unusually limited pollen dispersal and connectivity of Pedunculate oak (Quercus robur) refugial populations at the species' southern range margin

Low‐latitudinal range margins of temperate and boreal plant species typically consist of scattered populations that persist locally in microrefugia. It remains poorly understood how their refugial habitats affect patterns of gene flow and connectivity, key components for their long‐term viability and evolution. We examine landscape‐scale patterns of historical and contemporary gene flow in refugial populations of the widespread European forest tree Pedunculate oak (Quercus robur) at the species' southwestern range margin. We sampled all adult trees (n = 135) growing in a 20 km long valley and genotyped 724 acorns from 72 mother trees at 17 microsatellite loci. The ten oak stands that we identified were highly differentiated and formed four distinct genetic clusters, despite sporadic historical dispersal being detectable. By far most contemporary pollination occurred within stands, either between local mates (85.6%) or through selfing (6.8%). Pollen exchange between stands (2.6%) was remarkably rare given their relative proximity and was complemented by long‐distance pollen immigration (4.4%) and hybridization with the locally abundant Quercus pyrenaica (0.6%). The frequency of between‐stand mating events decreased with increasing size and spatial isolation of stands. Overall, our results reveal outstandingly little long‐distance gene flow for a wind‐pollinated tree species. We argue that the distinct landscape characteristics of oaks' refugial habitats, with a combination of a rugged topography, dense vegetation and humid microclimate, are likely to increase plant survival but to hamper effective long‐distance pollen dispersal. Moreover, local mating might be favoured by high tree compatibility resulting from genetic purging in these long‐term relict populations.     

甘肃小陇山森林植被碳库及其分配特征

为准确估计甘肃小陇山林区森林植被的碳库大小,应用干烧法对该地区主要林分类型的13种乔木、14种灌木、10种草本植物的不同器官和7类林分的枯落物有机含碳率进行了测定,同时利用生物量标准地资料对8类林分的乔木层平均含碳率及森林植被的储碳密度和碳储量进行了估算,并分析了林分各组分的碳储量分配特征.结果表明:锐齿栎、油松、栓皮栎、白桦、红桦、日本落叶松、华山松、云杉、秦岭冷杉、水曲柳、大叶椋子木、五角枫、辽东栎13种乔木树种的器官平均含碳率范围为0.4501～0.5049,14种灌木和10种草本的器官平均含碳率分别为0.4446和0.3270,7类林分枯落物平均含碳率为0.4221.该地区8类林分的乔木层平均含碳率范围为0.4676～0.4976;小陇山林区森林植被层平均储碳密度为39.4254 t hm-2,总碳储量为13.3579 Tg.8类林分总碳储量分配中,乔木层占98.07%±0.73%,灌木层占1.38%4±0.43%,草本层占0.17%4±0.08%,枯落物层占0.37%±0.37%.甘肃小陇山8类林分乔木层的平均储碳密度值与我国及世界各地森林平均储碳密度的一些估计值基本接近.     

崖柏群落优势乔木树种种间关系

采用样方调查法,对组成崖柏群落的乔木树种进行调查。通过计算重要值确定崖柏群落优势乔木树种,研究优势树种的总体联结性、2物种间的联结性和种间协变。结果表明:崖柏群落内的15个优势树种总体间存在负关联;2物种间具有显著正联结的种对有:高山栎-川陕鹅耳枥、高山栎-华中八角、大叶青冈-乌岗栎、铁杉-华西花楸、华千金榆-青榨槭、华千金榆-大叶青冈、大叶青冈-川鄂山茱萸、川陕鹅耳枥-华中八角;呈显著负联结的种对是高山栎-青榨槭;崖柏与其他优势树种的联结性均未达到显著程度;有26个种对表现出明显的正协变,12个种对表现出明显的负协变;种对正协变的存在是由于这些物种对环境资源的利用具有相似性所致。     

Evaluation of secondary dispersal in a large-seeded tree Aesculus turbinata: a test of directed dispersal

Gap characteristics and gap phase replacement of major tree species were examined in two primary old-growth (mean DBHs of the canopy trees were 45.2 and 56.1 cm) and four secondary developing (range of mean DBH of the canopy trees was 23.5–39.9 cm) beech (Fagus crenata) stands in the Daisen Forest Reserve, southwestern Japan, and these were analyzed in relation to stand development as expressed by the difference of mean DBH of canopy trees. Percentage gap area (percentage of total gap area to total surveyed area) and mean and maximum gap size varied widely and ranged from 1.7 to 20.0%, from 19.4 to 162.8 m2, and from 35.7 to 585.1 m2, respectively. Mean percentage gap area and mean gap size were significantly greater in old-growth than in developing stands. However, they and maximum gap size might not increase linearly with stand development, and the gap feature of less developed stands was greater than that of later stages in developing stands. The cause was a higher formation rate, in younger developing stands, of gaps formed by simultaneous death (multiple trees falling down in domino fashion) which tends to produce larger gaps. In developing stands mean DBH of gapmakers (canopy trees that formed a gap) was smaller than that of canopy trees, though the inverse trend might be found in old-growth stands. Three typical types of death or injury states of gapmakers such as standing dead, trunk broken and uprooted were found in every study stand and the difference in stand development may not cause stand-to-stand variations for them. Importance of F. crenata (the most dominant species) in the canopy layer increased and its importance in the understory layer decreased with stand development. Shade-intolerant Quercus mongolica in the canopy layer was more important in younger than in old-growth stands, and there was no Quercus regeneration in old-growth stands. Acer mono consistently appeared, though in much less abundance than other species, in both canopy and understory layers of all study stands. Sub-canopy layers, which are mainly formed by sub-canopy tree species such as Acanthopanax sciadophylloides and Acer japonicum, may gradually develop with stand development.p>     

Radial growth response of Pinus densiflora and Quercus spp. to topographic and climatic factors in South Korea

Aims This study aimed to develop radial growth models and to predict the potential spatial distribution of Pinus densiflora (Japanese red pine) and Quercus spp. (Oaks) in South Korea, considering topographic and climatic factors.Methods We used a dataset of diameter at breast height and radial growth estimates of individual trees, topographic and climatic factors in systematic sample plots distributed over the whole of South Korea. On the basis that radial growth is attributed primarily to tree age, we developed a radial growth model employing tree age as an explanatory variable. We estimated standard growth (SG), defined as radial growth of the tree at age 30, to eliminate the influence of tree age on radial growth. In addition, SG estimates including the Topographic Wetness Index, temperature and precipitation were calculated by the Generalized Additive Model.Important findings As a result of variogram analysis of SG, we found spatial autocorrelation between SG, topographic and climatic factors. Incremental temperature had negative impacts on radial growth of P. densiflora and positive impacts on that of Quercus spp. Precipitation was associated with positive effects on both tree species. Based on the model, we found that radial growth of P. densiflora would be more vulnerable than that of Quercus spp. to climatic factors. Through simulation with the radial growth model, it was predicted that P. densiflora stands would be gradually replaced with Quercus spp. stands in eastern coastal and southern regions of South Korea in the future. The models developed in this study will be helpful for understanding the impact of climatic factors on tree growth and for predicting changes in distribution of P. densiflora and Quercus spp. due to climate change in South Korea.     

Diversity and specificity of ectomycorrhizal fungi retrieved from an old-growth Mediterranean forest dominated by Quercus ilex

We analysed the ectomycorrhizal (ECM) fungal diversity in a Mediterranean old-growth Quercus ilex forest stand from Corsica (France), where Arbutus unedo was the only other ECM host. On a 6400 m2 stand, we investigated whether oak age and host species shaped below-ground ECM diversity. Ectomycorrhizas were collected under Q. ilex individuals of various ages (1 yr seedlings; 3-10 yr saplings; old trees) and A. unedo. They were typed by ITS-RFLP analysis and identified by match to RFLP patterns of fruitbodies, or by sequencing. A diversity of 140 taxa was found among 558 ectomycorrhizas, with many rare taxa. Cenococcum geophilum dominated (35% of ECMs), as well as Russulaceae, Cortinariaceae and Thelephoraceae. Fungal species richness was comparable above and below ground, but the two levels exhibited < 20% overlap in fungal species composition. Quercus ilex age did not strongly shape ECM diversity. The two ECM hosts, A. unedo and Q. ilex, tended to share few ECM species (< 15% of the ECM diversity). Implications for oak forest dynamics are discussed.     

Epiphytic lichens and bryophytes of forest ecosystems in Tuscany (Central Italy)

The distribution of epiphytic bryophytes and lichens on three species of trees (Quercus ilex, Quercus cerris and Fagus sylvatica, representing the forest ecosystems of central Italy at low, medium and high elevations, respectively) was examined with respect to host specificity and habitat differences. Although most species were host-specific, the results suggested that habitat characteristics are more important than phorophyte properties in determining the distribution of lichens and bryophytes. Factors affecting the diversity of the epiphytic flora were different for lichens and bryophytes. In the case of lichens, climatic parameters and forest monospecificity were important factors, whereas tree cover (i.e. shade) and to a certain extent the number of available trees seemed more important for bryophytes.     

Stand and self-thinning dynamics in natural <Emphasis Type="Italic">Abies</Emphasis> stands in northern Hokkaido,Japan

Stand dynamics and self-thinning were analyzed in relation to the dynamics of above-ground biomass in natural Abies sachalinensis stands growing on sand dunes in northern Hokkaido, Japan. This was done in order to examine wave-type regeneration in the stands. Fifty-two plots were established in almost pure Abies stands that ranged from saplings to the mature and collapsing growth stages. Above-ground biomass and tree height reached asymptotic levels prior to the collapsing phase, unlike wave-regeneration Abies stands in central Japan and North America. Stand density was high in the young growth stages, but the self-thinning rate, that is, the density decrease per biomass growth in the study stands was greater than in wave-regeneration stands in central Japan, as indicated by a large self-thinning exponent (–1.26 by reduced major axis regression). The range of tree height distribution was very narrow, and the stands vertical structure was typically single-layered. The slenderness ratio of trees was large, except in young stands. In mature and collapsing stands, advanced seedling density increased markedly. These stand and tree characteristics were considered to be correlated with the wave-type regeneration in the study stands, and it is assumed that prevailing winds affect tree mortality.