Genetically-and environmentally-dependent processes drive interspecific and intraspecific divergence in the Chinese relict endemic genus Dipteronia

China is a hotspot of relict plant species that were once widespread throughout the Northern Hemisphere.Recent research has demonstrated that the occurrence of long-term stable refugia in the mountainous regions of central and south-western China allowed their persistence through the late Neogene climate fluctuations.One of these relict lineages is Dipteronia,an oligotypic tree genus with a fossil record extending to the Paleocene.Here,we investigated the genetic variability,demographic dynamics and diversification patterns of the two currently recognized Dipteronia species(Dipteronia sinensis and D.dyeriana).Molecular data were obtained from 45 populations of Dipteronia by genotyping three cpDNA regions,two single copy nuclear genes and 15 simple sequence repeat loci.The genetic study was combined with niche comparison analyses on the environmental space,ecological niche modeling,and landscape connectivity analysis.We found that the two Dipteronia species have highly diverged both in genetic and ecological terms.Despite the incipient speciation processes that can be observed in D.sinensis,the occurrence of long-term stable refugia and,particularly,a dispersal corridor along Daba Shan-west Qinling,likely ensured its genetic and ecological integrity to date.Our study will not only help us to understand how populations of Dipteronia species responded to the tectonic and climatic changes of the Cenozoic,but also provide insight into how Arcto-Tertiary relict plants in East Asia survived,evolved,and diversified.

Evaluation and analysis of intraspecific competition in maize: A case study on plant density experiment

Intraspecific competition is a common phenomenon in agricultural production,and maize is one of the most sensitive grass species to intraspecific competition due to its low tillering ability.This study evaluated and analyzed intraspecific competition in maize,and screened competitive indices that could be used to evaluate intraspecific competition in a maize population.A 2-year field experiment was conducted using the maize hybrid Zhongdan 2 at 12 plant densities ranging from 1.5 to 18.0 plants（pl）m-2.The results showed that the response of single-plant grain yield and dry matter at harvest to increased plant density decreased exponentially and that the harvest index decreased linearly.The response of population-level grain yield to plant density was curvilinear,producing a maximum value at the optimum population density.However,the yielddensity equation agreed well with the Steinhart-Hart equation curves,but not with the quadratic equation curves reported by most previous studies.Competitive indices are used to evaluate competition in a plant population or plant species.The present results show that competitive intensity（CI）and absolute severity of competition（ASC）increased with increasing plant density;however,relative yield（RY）and relative reproductive efficiency（RReff）decreased.The different responses of these indices reflect different aspects of competition.According to the analysis of CI,ASC,RY,and RReff,higher CI and ASC values indicate higher intraspecific competition,whereas higher RY and RReff values indirectly reflect lower intraspecific competition.These competitive indices evaluate not only the intraspecific competitive intensity under different plant densities of the same cultivar but also those of different cultivars under the same plant density.However,some overlap exists in the calculations of ASC,CI,and RY,so one could simply select any one of these indices to evaluate intraspecific competition in a maize population.In conclusion,the present study provides a method to evaluate intraspecific competition in maize populations,which may be beneficial for breeding high-yield maize varieties in the future.

INTRASPECIFIC CROSSINGS OF UNDARIA PINNATIFIDA (HARV.) SUR.-A POSSIBLE TIME-SAVING WAY OF STRAIN SELECTION

Unicellular gametophytes of Undaria pinnatifida (Harv.) Sur. were isolated in Qingdao, P. R. China in April 1993 and in Tokushima, southem Japan in March 1995. Diferent intraspecific crossings by using unicellular male and female gametophytes were successfully undertaken in Sept. of 1995 in Qingdao.Sporophytes were transplanted to two different locations for open an cultivation. One was at Zhanqiao (ZQ) Bay where the water current was slower than that of another location - Taipingjiao(TPJ). A total of218 adult sporophytes were harvested on January 12 and 18 from TPJ and ZQ repectively. For eacncombination, 10 sporophytes un cultivated. Analysis of the mprpholgical characteristics of adult spprophytes indicated that the longest length between two bases of the serration of pinnate bldes (W2) is a morphological characteristic that can be transferred from the parent plant to the next generations regardless of environmental variations. There was evidence that W2 was apparently determined by sex-linked factors, i.e, by male parental grametophyte.. Sporophytes from certain crossing combinations showed more vigorous growth than those from other crossing combinations. It is therrfore possible to select grametophyte strains which can be used as parental gametophytes for the seeding production of sporophytes with more vigorous growth within shorter cultivation period. The morphology of hybrids from a Qingdao strain and a Tokushima cultivated strain resembled that of both parental plants in frond feaures (wrinkled or smooth) and W2. Sporophyll formation also varied with strains. The fact that adult sporophytes resulting from the same crossing combinations have identical morphological characteristics under the same environmental conditions indicates the possibility of a new way to select strains which are expected to be ideal for commerical production by purposely selecting, propagating, and seeding unicellular gametophytes for sporeing production through freeliving techniques of gametophytes.

Potential tradeoffs between intraspecific and interspecific trait variations along an environmental gradient in a subtropical forest

There has been growing attention to intraspecific variation in trait-based plant ecology. However, studies on these changes across ontogenetic stages and the potential trade-offs with interspecific traits along environmental gradients are rare. In this study, we measured six wood and bark traits of 1030 trees of six species(Castanopsis nigrescens; C. carlesii; Lithocarpus polystachyus; L. synbalanos; Ormosia glaberrima; O. pachycarpa) from a10-ha plot in a subtropical forest. Mean intraspecific variation in bark thickness and bark percentage to DBH was more than twice that for wood density and bark density.Bark thickness and bark percentage showed a consistent trend with increasing tree size. Small-tree traits were more variable than the same traits in larger trees. Altitude,convexity and soil nutrients explained the majority of the variations in the six traits, while sibling species had similar relationships between traits and environmental variables.Trees with dense wood and thin bark were usually found on steep slopes at lower altitudes. Our findings show intraspecific trait variability has different spatial patterns compared with interspecific variabilities along an environmental gradient.

Comparative genome analysis on intraspecific evolution and nitrogen fixation of Leptospirillum ferriphilum isolates

To reveal the intraspecific evolution of Leptospirillum ferriphilum isolates which thrived in industrial bioleaching ecosystems and acid mine drainages,genome sequences of L.ferriphilum YSK,L.ferriphilum DX and L.ferriphilum ZJ were determined to compare with complete genome of L.ferriphilum ML-04.The genome comparisons reveal that extensive intraspecific variation occurs in their genomes,and that the loss and insertion of novel gene blocks of probable phage origin may mostly contribute to heterogeneity of gene content among L.ferriphilum genomes.Surprisingly,a nif gene cluster is identified in L.ferriphilum YSK and L.ferriphilum ZJ genomes.Intensive analysis and further experiments indicate that the nif gene cluster in L.ferriphilum YSK inherits from ancestor rather than lateral gene transfer.Overall,results suggest that the population of L.ferriphilum undergoes frequent genetic recombination,resulting in many closely related genome types in recent evolution.The combinatorial processes profoundly shape their physiologies and provide the basis for adaptation to different niches.

Strong intraspecific trait variation in a tropical dominant tree species along an elevational gradient

Functional trait variation of plant species includes both inter-and intraspecific variation;however,traitbased plant ecology generally considers only interspecific variation while ignoring intraspecific variation.One reason for this neglect is that intraspecific variation may be negligible when compared to interspecific variation;however,direct comparisons between inter-and intraspecific variation of plant species are lacking,especially in tropical forests.Here we investigated intraspecific leaf trait variation(leaf area,specific leaf area,leaf thickness,leaf density,leaf chlorophyll content) of Pittosporopsis kerrii Craib(Icacinaceae),the most abundant tree species in the Xishuangbanna tropical seasonal rainforest in southwestern China,along an elevational gradient(703-824 m).We found a substantial range of intraspecific variation in P.kerrii that was never less than 22.1% of range of the interspecific variation among 462 tree species reported before in the same community.Moreover,with increased elevation,both leaf thickness and density increased and specific leaf area decreased significantly.It could be more important for the individuals of P.kerrii to produce thicker and denser leaves to tolerate environmental stress(e.g,soil water availability) rather than having high growth rates at the places with higher elevation in the Xishuangbanna tropical seasonal rainforest.

Identifying Intraspecific Variation in Venom Yield of Chinese Cobra(Naja atra) from Ten Populations in China's Mainland

Detailed information on venom yield is helpful in preparing antivenoms and treating snakebites, but such information is lacking for many species of venomous snakes. The Chinese cobra(Naja atra) is a large sized, venomous snake commonly found in southeastern China, where it causes a heavy burden of snakebites. To examine the effects of various factors(morphology, sex, age, season, and geographical origin) on the venom yield in this snake, we collected venom samples of 446 individuals(426 adults and 20 neonates) from 10 populations of N. atra over an eightyear period. We used two variables, lyophilized venom mass(venom yield) and solid content of venom(% solids), to quantify the venom yield. We used linear regression analysis to check if venom yield was related to morphological factors, one-way ANOVA and one-way ANCOVA to detect the sexual, ontogenetic, and geographic variation in venom yield, and repeated-measures ANOVA to examine seasonal shifts in venom yield. Our results indicate that venom yield of N. atra is positively related to the morphological traits examined, with male snakes expelling more venom than females. Venom yield in N. atra was age-related, with elder snakes always expelling more venom than younger ones. Geographic variation in venom yield was also observed, while seasonal variation was not. The solid content of venom was lower in males than in females, but this was not related to morphology, season, age, or geography. Our findings suggest that venom yield in N. atra is influenced by multiple factors, as well as by the interactions among these factors.

Intraspecific diversity: adaptive differentiation of Picea mongolica W. D. Xu ecotypes

Picea mongolica W. D. Xu is an endemic and endangered species which is only found in semi-arid areas of northern China. It has been widely used as an afforestation tree in the establishment of the Three-North Shelterbelt System for its adaptation to arid soils and as a virescent tree in urban gardens and streets for its beautiful shape. Due to different microenvironments, P. mongolica populations may differentiate into many ecological groups with different adaptive abilities. Long-term adaptation to a dry environment makes P. mongolica differentiate into different ecotypes. Typical ecotypes are P. mongolica f. purpurea （Fp）, P. mongolica f. rubra （Fr） and P. mongolica f. viridis （Fv）. Our results show that cone size is clearly not uniform among different ecotypes; the largest cones are found in the Fv ecotype and the smallest in Fp. There were also distinct differences between these ecotypes in terms of height and length of squama. At the molecular level, the zymograms of peroxidase and lipase prove the existence of different ecotypes in P. mongolica. The results are useful for investigating and managing this rare spruce species in China.

Intraspecific leaf morphological variation in Quercus dentata Thunb.:a comparison of traditional and geometric morphometric methods,a pilot study

To compare the application of traditional morphometric methods(TMMs)and geometric morphometric methods(GMMs)in the study of intraspecific leaf morphological characters of Quercus dentata,fifteen linear measurement indices and thirteen landmarks of leaves were used to study leaf morphology of three provenances(H1,H2,and H3).In TMMs,principal component analysis(PCA)showed that leaf size–related indices played an important role in population classification.Partial least square(PLS)analysis showed that the main morphological characters affecting leaf size were the average depth of the lobes and the length–width ratios.However,the important indices to distinguish the provenances were circularity,leaf width,and length–width ratio.The results of discriminant analysis(DA)showed that 74.0%of H1,68.0%of H2,and 74.0%of H3 were correctly classified.Cluster analysis showed that the Mahalanobis distances between H1 and H2,H1 and H3,and H2 and H3 were 4.3761,11.4629,and 10.2067,respectively.In GMMs,PCA based on symmetrical components showed that the difference in leaf morphology was mainly due to the changing trend of the leaf apex and base,petiole length,and degree of leaf cracking.PLS analysis showed that there was a significant covariation between the leaf symmetrical components and size:as the leaf enlarged,the widest part gradually moved up,and the shape changed from nearly oval to lanceolate.DA results showed that 86.0%and 78.0%of H1 and H2,70.0%and 80.0%of H1 and H3,and 82.0%and 76.0%of H2 and H3 were correctly classified.Canonical variate analysis results showed that the Mahalanobis distances between H1 and H2,H1 and H3,and H2 and H3 were 1.7238,1.5380,and 1.6329,respectively.Both GMMs and TMMs showed significant differences in morphology among the three Q.dentata provenances,but GMMs had higher classification accuracy and could provide more information about leaf shape,whereas TMMs could provide more information about leaf size.Based on our results,GMMs are promising in the study of leaf morphological variation within Q.dentata provenances.

On the intraspecific variability in basal metabolism and the food habits hypothesis in birds

The food habits hypothesis （FHH） stands as one of the most striking and often-cited interspecific patterns to emerge from comparative studies of endothermic energetics. The FHH identifies three components of diet that potentially produce variability in mass-independent BMR, i.e. food quality, food availability, and food predictability or environmental productivity. The hypothesis predicts that species with diets of low energy content and/or low digestibility should evolve low mass-independent BMRs. The effects of food habits on BMR have been widely investigated at the interspecific level, but the variation between individuals and populations has been largely ignored. Our focus is to compare predictions derived from interspecific studies with data collected from within-species studies to explore the mechanisms and functional significance of adaptive responses predicted by the food-habits hypothesis among birds. We conclude that if BMR is correlated with daily energy expenditure, then organisms that can lower BMR will reduce daily energy expenditure and hence, food requirements. Birds that lower BMR in stressful environments may increase survival. Nevertheless, the mechanism （s） by which birds eating a low quality diet reduce BMR and whether lower BMR affects fitness remain to be determined [Current Zoology 56 （6）： 759-766, 2010].

Intraspecific morphometric diversity of barramundi (Lates calcariferBloch, 1790) in the waters of southern Bangladesh

A total of 160 barramundi's (Lates calcarifer Bloch, 1790) sampled from four rivers (Tentulia, Balaswar, Bakkhali, and Andarmanik) along the southern coastal region of Bangladesh were investigated in terms of morphometric characters to reveal the intraspecific variation. Twenty-five morphometric measurements were extracted using the conventional method and subjected to multivariate analyses (i.e., principal component analysis (PCA), discriminate function analysis (DFA), cluster analysis (CA)) to distinguish individuals from different rivers. The result demonstrated that twenty-two out of 25 measurements was statistically significant (Univariate ANOVA) among all four populations. PCA analysis of morphometric characters resulted in two principal components, PC I and PCⅡ which accounted for 79.25% and 4.28% of the total data variance. PC I-PC Ⅱ plot explained 83.53% of total variance differentiated the population of L. calcarifer into two groups. Discriminate analysis correctly classified about 88.1% of the examined fish into the four areas. The UPGMA dendrogram showed that Bakkhali populations were the most morphologically different populations in comparison to other populations, while Andarmanik and Balaswar populations were very close to each other. The strong morphometric variation between Bakkhali and Tentulia, Andarmanik and Balaswar was observed in the present study, suggested the evidence of the separate stock population of barramundi in these locations, which might require distinct stock management strategies for resource sustainability in the waters of southern Bangladesh. However, if these findings are supported by further molecular markers and geometric morphometry, this would be a strong indication of different stocks of this population in the four rivers of southern Bangladesh.

Cell number explains the intraspecific spur-length variation in an Aquilegia species

Variations of nectar spur length allow pollinators to utilize resources in novel ways,leading to the different selective pressures on spurs and allowing taxa to diversify.However,the mechanisms underlying spur length variation remain unclear.Interspecific comparisons of spur length suggest that both cell division and anisotropic expansion could explain the changes of spur length,and that hormone-related genes contribute to the process of spur formation.In contrast,little is known about intraspecific spur length variation.In Aquilegia rockii,spur length varies strikingly,ranging from 1 mm to 18 mm.To examine the potential mechanisms underlying spur length variation in A.rockii,we observed cell morphology and analyzed RNA-seq of short-and long-spurred flowers.Scanning electron microscopy revealed that at two positions on spurs there were no differences in either cell density or cell anisotropy between short-and long-spurred flowers,suggesting that in A.rockii changes in cell number may explain variations in spur length.In addition,we screened transcriptomes of short-and long-spurred flowers for differentially expressed genes;this screen identified several genes linked to cell division(e.g.,F-box,CDKB2-2,and LST8),a finding which is consistent with our analysis of the cellular morphology of spurs.However,we did not find any highly expressed genes involved in the hormone pathway in long-spurred flowers.In contrast to previous hypotheses that anisotropic cell expansion leads to interspecific spur variation in Aquilegia,our results suggest that cell number changes and related genes are mainly responsible for spur length variations of A.rockii.Furthermore,the underlying mechanisms of similar floral traits in morphology may be quite different,enriching our understanding of the mechanisms of flower diversity in angiosperms.

Intraspecific Variability and Problematic Dimorphism in the Early Cretaceous(Valanginian)Ammonite Saynoceras verrucosum(d'Orbigny,1841)

The population of Saynoceras verrucosum （d＇Orbigny） from the Polish basin （Wawal section, central Poland） shows no significant intraspecific variability. Dimorphism has not been found in this population. Statistical analyses of material from the shallow epicratonic Polish basin and the relatively deep Vocontian basin （southeastern France） indicated two populations. Forms from the Polish basin are somewhat more inflated and smaller than forms from the Vocontian basin.

Role of intraspecific competition in intrinsic growth rate regulation in an Oikopleura dioica(Tunicata)population

Planktonic Oikopleura dioica respond almost instantly to opportunistic algal blooms,but it is unknown whether the population increase can change from an exponential to a logistic model to avoid a final sudden collapse.To test the hypothesis that intraspecific competition regulates the intrinsic rate of natural increase(r),density-dependent effects on growth and reproduction performance were investigated in O.dioica via laboratory incubations.Over a large food concentration range,batch maturation was observed above the per capita food supply(PFS)of 8.1µgC/ind.in 4.5 d.Somatic growth was saturated beyond this PFS value,whereas gonad length increased continuously.Below this threshold,individuals reached small body and gonad lengths,and maturation was rarely observed during the incubation period.The gonad/body volume and maturation ratios also increased with the PFS.Instead of the food concentration,the r values were regulated by competition pressure via variability in maturation duration and the proportion of mature individuals in the cohorts.When the minimum food demand was satisfied in the designated generation time,the r value tended to be regulated by the spawning proportion in the population.Otherwise,prolonged development duration and decreased r values were expected.

Intraspecific trait variation of woody species reduced in a savanna community,southwest China

Plants deploy various ecological strategies in response to environmental heterogeneity.In many forest ecosystems,plants have been reported to have notable inter-and intra-specific trait variation,as well as clear phylogenetic signals,indicating that these species possess a degree of phenotypic plasticity to cope with habitat variation in the community.Savanna communities,however,grow in an open canopy structure and exhibit little species diversification,likely as a result of strong environmental stress.In this study,we hypothesized that the phylogenetic signals of savanna species would be weak,the intraspecific trait variation(ITV)would be low,and the contribution of intraspecific variation to total trait variance would be reduced,owing to low species richness,multiple stresses and relatively homogenous community structure.To test these hypotheses,we sampled dominant woody species in a dry-hot savanna in southwestern China,focusing on leaf traits related to adaptability of plants to harsh conditions(yearround intense radiation,low soil fertility and seasonal droughts).We found weak phylogenetic signals in leaf traits and low ITV(at both individual and canopy-layer levels).Intraspecific variation(including leaf-,layer-and individual-scales)contributed little to the total trait variance,whereas interspecific variation and variation in leaf phenology explained substantial variance.Our study suggests that intraspecific trait variation is reduced in savanna community.Furthermore,our findings indicate that classifying species by leaf phenology may help better understand how species coexist under similar habitats with strong stresses.

Evolutionary history of a desert perennial Arnebia szechenyi(Boraginaceae):Intraspecific divergence,regional expansion and asymmetric gene flow

The complex interactions of historical,geological and climatic events on plant evolution have been an important research focus for many years.However,the role of desert formation and expansion in shaping the genetic structures and demographic histories of plants occurring in arid areas has not been well explored.In the present study,we investigated the phylogeography of Arnebia szechenyi,a desert herb showing a near-circular distribution surrounding the Tengger Desert in Northwest China.We measured genetic diversity of populations using three maternally inherited chloroplast DNA(cpDNA)fragments and seven bi-paternally inherited nuclear DNA(nDNA)loci that were sequenced from individuals collected from 16 natural populations across its range and modelled current and historical potential habitats of the species.Our data indicated a considerably high level of genetic variation within A.szechenyi and noteworthy asymmetry in historical migration from the east to the west.Moreover,two nuclear genetic groups of populations were revealed,corresponding to the two geographic regions separated by the Tengger Desert.However,analysis of cpDNA data did not show significant geographic structure.The most plausible explanation for the discrepancy between our findings based on cpDNA and nDNA data is that A.szechenyi populations experienced long periods of geographic isolation followed by range expansion,which would have promoted generalized recombination of the nuclear genome.Our findings further highlight the important role that the Tengger Desert,together with the Helan Mountains,has played in the evolution of desert plants and the preservation of biodiversity in arid Northwest China.

Intraspecific Phylogenetic Relationships of Caryopteris incana in the Tsushima Islands, Japan, Using DNA Sequence Analysis

Caryopteris incana is a perennial shrub distributed in the temperate zone of the East Asia. It is found in West Kyushu in Japan, where it is designated as an endangered species. Tsushima, Nagasaki, which experienced repeated connection and fragmentation between the Korean Peninsula and Japan, is an island on the route along which C. incana moved to Japan from continental Asia. We conducted field work and confirmed the genetic structure of populations using DNA sequence analysis to construct a detailed distribution map and clarify the intraspecific phylogenetic relationships of C. incana in Tsushima Island. We confirmed 72 populations in Tsushima. Using the leaves of individuals cultivated from seeds collected from each natural population, we analyzed the chloroplast and nuclear DNA sequence variations. Among the populations, sequence variations were confirmed in six regions of chloroplast DNA, and six haplotypes, including base substitutions, were distinguished. Two haplotypes were mainly divided at the border of the northern part of the southern island in Tsushima. One population in the northwestern part of the north island showed a haplotype derived from the southern part. This finding revealed that the distribution of C. incana had been artificially influenced. Several haplotypes were confirmed by sequence variations in the northern populations, but only one haplotype in the southern populations, suggesting that C. incana on the north island had separated early from the south island in Tsushima.

Intraspecific Relationship of Sheep Based on Mitochondrial Cytochrome b Gene

[ Objective] To preliminarily explore the intraspecific relationship of sheep based on cytochrome b （ Cyt b） gene of mitochondrial DNA （mtDNA）. [Method] The Cyt b gene sequences in 112 sheep individuals of two local sheep breeds were amplified by PCR. Then the amplified products were digested with EcoR I and analyzed by restriction fragment length polymorphism （RFLP）. [ Result] As many as 56 samples from Tan sheep and 56 samples from Wadi sheep were detected. The results showed that the amplified Cyt b gene in 51 individuals of Tan sheep had one EcoR I restriction site and no EcoR I restriction site in other five individuals, thus the Cyt b gene in Tan sheep showed two restriction morphs; the Cyt b gene in all 56 individuals of Wadi sheep had one EcoR I restriction site and showed one restriction morph. [ Conclusion] The polymorphism of mitochondrial Cyt b gene in Tan sheep and Wadi sheep is poor, and the Cyt b gene in sheep breeds is very conservative. Therefore, using Cyt b gene as gene marker to study the intraspecific relationship of sheep has some limitations.

Competitive interactions between invasive fall armyworm and Asian corn borer at intraspecific and interspecific level on the same feeding guild

Interspecific competition is an important factor in the population dynamics and geographical distribution of insect populations.Fall armyworm(FAW),Spodoptera frugiperda,an invasive species,and the Asian corn borer(ACB),Ostrinia furnacalis,a native pest species are major pests on maize in China,posing a threat to maize yield and grain quality.A series of laboratory,greenhouse,and field experiments were conducted to elucidate the competitive interactions between FAW and ACB.In the laboratory experiments,FAW exhibited aggressive behaviors more frequently when compared with ACB,while the latter species exhibited defense behaviors more frequently.Higher intraspecific competition was recorded in the FAW conspecific rather than in ACB,particularly in the 6th larval stage.FAW had a higher interspecific competitive advantage through intraguild predation over ACB,resulting in partial or complete displacement ACB when initially the ratio of the 2 species was 1:1.The interspecific competition also had significantly influenced on the population parameters,defensive enzymes,and nutrient of these 2 species.Competitive interaction proved that the response of superoxide,catalase,and soluble protein in FAW were significantly increased,whereas the total sugar content in both species was substantially decreased.Survival rate,and the plant damage that co-infested by both species varied significantly among the sequential combinations under greenhouse and field conditions.FAW consistently exhibited stronger intraspecific aggression than ACB under laboratory and field conditions when co-existing on the same feeding guild.These findings contribute to efforts toward the improvement of integrated pest management programs for FAW,in decision making for invasive and native pests’management strategies to reduce the high risks of FAW and ACB outbreaks.

Environment-driven intraspecific variation shows coordination of functional traits of deciduous oaks among and within different biological levels

Deciduous oaks(Quercus spp.)are distributed from subalpine to tropical regions in the northern hemi-sphere and have important roles as carbon sinks and in climate change mitigation.Determining variations in plant functional traits at multiple biological levels and linking them to environmental variables across geographical ranges is important for forecasting range-shifts of broadly-distrib-uted species under climate change.We sampled leaves of five deciduous Quercus spp.covering approximately 20°of latitude(~21°N-41°N)and 20 longitude(~99°E-119°E)across China and measured 12 plant functional traits at different biological levels.The traits varied distinctively,either within each biological level or among different levels driven by climatic and edaphic variables.Traits at the organ level were significantly correlated with those at the cellular and tissue levels,while traits at the whole-plant level only correlated with those at the tissue level.The Quercus species responded to changing environments by regulating stomatal size,leaf thickness and the palisade mesophyll thickness to leaf thickness ratios with contrasting degree of effect to adjust the whole-plant functioning,i.e.,intrinsic water use efficiency(iWUE),carbon supply and nitrogen availability.The results suggest that these deciduous Quercus spp.will maintain vigour by increasing iWUE when subjected to large temperature changes and insufficient moisture,and by accu-mulating leaf non-structural carbohydrates under drought conditions.The findings provide new insights into the inher-ent variation and trait coordination of widely distributed tree species in the context of climate change.