Within-population variability in architectural traits and suitability to forestry conditions in Nothofagus nervosa (= N. alpina; Nothofagaceae)

? Context

There is evidence that Nothofagus nervosa (= N. alpina) is suitable for timber production in temperate regions due to its wood quality and fast growth. However, high intra-specific variability in the architectural traits of this species limits its usefulness.

? Aims

This study was aimed at evaluating intra-specific variability in the architecture of N. nervosa trees at the population level, with emphasis on traits related to the suitability of trees for timber production.

? Methods

The size of 13-year-old trees installed in an experimental field population and the extent of differentiation between trunk and branches were compared among families (trees derived from different mother trees).

? Results

The large majority of trees exhibited architectural features indicative of high suitability for timber production: clear differentiation between trunk and main branches and few occurrences of permanent trunk forks, apex deaths, and sylleptic branching. Trees belonging to different families differed in height and trunk diameter. Micro-environmental conditions had a very significant effect on most architectural traits, including the trunk differentiation.

? Conclusions

N. nervosa emerges as a suitable option for timber production in temperate regions. Trunk growth seems to be under both environmental and genetic control in this species.     

Hydraulic properties and embolism in small-diameter roots of five temperate broad-leaved tree species with contrasting drought tolerance

? Context

It has been estimated that about half of a plant??s total hydraulic resistance is located belowground, but it is not well known how temperate tree species differ in root hydraulic properties and how these traits vary with the species?? drought tolerance.

? Aims

We examined root anatomical and hydraulic traits in five broad-leaved tree species with different drought tolerance, analyzed the relation between root anatomy and hydraulic conductivity and root embolism, and investigated the relation of these traits to the species?? drought tolerance.

? Methods

In small-diameter roots (2?C6?mm), we measured vessel diameters and vessel density, specific hydraulic conductivity, and the percental loss of conductivity (??native?? embolism) during summer in a mixed forest.

? Results

Specific conductivity was positively related to vessel diameter but not to vessel density. Drought-tolerant Fraxinus showed the smallest mean vessel diameters and drought-sensitive Fagus the largest. Specific conductivity was highly variable among different similar-sized roots of the same species with a few roots apparently functioning as ??high-conductivity roots??.

? Conclusion

The results show that coexisting tree species can differ largely in root hydraulic traits with more drought-sensitive trees apparently having larger mean vessel diameters in their roots than tolerant species. However, this difference was not related to the observed root conductivity losses due to embolism.     

Light transmissivity of tube shelters affects root growth and biomass allocation of Quercus ilex L. and Pinus halepensis Mill

Context

Tube shelters have been shown to enhance field performance of several Mediterranean species, but responses of newly planted seedlings to the microenvironment induced by shelter walls with different light transmissivity are still poorly documented.

Aims

We studied effects of a range of shelters with varying light transmissivity on post-planting seedling responses during the wet season establishment phase for two Mediterranean trees of contrasting functional ecology.

Methods

Root growth, biomass allocation, water potential, and chlorophyll fluorescence of Quercus ilex and Pinus halepensis seedlings were evaluated across shelters varying in light transmissivity (80, 40, 20, and 10 % plus a mesh shelter) with irrigation.

Results

Plants in dark tubes (20 and 10 % light transmissivity) had less above- and belowground growth and more than two times greater leaf to protruding roots mass ratio, with shoot growth response of Q. ilex being less plastic. Ratio of leaf area/protruding roots area decreased when light transmissivity increased, although no differences were found at ≥40 % transmissivity. Xylem water potential indicated lack of water stress, and high maximum photosynthetic efficiency (F _v/F _m) values show no photoinhibition symptoms irrespective of light transmissivity.

Conclusion

Shelter transmissivity ≥40 % promotes rapid and vigorous root growth immediately after planting for these species. This minimum transmissivity should be considered as a target when designing shelters to help root development and improve water balance of Mediterranean seedlings.     

Back to the roots: how do seedlings of native tree species react to the competition by exotic species?

Context

Identifying the traits of exotic species may explain their invasiveness and help control the negative impacts of these species on native ecosystems. In this study, we investigated root competition as an important driver for the competitiveness of two exotic tree species in the seedling stage.

Aims

In a pot experiment, carried out in Central Europe, we studied the effect of root competition of two exotic tree species (Prunus serotina Ehrh. and Robinia pseudoacacia L.) on biomass allocation of two native ones (Quercus robur L. and Carpinus betulus L.).

Methods

Seedlings of these species were exposed to intra- and interspecific mixtures with and without the effect of root competition. For this, belowground plastic partitions were installed in the pots.

Results

Competition intensity in the pots increased over time, irrespective of mixture type. However, this increase was much higher for the native species if mixed with the exotic species compared to monocultures or inter-native mixtures. In addition to ontogeny, competition affected biomass allocation patterns. Under root competition by the exotic species, the native species preferentially allocated their biomass to the roots. A higher allocation to the roots was mainly achieved at the expense of leaf and branch biomass.

Conclusion

Root competition of P. serotina and R. pseudoacacia may be a reason for the lack of Q. robur and C. betulus in the seedling stage in natural environments where all four species occur.     

Genotypic parameters of wood density and fiber traits in triploid hybrid clones of Populus tomentosa at five clonal trials

? Context

A clear understanding of the genetic control of wood properties is a prerequisite for breeding for higher wood quality in Populus tomentosa Carr. hybrid clones.

? Aims

The experiments aimed at unraveling genetic and environmental effects on wood properties among triploid hybrid clones of P. tomentosa.

? Methods

We used 5-year-old clonal trials established in Northern China to assess the heritability of wood density and fiber traits. Two hundred seventy trees from nine clones were sampled in five sites.

? Results

Site had a very significant effect on all recorded traits. Despite this large site effect, a tight genetic control was detected and clonal repeatability varied between 0.53 and 0.95. Significant genotype?×?environment interactions were detected for most of the traits. Moderate to tight correlation were evidenced among traits but they were not consistent with that in several cases that were site-dependent.

? Conclusions

Our results revealed a tight genetic control over several wood properties and therefore breeding programs might be able to improve wood density, fiber length, and coarseness in these hybrids.     

Low light acclimation in five temperate broad-leaved tree species of different successional status: the significance of a shade canopy

?Context

Tree species differ largely in their capability to produce characteristic shade leaves with effective morphological and physiological acclimation to low light.

?Aims

By examining the sun/shade leaf differentiation in leaf morphology, foliar nitrogen and photosynthetic capacity in five temperate tree species of different successional status, we aimed at identifying those leaf traits that determine the development of a typical shade crown with low light-acclimated leaves.

?Methods

Leaf morphology, foliar N content, photosynthetic capacity (V _cmax, J _max and A _max) and leaf dark respiration (R _d) were measured in the canopies of 26 adult trees of Fraxinus, Acer, Carpinus, Tilia and Fagus species.

?Results

Six traits (the sun/shade leaf differentiation in specific leaf area, leaf size, A _max per leaf area or per mass, photosynthetic N use efficiency and R _d) were found to characterise best the degree of low light acclimation in shade leaves. All five species exhibited certain modifications in leaf morphology and/or physiology in response to low light; Fagus sylvatica showed the highest and Fraxinus excelsior the lowest shade leaf acclimation.

?Conclusions

Our results indicate that the five early/mid- to late-successional species have developed species-specific low light acclimation strategies in their shade crowns which differ in terms of the relative importance of leaf morphological and physiological acclimation.     

Flammability of some companion species in cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis> L.) forests

Key message

The high flammability of some companion species in Quercus suber forests, estimated in laboratory tests, could potentially generate an increase in fire vulnerability and in fire risk.

Context

Recurrent wildfire is one of the main causes of forest degradation, especially in the Mediterranean region. Increased fire frequency and severity due to global change could reduce the natural resilience of cork oak to wildfire in the future. Hence, it is important to evaluate the flammability of companion species in cork oak forests in the particularly dry bioclimatic conditions of North Africa.

Aims

This study aimed to assess and compare flammability parameters at laboratory scale among ten companion frequent species in cork oak forests.

Methods

Fuel samples were collected in a cork oak (Quercus suber L) forest in the southern part of the mountains of Tlemcen (Western Algeria). A series of flammability tests were carried out using a Mass Loss Calorimeter device (FTT ®). A cluster analysis to classify flammability of the selected species was conducted using the K-means algorithm.

Results

The results revealed differences in the four flammability parameters (ignitability, sustainability, combustibility and consumability), in both fresh and dried fine fuel samples from Quercus suber, Pinus halepensis, Quercus ilex, Quercus faginea, Erica arborea, Arbutus unedo, Pistacia lentiscus, Calicotome spinosa, Juniperus oxycedrus and Tetraclinis articulata. Application of the K-means clustering algorithm showed that C. spinosa, T. articulata, J. oxycedrus and P. halepensis are highly flammable because of their high combustibility and sustainability.

Conclusion

The findings identify species that could potentially increase the vulnerability of cork oak forests to forest fires.

     

Performance of <Emphasis Type="Italic">Quercus suber</Emphasis> L. at nursery stage—application of two bio-inoculants under two distinct environments

Key message

Despite the fact that the technique of application of bioinoculants improved the quality of Quercus suber L. seedlings produced in nurseries, these benefits are dependent on the ecological conditions of the site and the composition of the applied inoculum, which interferes with the profile of the local fungal community.

Context

Quercus suber L. plays a key ecological and socio-economical role in the Iberian Peninsula. Symbiotic ectomycorrhizal fungi-ECM are crucial partners of several tree species, and assessing the efficacy of bioinoculants at nursery stage helps devising tools to increase plant resilience.

Aims

The aim of this study was to compare the effects of two inocula formulations of mixed ECM fungi and bacteria on the quality of seedlings produced in two forest nurseries, differing in environmental conditions and forest embedment.

Methods

Quercus suber L. seedlings were inoculated with a commercial product containing Pisolithus tinctorius (Pers) Coker & Couch, Scleroderma sp., and six bacterial species and with a non-commercial fungal and bacterial dual inoculum (Suillus granulatus (L.) Roussel + Mesorhizobium sp.). Biometric and nutritional parameters and morphological quality indexes were determined on seedlings. The ECM community was assessed by denaturing gradient gel electrophoresis and cloning-sequencing.

Results

In both nurseries, the seedling quality index in inoculated was up to 2-fold higher than in non-inoculated seedlings. Plant biomass differed significantly among nurseries. The inoculum influenced the profile of the fungal community. S. granulatus and P. tinctorius persisted for 6 months in the inoculated seedlings.

Conclusion

The nursery ecosystem influenced plant growth. Inoculation treatments increased plant performance; however, the dual inoculum resulted in more consistent improvements of Q. suber at nursery stage, highlighting the importance of inocula selection.

     

Mast seeding in deciduous forests of the northern Apennines (Italy) and its influence on wild boar population dynamics

Context

Pulsed food resources may strongly affect the population dynamics of several consumer species, with consequences on the ecosystem. One of the most common pulsed resources is forest mast seeding.

Aims

We analysed mast seeding in deciduous forests in a mountainous area of northern Apennines and its effect on population dynamics of wild boar (Sus scrofa L.).

Methods

We performed a quantitative, 20-year analysis on annual seed production in Turkey oak (Quercus cerris L.), beech (Fagus sylvatica L.) and chestnut (Castanea sativa Mill.) forest stands using litter traps. The wild boar population density was estimated by means of drive censuses and hunting bag records. The role of other biotic (density of predators) and abiotic (climate) factors potentially affecting wild boar mortality was also investigated.

Results

Turkey oak and chestnut showed high levels of seed production, whereas lower levels were found in beech. The pulsed resources of chestnut and Turkey oak positively affected piglet density. Analyses also highlighted the influence of snow cover and wolves on wild boar population dynamics.

Conclusion

Wild boar can be considered a pulse rate species, the management of which can be improved by annual monitoring of seed production.     

Estimation of chlorophyll in Quercus leaves using a portable chlorophyll meter: effects of species and leaf age

? The potential of a chlorophyll meter (CCM-200, Opti-Sciences, Tyngsboro, MA, USA) for the estimation of total leaf chlorophyll contents was evaluated in leaves of different developmental stages for Quercus pyrenaica Willd., Q. faginea Lam., Q. suber L. and Q. ilex L. subsp. ballota (Desf.) Samp.

? For all species and leaf stages, a strong non-linear relationship (P < 0.0001) was found between the chlorophyll content index (CCI) and leaf chlorophyll content per unit area. In all species, the slope of the CCI-chlorophyll content relationship varied during leaf development, suggesting that caution should be exercised when using the CCM-200, since the interpretation of CCI readings should be limited to samples of similar leaf age. Also, the data indicate that the models should also be speciesspecific, owing to differences in the intercept and/or slope of the equations.

? The addition of fresh LMA as a complementary parameter improved the accuracy of chlorophyll estimation in non-species-specific equations to a considerable extent when leaf samples of similar development were considered.

? After due consideration, it may be concluded that the CCM-200 chlorophyll meter is a reliable method for acquiring an estimation of chlorophyll contents in Mediterranean Quercus species, with potential use as an effective tool in forest management.

     

Interspecific differences in the oleoresin production of Copaifera L. (Fabaceae) in the Amazon rainforest

? Context

Copaifera species produce an oleoresin of commercial importance that is widely extracted in Amazon communities.

? Aims

This paper addresses two questions. (1) What are the morphological characteristics of Copaifera species that influence oleoresin production? (2) How do different Copaifera species respond to repeated harvests?

? Methods

We performed a large-scale experiment in the Brazilian Amazon. We tapped 110 Copaifera trees belonging to four species, and several morphological tree characteristics were measured to determine their effect on oleoresin production.

? Results

The proportion of Copaifera reticulata and Copaifera paupera trees that yielded more than 1 ml oleoresin was higher than the other species. The oleoresin volumes from yielding Copaifera pubiflora trees were significantly higher than those from C. reticulata and C. paupera, with Copaifera multijuga yielding intermediate values. Interestingly, none of the studied morphological tree characteristics had a significant effect on the proportion of yielding trees. Hollowed trees yielded significantly smaller volumes than non-hollowed trees. Both the proportion of yielding trees and oleoresin volumes decreased from the first to the second harvests for C. reticulata and C. paupera; however, the opposite pattern was observed for C. pubiflora.

? Conclusions

Oleoresin production capacity varies by species, and management protocols should account for these differences.     

Variation in carbon concentration and basic density along stems of sessile oak (Quercus petraea (Matt.) Liebl.) and Pyrenean oak (Quercus pyrenaica Willd.) in the Cantabrian Range (NW Spain)

Context

Information about variations in basic density (BD) and carbon content (%C) along tree stems is key to assessing forest carbon sinks.

Aims

The aim of the study was to determine any differences in %C and BD between different woody tissues (bark, sapwood and heartwood) in two widespread European oak species (Quercus pyrenaica and Quercus petraea).

Methods

Twenty trees were felled in northern Spain, and 317 discs cut from the trees were dried and analysed to determine %C and BD.

Results

There were significant differences in %C between bark, heartwood and sapwood, and between species. There were also significant differences in BD between the tissues (heartwood?>?sapwood?>?bark), and the BD was higher in Q. petraea than in Q. pyrenaica. Both %C and BD varied along the stem.

Conclusion

More accurate estimates of carbon contents were obtained by using specific values for different heights and anatomical parts, than by using single values.     

Assessing changes in species distribution from sequential large-scale forest inventories

? Context

It is assumed that global change is already affecting the composition, structure and distribution of forest ecosystems; however, detailed evidences of altitudinal and latitudinal shifts are still scarce.

? Aims

To develop a method based on National Forest Inventory (NFI) to assess spatio-temporal changes in species distributions.

? Methods

We develop an approach based on universal kriging to compare species distribution models from the different NFI cycles and regardless of the differences in the sampling schemes used. Furthermore, a confidence interval approach is used to assess significant changes in species distribution. The approach is applied to some of the southernmost populations of Pinus sylvestris and Fagus sylvatica in the Western Pyrenees over the last 40 years.

? Results

An increase of the presence of the two species in the region was observed. Scots pine distribution has shifted about 1.5 km northwards over recent decades, whereas the European beech has extended its distribution southwards by about 2 km. Furthermore, the optimum altitude for both species has risen by about 200 m. As a result, the zone in which the two species coexist has been enlarged.

? Conclusions

This approach provides a useful tool to compare NFI data from different sampling schemes, quantifying and testing significant shifts in tree species distribution over recent decades across geographical gradients.     

Effect of selective logging on stand structure and tree species diversity in a subtropical evergreen broad-leaved forest

?Context

Selective logging followed by natural regeneration is rarely employed for restocking subtropical evergreen broad-leaved forests in East Asia compared with the use of clear-cutting.

?Aims

To clarify the succession of these forests, the effects of selective logging on stand structure, species diversity, and community similarity were studied in a mature and regenerating forest in Okinawa, Japan.

?Methods

Four study plots were established, and trees ≥1.2 m height were identified by species name, tree height, and diameter at breast height.

?Results

The results showed that the species composition of regenerating forest was similar to mature forest; however, the former had a greater species density and Shannon–Wiener index than the latter. Castanopsis sieboldii and Distylium racemosum, the predominant trees in the mature forest, continued to dominate the regenerating forest, with a broad layer distribution. High Sørensen and Jaccard community similarity indices for mature and regenerating forest indicated that the regeneration occurred in a progressive succession.

?Conclusion

The similar species composition and stand structure for both mature and regenerating forest, and the higher species diversity for the latter, provided no evidence of forest degeneration and suggested that the regenerating forest may develop into a stand similar to preselective logging forest.     

Effects of region, soil, land use, and terrain type on fuelwood properties of five tree/shrub species in the Sahelian and Sudanian ecozones of Mali

? Context

There is little information about inter- and intra-specific variation in fuelwood properties of trees/shrubs in West Africa.

? Aims

We studied variation in fuelwood properties of Balanites aegyptiaca, Combretum glutinosum, Guiera senegalensis, Piliostigma reticulatum, and Zizyphus mauritiana in the Sahelian and Sudanian ecozones of Mali.

? Methods

Trees were sampled on different soil (sandy, sandy/loam, or rocky), land use (parkland agroforest or woodland), and terrain (flat, temporarily flooded, or hill slope) types in five regions extending from the drier eastern to the more humid western parts of Mali. Basic density, volatile matter, fixed carbon, ash content, moisture content, gross calorific value, and fuel value index were measured for each tree and adjusted for tree age.

? Results

All fuelwood properties differed significantly among species, but there were significant species by region interactions. Most fuelwood properties of four species differed significantly among regions. Soil and/or land use type had significant effects on a few fuelwood properties of four species.

? Conclusion

In general, fuelwood production is recommended in all regions for G. senegalensis, in drier regions for B. aegyptiaca and C. glutinosum, in more humid regions for Z. mauritiana, and on rocky soils for all species. Fuelwood production of P. reticulatum is not recommended.     

Characterization and photodegradation mechanism of three Algerian wood species

Aging is the irreversible change of mechanical, physical, and chemical properties of materials; the main objective of this work was to study the photochemical degradation and structural changes of three major Algerian wood species. For this, we evaluated the photodegradation mechanism for Maritime Pine (Pinus pinaster), zeen oak (Quercus canariensis), and afares oak (Quercus afares) by accelerated aging in a Xenon test chamber. Degradation of the samples was established by an initial color change (after 30 h exposure), followed by roughening and cracking (120 h exposure) as translated by scanning electron microscopy and Fourier transform infrared spectroscopy. The discoloration of irradiated wood samples was primarily related to the decomposition of lignin which is the key structure in wood photodegradation. As expected, a decrease in mechanical properties was observed; for all samples, the modulus of elasticity decreased after aging, indicating that the wood specimens loss some of their stiffness.     

Hybridization between Quercus robur and Q. petraea in a mixed oak stand in Denmark

? Hybridization and mating pattern between Quercus robur and Q. petraea was studied in a 5.8 ha mixed forest stand in Jutland, Denmark which comprises in total 135 Quercus robur and 230 Q. petraea trees. Classification of the oak trees into species was performed using canonical discriminant analysis of a range of leaf morphological traits. Adult trees (365) and offspring (582) were genotyped with eight microsatellite markers. Seedlings were sampled in 2003 and acorns were collected in 2004.

? Mating patterns of Q. robur and Q. petraea are expected to be different in the northern range of the distribution area and a larger hybridization rate is expected. It is further expected, that pollination from outside sources will be relatively less in small fragmented forest management systems compared to large scale oak forest. The conclusions should be verified through repeated year to year analysis of the mating pattern.

? Phenological studies revealed that there was no difference in flowering time between species. Data for the adult trees revealed no significant departures from Hardy-Weinberg proportions and there was weak, but significant spatial genetic structure, which supports the idea that the stand is of natural origin. Spatial genetic structure in the first distance class is stronger for Q. petraea. The genetic composition of the offspring was remarkably consistent from year to year. Paternity analysis revealed that, on average, 85% pollination came from fathers within the stand. The direction of the pollen flow varied from year to year. Inter-specific hybridization was high and ranged from 15–17% and from 48–55% for Q. petraea and Q. robur mothers respectively. Paternity analysis revealed that the population was basically outcrossing and only 3.7% of the analysed progeny were the product of selfing. Over the two years of study, approximately 200 trees contributed to the paternity of the next generations.

? The study confirms earlier studies with a greater tendency for Q. robur mothers to produce hybrid seeds than Q. petraea mothers. The rate of hybridization is higher in this Danish stand than in comparable studies elsewhere in Europe. Gene flow from outside sources are relatively low.

     

Summer drought and canopy opening increase the strength of the oak seedlings–shrub spatial association

? Context

The knowledge of how shrub–seedling interactions vary with summer drought, canopy opening, and tree species is crucial for adapting forest management to climate change.

? Aims

The aim of this study was to assess variation in shrub–oak recruitment associations along a south–north drought climate gradient and between two levels of canopy cover in coastal dune forest communities in a climate change-adapted forest management perspective.

? Material and methods

Mapped data of associational patterns of seedlings of three oak species with interspecific pooled shrubs were analyzed using a bivariate pair correlation function in 10 (0.315 ha) regeneration plots located in forest and recent gap sites along the climate gradient. An index of association strength was calculated in each plot and plotted against a summer moisture index.

? Results

The association strength increased with increasing summer drought from wet south to dry north and from closed forests to gaps.

? Conclusion

Consistent with facilitation theory, our results suggest that climate change may shift associational patterns in coastal dune forest communities towards more positive associations, in particular in canopy gaps. In a perspective of climate change, foresters may need to conserve understory shrubs in gaps in order to promote oak species regeneration.     

Intra- and interspecific competition differently influence growth and stem quality of young oaks (Quercus robur L. and Quercus petraea (Mattuschka) Liebl.)

Context

Cluster planting has become a conventional establishment method for oaks in Central Europe, where the spacing of seedlings within clusters varies between ‘nests’ (0.2?×?0.2 m) and ‘groups’ (1?×?1 m). Although the space between clusters is expected to fill with voluntary regeneration, its competitive effect on oak growth and quality had not been studied yet.

Aims

The aim of the study was to analyse the effects of inter- and intraspecific interactions on growth and quality of oaks grown in cluster plantings by quantifying the influence of neighbouring trees. In addition, we analysed whether the spatial position of oaks within groups (inner section or periphery) influenced their quality development.

Methods

Using Hegyi’s competition index, the influence of competition from intra- and interspecific trees from early, mid- and late-successional species, on diameter, height, slenderness and quality (length of branch-free bole) of 10- to 26-year-old oaks grown in cluster planting stands was quantified at seven sites in Baden-Württemberg and Hessen, Germany.

Results

In general, mid- and late-successional trees exerted a stronger competitive influence on growth of target oaks in clusters than the conspecific oaks and pioneer tree species. Oak quality development benefited from intraspecific competition, but self-pruning was not further promoted through additional interspecific competition. Within groups, inner oaks had a higher probability of developing into potential future crop trees than outer oaks.

Conclusion

Our study showed that intra- and interspecific competition had different effects on target oak trees and that these effect differed between nest and group plantings. The development of naturally regenerated and planted trainer trees in group plantings should be monitored carefully and if necessary be controlled through thinning or pollarding.     

Genetic structures of Calophyllum inophyllum L., a tree employing sea-drift seed dispersal in the northern extreme of its distribution

? Context

Little is known about the potential of gene flow and resulting genetic structures of trees employing sea-drifting seed dispersal in island populations.

? Aims

Current genetic structure and the magnitude of historical gene flow were estimated in island populations of Calophyllum inophyllum L., a typical plant employing sea-drifting seed dispersal.

? Methods

Samples were collected from the northern extreme of the species’ distribution (Taiwan and the Sakishima, Daito, and Ogasawara Islands, Japan) and genotyped using 15 EST-SSR markers. Genetic differentiation (F _ST and AMOVA), genetic structure (STRUCTRE analysis), and historical gene flow (assignment testing) were determined.

? Results

Frequent gene flow within and rare gene flow among island groups was determined using assignment testing. Clear genetic structures were also detected using the STRUCTURE analysis, which demonstrated differentiation between dominant clusters among geographically constructed island groups.

? Conclusions

The potential for gene flow via sea-drifting seed dispersal was high, and this was possible even among small islands. However, the extent and frequency of gene flow were not great enough to prevent genetic differentiation in a range of over a few hundred kilometers.