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.

     

Intrapopulation and Interpopulation Genetic Variation of Quercus in Denmark

Native Danish oak stands are fragmented and decreasing because of the extensive use of foreign seed sources. Therefore, the population structure of natural Danish oak stands was analysed by means of six polymorphic enzyme loci. A total of 17 stands of Quercus robur L. (including an ''outgroup'' sample from The Netherlands), seven Quercus petraea [Matt.] Liebl. and two putative hybrid stands were included in the study. The average genetic diversity was similar for the two species as well as for the putative hybrid stands: 0.25 for Q. robur, 0.27 for Q. petraea and 0.26 for the hybrid stands. The genotypic proportions at two (Pgm and Mnr) of the six loci showed many significant deviations from Hardy-Weinberg expectations, always with an excess of homozygotes, whereas the remaining four loci accorded to Hardy-Weinberg proportions, suggesting a low level of inbreeding. The differentiation of the Q. petraea and Q. robur populations was quantified with Wright's F-statistics. The within-species component was low, 0.022, reflecting the wind-pollinated reproductive mode of the two species. The betweenspecies component was 10-fold higher, 0.235, indicating two separate groups. A phylogenetic tree estimated from allele frequencies also supported the presence of these two groups. Despite the two well-separated groups in the tree, it was not possible to assign all individuals to the species to which they supposedly belonged. In total, 10% of the Q. robur individuals and 14% of the Q. petraea individuals were assigned to the other species, suggesting a limited amount of introgression between the two species.     

AN ENUMERATION OF THE MATURE OAK IN THE FOREST OF BERE

The paper describes the methods used in the field and in theoffice to determine the standing volumes and increments of 100-and 130-year old oak in the Forest of Bere, Hampshire, duringworking-plan revision. The figures obtained show that the standingvolumes and increments of sessile oak (Quercus petraea Liebe)are considerably higher than those of pedunculate oak {Q. roburL.) on the three sites investigated, namely, clays, sands, andgravels, while for both species sands appear to be the mostfavourable and gravels the least favourable soil types. Therewas little difference in the mean values for increment and standingvolume between the 100- and the 130-year old pedunculate oak.     

PRELIMINARY LIGHT-INTENSITY STUDY UNDER SESSILE AND PEDUNCULATE OAK

While study by continental and American workers has continuedover a considerable period, little has been published in connexionwith forestry. A method is described using photoelectric cellphotometers and the results given of measurements made undercomparable stands of sessile and pedunculate oak (Quercus petraeaLiebe and Q. robur L.). These results confirm that sessile oakcasts more shade than pedunculate oak.     

Variation in wood volatile compounds in a mixed oak stand: strong species and spatial differentiation in whisky-lactone content

The effect of species and ecological conditions on oak volatile extractive content was investigated in an evenaged (100 years) stand located in western France. The sample included a total of 286 trees (118 sessile, 158 pedunculate and 10 oaks with an intermediate morphology) growing in contrasted environments (plateau, intermediate slope, small valley). The main factor influencing oak extractives level is species. The effect of the local environment appears negligible. No correlation between ring width and volatile extractive content was found. Q. petraea is significantly richer than Q. robur in eugenol and whisky-lactone (10.8 vs. 0.6 μg/g). However, two groups of sessile oaks could be identified, one poor and one rich in whisky-lactone. Among the latter, either the cis or the trans stereoisomer was predominant, suggesting that their production is not independent. A strong spatial structure was detected for whisky-lactone (cis-, trans- and total whisky-lactone, for the two species combined but also for Q. petraea alone in the case of the cis isomer).     

OAK

Four basic stages in the life of the oak wood (Quercus roburand Q. petraea) are defined and the factors which should influencethe manager's mind when he weighs density of stocking and lengthof bole against rotation to meet his particular requirementsare considered briefly in non-technical language.     

Oak in Wessex An Account of Field Studies 1982-4

The Wessex Silvicultural Group spent three years studying themanagement and silviculture of the oaks (Quercus robur L andQ petraea (Matt) Leibl) in the Wessex area. Low input methodsof establishment including the use of tree shelters and thetending and thinning of younger oak stands together with controlof timber quality were examined in twenty-one forests and estates.Environmental values were acknowledged. While Q robur has previouslypredominated, Q petraea is now generally preferred.     

The effect of heavy or 'free growth' thinning on oak (Quercus petraea and Q. robur)

Free growth is a type of heavy thinning which aims to maximizediameter increment and produce valuable timber on a relativelyshort rotation. An experiment is described which investigatedthe application of free growth to a stand of oak {Quercus petraeaand Q. robur) planted in 1930. At age 58 free growth thinninghad resulted in a mean diameter at breast height of 39.0 cmwith an estimated mean tree volume of 0.98 m³, compared with29.3 cm and 0.52 m³ for equivalent crown thinned trees. Discountedcash flow calculations showed that free growth thinning of oakcould be justified using a 3 per cent discount rate assumingthat the increased intensity of pruning results in a large proportionof veneer quality timber; an independent assessment indicatedthis may be possible. Free growth thinning of oak is not a commonpractice in British broadleaved silviculture probably becauseof the cost of controlling epicormic shoots. It is suggestedthat it may be more appropriate to other species such as ash,sycamore and wild cherry.     

Genetic control of wood colour, density and heartwood ellagitannin concentration in European oak (Quercus petraea and Q. robur)

Pressler cores of young clones of Quercus petraea and Q. roburwere analysed from two sites in Germany. Variation of wood colour(defined by the CIELAB system), density and heartwood ellagitanninswas examined between clones, species and sites. Similar studieswere made of cores from another trial of 20 half-sib familiesof parent trees deriving from five German forests. The resultsfrom both the clonal and progeny trials indicate that heartwoodellagitannin content and wood density are under strong geneticcontrol, while sapwood and heartwood colour are less so. A largeproportion of between-clone variation in wood density and ellagitanninlevels occurred between the two species, while these propertiesvaried little among ramets of the same clone grown on two contrastingsites, despite a difference in growth rates. A large proportionof the total variation among progeny was attributed to forestorigins and could not be attributed clearly to either geneticor environmental causes because of limitations of the samplingdesign     

Detecting interspecific and geographic differentiation patterns in two interfertile oak species (Quercus petraea (Matt.) Liebl. and Q. robur L.) using small sets of microsatellite markers

Genetic analysis was carried out in order to provide insights into differentiation among populations of two interfertile oak species, Quercus petraea and Quercus robur. Gene flow between the two species, local adaptation and speciation processes in general, may leave differential molecular signatures across the genome. Three interspecific pairs of natural populations from three ecologically different regions, one in central Europe (SW Germany) and two in the Balkan Peninsula (Greece and Bulgaria) were sampled. Grouping of highly informative SSR loci was made according to the component of variation they express—interspecific or provenance specific. ‘Species’ and ‘provenance discriminant’ loci were characterized based on F_STs. Locus specific F_STs were tested for deviation from the neutral expectation both within and between species. Data were then treated separately in a Bayesian analysis of genetic structure. By using three ‘species discriminant’ loci, high membership probability to inferred species groups was achieved. On the other hand, analysis of genetic structure based on five ‘provenance discriminant’ loci was correlated with geographic region and revealed shared genetic variation between neighbouring Q. petraea and Q. robur. Small sets of highly variable nuclear SSRs were sufficient to discriminate, either between species or between provenances. Thus, an effective tool is provided for molecular identification of both species and provenances. Furthermore, data suggest that a combination of gene flow and natural selection forms these diversity patterns. ‘Species discriminant’ loci might represent genome regions affected by directional selection, which maintains species identity. ‘Provenance specific’ loci might represent genome regions with high interspecific gene flow and common adaptive patterns to local environmental factors.     

The reproductive success of a Quercus petraea × Q. robur F1-hybrid in back-crossing situations

A 56 year old Quercus petraea × Q. robur F1-hybrid was back-crossed to both parental species. Pollen mixes were applied and paternity assigned to offspring based on microsatellite markers. The studied Q. petraea × Q. robur hybrid proved highly fertile and back-crossed well with both Q. robur and Q. petraea with slight but not significant preference for Q. robur. The results do not support the hypothesis about highly unidirectional gene flow between Q. robur and Q. petraea towards Q. robur as the observed back-crossing ability of the hybrid opens a route for nuclear gene flow from Q. robur to Q. petraea. However, Q. petraea × Q. robur hybrids may be rare in nature and the results do not tell us if the (probably more common) reciprocal hybrid also back-crosses easily to Q. petraea.     

The Dependence of Quercus robur and Q. petraea Seeding on Cotyledon Potassium, Magnesium, Calcium and Phosphorus during the First Year of Growth

Acorns collected from a range of Quercus robur L. and Q. petraea(Matt.) Liebl. populations were germinated and grown in an unheatedgreenhouse using a standard commercial compost. Total seedlingdry weight did not increase until the end of the growing season,with Q. robur and Q. petraea seedlings showing mean dry weightincreases of 211 per cent and 414 per cent respectively. Oakseedlings appear to depend upon cotyledon reserves of carbohydrate,P, K and Mg during most of the first year of growth, but relyon external sources of Ca. Seedlings become independent of thecotyledons at the end of the first growing season.     

A comparison of the autecology of Quercus robur L. and Q. pyrenaica Wild.: present habitat in Galicia, NW Spain

The biotopes occupied by pedunculate oak (Quercus robur L.)and rebollo oak (Quercus pyrenaica Willd.) in Galicia were characterizedusing different parameters, enabling a comparative analysisof its habitats. Thirty-nine plots of pedunculate oak and 40of rebollo oak were inventoried and 25 ecological parameters(physiographic, climatic and edaphic) were measured to describethe biotopes, along with 14 stand parameters to characterizethe structure and the silviculture of these forests; some ofthem were used in previous works about the autecology of bothspecies. Quercus robur was found to occupy a wider range ofecological habitats and their distribution is oceanic; Q. pyrenaicapresents a more Mediterranean phytoclimatic position. The resultsindicated that the distribution of oak forests in Galicia ismore closely related to physiographic and climatic featuresthan to edaphic, because the nature of the substrates that theyoccupy is similar. Analysis of fragility/aggressiveness betweenboth species and chestnut shows that the aggressiveness of bothoaks, mainly pedunculate, was much higher, except at high altitudes,and in areas with a high mean temperature, where rebollo ismore aggressiveness. Comparison of the two Quercus shows formost of the parameters that Q. robur is the more aggressivespecies.     

A Possible Indicator of Shake in Oak: Relationship between Flushing Dates and Vessel Sizes

Previous work by Savill (1986) has shown that oak trees (Quercusrobur and Q. petraea) with large earlywood vessels appear tohave a much greater predisposition to shake than trees withsmaller vessels. The aim of the investigation described herewas to determine whether there are any externally visible characteristicscorrelated with vessel size, so that shake-prone trees can berecognized in the field. It was found that trees that flushlatest within a population tend to have the biggest vessels.Possible reasons for this correlation are discussed briefly.The finding has the practical value that oaks which are predisposedto shake can be marked at flushing time and removed in thinningoperations.     

A Comparison Between European Continental and British Provenances of Some British Native Trees: Growth, Survival and Stem Form

Data describing the growth, survival and stem form of: a) Europeancontinental, and b) British provenances of trees native to Britainwere collated from Forestry Commission records. The growth ratesof European continental provenances of Scots pine (Pinus sylvestris),silver birch (Betulo pendula), sessile oak (Quercus petraea)and common alder (Alnus glutinosa) were inferior to those ofBritish provenances in 90 per cent of cases. European provenancesof these species also showed inferior survival. The growth ratesof continental provenances of beech (Fagus sylvatica) were superiorto British provenances in about 50 per cent of cases, provenancesfrom northern France, Belgium and Holland being the fastestgrowing. Relationships between the relative height growth offoreign provenances and latitude were demonstrated for Scotspine, silver birch and beech (relative height growth = the heightgrowth of a foreign provenance expressed relative to that ofBritish provenances at the same site). The continental provenancesthat grew best originated at 0–4° south of the latitudesof the planting sites. Relative height growth decreased bothnorth and south of these latitudes. In the case of oak, relativeheight growth did not vary widely between origins. The survivalof continental provenances of Scots pine was particularly pooron exposed upland sites. The stem form of British provenancesof Scots pine was intermediate between Fennoscandian provenancesand provenances from the rest of Europe. In oak and beech therewas generally little difference in stem form between Britishand continental provenances, but the best continental provenances(Belgian beech; French oak) were most reliable. The growth andsurvival of native trees is interpreted in the light of thepost-glacial history of these species. The implications regardingthe continued use of seed of native hardwoods imported fromcontinental Europe are discussed.     

Differences in composition of endophytic mycobiota in twigs and leaves of healthy and declining Quercus species in Italy

The composition of the endophytic fungal assemblages in Quercus cerris, Q. pubescens and Q. robur, three oak species most susceptible to decline, was investigated in Fagarè (Padova), Ulignano (Pisa), and Radda in Chianti (Firenze). The endophytic assemblages were studied as a function of health status of the whole tree and of twigs and leaves. The effect of station altitude was tested on Q. cerris. Samples were gathered from healthy and declining trees. In April (bud opening time) 20 twigs and 10 leaves were collected from each tree. Fungi were isolated from twig segments and leaf portions. A total of 23 fungal genera with 27 different species were isolated, in addition to a few sterile mycelia. Eleven species were found to be common to all three oak species, while other species were found only on one oak species, suggesting host specificity. Among the fungal species isolated, some were known to be pathogenic (Apiognomonia quercina, Colpoma quercinum, Diplodia mutila, Phomopsis quercina). The colonization frequency (CF) of pathogenic species varied between 0.9% for A. quercina in Q. cerris and 60.2% for P. quercina in Q. robur. Analysis of variance showed a statistically significant difference among the oak species tested. The CF was higher in declining trees in comparison with healthy trees, and also in twigs vs. leaves. The CF was found to be higher for Q. cerris trees growing at sea level as compared with those situated at an elevation of 350–400 m. In stands where, on account of particular ecological conditions, pathogenic behaviour is displayed simultaneously by more than one endophytic fungal species, trees accentuate their decline.     

THE STORAGE OF ACORNS IN WATER

Acorns of sessile oak (Quercus petraea) will not tolerate prolongedsubmergence, but those of pedunculate oak (Quercus robur) survivevery well in regularly changed distilled water or tap-water.Survival is considerably worse in basic river water; the differencefrom tap-water is likely to be due either to the greater activityof organisms promoting decay, or to the prevalence of loweroxygen-tensions in river water. Storage in water under suitablycontrolled conditions might be a useful practical procedurewith pedunculate acorns.     

Growth stimulation of young oak plantations inoculated with the ectomycorrhizal fungus Paxillus involutus with special reference to summer drought

Nursery-grown Quercus petraea and Q. robur seedlings were inoculated with the ectomycorrhizal fungi Paxillus involutus, Hebeloma crustuliniforme or Laccaria laccata. At the end of the first growing season, the seedlings with roots colonized with the inoculated fungi were outplanted at two sites in northeastern France in 1985 or 1988. P. involutus was found to be the most competitive and efficient of the three fungi tested; its mycorrhizas were present 7 yrs after outplanting and it markedly improved the growth of both oak species. In the case of Q. robur, the growth stimulation was more marked on years with a dry summer. These results are discussed in terms of water relations and of competitiveness of the introduced versus resident symbiotic fungi.     

Hosts and distribution of Collybia fusipes in France and factors related to the disease's severity

Using data collected by two different observation networks, it was possible to clarify the distribution of Collybia fusipes in France. It is frequently found in the western and northern parts of France, mainly on Quercus rubra and Quercus robur, and to a somewhat lesser extent on Quercus petraea. When the relative areas covered by these species are compared, it is evident that Q. rubra is much the most susceptible of the three species. The fungus has been less frequently reported in the south‐eastern quarter of France, where Quercus ilex and Quercus pubescens are the dominant species. Complementary observations carried out at the National Arboretum of Les Barres indicate that a large number of oak species are susceptible to infection by this fungus, particularly those in the subgenus Erythrobalanus. These observations are discussed with reference to reports of this fungus throughout Europe. In a Q. rubra stand, fruit‐body clumps were monitored and root damage was assessed on 73 final‐crop trees and 40 codominant trees. The results suggest that C. fusipes does not preferentially infect weakened trees. Both the severity of root damage and the number of fruit‐body clumps increased with increasing depth to signs of waterlogging.     

Characteristics of the Cryphonectria parasitica isolated from Quercus in Slovakia

The occurrence of chestnut blight (Cryphonectria parasitica) on oaks in mixed chestnut‐oak forests was studied in 2003–2008 in Slovakia. Infections on living Quercus trees were found at four of seven localities. The disease incidence on oaks ranged from 1.3% to 15.8%. The symptoms on infected oaks were similar to those on chestnut, but less conspicuous. Cankers of C. parasitica were found only on Quercus robur and Q. petraea. A total of 22 isolates of C. parasitica, all virulent, were isolated. Each site yielded only a single vc type (EU12 or EU13). Field inoculation experiments on chestnuts with seven strains of C. parasitica from oaks and an isolate from Castanea sativa showed no differences in virulence. On Quercus robur stems, the cankered area was significantly smaller than on C. sativa and the cankers developed very slowly.