Báo cáo khoa học: " Intraspecific variation of growth and adaptive traits in European oak species"

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Review article Intraspecific variation of growth and adaptive traits in European oak species J Kleinschmit Lower Saxony Forest Research Institute, Department of Forest Tree Breeding, W-3513 Staufenberg OT Escherode, Germany Summary — According to various reports, 200-450 oak species including hybrid populations exist worldwide, with 24 of these - including 3 hybrid forms - having their natural range within Europe. all belong to the subgenus Lepidobalanus. The most important section is robur with 21 spe- They cies. The European species belong to both deciduous (15 species) and evergreen (9 species) oaks. Some difficulties in clear morphological and physiological definition of the species involved is caused by hybridization. Most studies into intraspecific variation exist for the species Quercus robur L and Quercus petraea (Matt) Liebl. Some information is available for Quercus cerris L, Quercus virginiana Ten (= Q dalechampii Wenz), Quercus ilex L and Quercus suber L, based on comparative planta- tions. In addition, a considerable number of morphological, physiological and biochemical studies based on natural populations exist which are not completely covered in this review paper. For most characters observed, oaks exhibit a wide variation. This is not only the case for morphological traits of pollen, seed, wood and plants but also for physiological traits and phenology which have great adaptive importance. Provenance experiments and progeny tests started as early as 1877. Most of these were only of local importance. This is partly due to the fact that acorns can only be stored for a limited period and flowering is irregular. However, the results available show that the choice of prov- enance can be important for the successful economic management of plantations. Variability of leaves, phenology, form, growth, wood and bark, roots, seed and flowering has been discussed sep- arately. Improvement via selection and testing seems to be promising due to the considerable be- tween-population and within-provenance variation. Vegetative propagation has been developed for some species by grafting, cutting propagation and in vitro propagation. Tree breeding approaches have also been discussed. Quercus / morphology / provenance / progeny test / intraspecific variation / vegetative propa- gation Résumé — Variabilité des caractères de croissance et d’adaptation chez les intraspécifique de chênes. Selon les auteurs, de 200 à 450 espèces de chêne, y compris espèces européennes les populations hybrides, ont été identifiées sur le globe. Vingt-quatre d’entre elles, comprenant 3 formes hybrides, ont été reconnues en Europe. Elles appartiennent toutes au sous-genre Lepidoba- lanus. La section la plus représentée est robur, avec 21 espèces. Les espèces européennes sont à feuilles caduques (15 espèces) ou persistantes (9 espèces). L’hybridation naturelle rend la classifi-
cation difficile sur la base de critères morphologiques ou physiologiques. La majorité des études de variabilité iniraspécifique concerne Quercus robur L et Quercus petraea (Matt) Liebl. Des informa- tions partielles, issues de plantations comparatives, sont disponibles pour Quercus cerris L, Quercus virginiana Ten (= Q dalechampii Wenz), Quercus ilex L et Quercus suber L. Par ailleurs, de nom- breuses références relatives à des études de variabilité in situ de caractères morphologiques, physio- logiques et biochimiques existent dans la littérature; elles ne sont qu’incomplètement évoquées dans cette revue. Pour la majorité des caractères, l’amplitude de variation est très grande. Il s’agit non seu- lement des caractères relatifs au pollen, à la graine, au bois, aux arbres, mais aussi aux caractères physiologiques et phénologiques, qui revêtent une grande importance adaptative. Les premiers tests de provenances et de descendances remontent à 1877. Ils ne comprenaient que les provenances lo- cales, à cause de la difficulté à conserver les graines et l’irrégularité des fructifications. Les résultats de ces plantations montrent cependant que le choix de la provenance est primordial pour le succès économique du reboisement. La variabilité de la morphologie des feuilles, de la phénologie, de la croissance, de la forme, du bois et de l’écorce, des racines, des graines et de la floraison est égale- ment évoquée dans une partie séparée. L’amélioration dans des programmes de sélection peut abou- tir à des gains élevés compte tenu de l’importance de la variabilité intraspécifique et individuelle. La multiplication végétative par greffage, bouturage et culture in vitro a été mise au point pour certaines espèces. Les méthodes d’amélioration génétique sont également mentionnées. Quercus / morphologie / provenance / test de descendance / variabilité intraspécifique / multi- plication végétative INTRODUCTION question whether Q robur and Q petraea are separate species in this sense, since intermating frequently. occurs The genus Quercus is represented by 200 The oak population distribution appears (Neger and Münch, 1950), 320 (Krahl- to be related to ecological site types, ie Urban, 1959) or 450 (Krüssmann, 1978) taxonomic speciation and ecological segre- species from the temperate to the tropical gation are closely linked (Grandjean and zones. The differences in the numbers are Sigaud, 1987). Species evolution is still un- partly explained by the definition of hybrid derway in many cases and genetic isola- forms as separate species (Krüssmann, tion is not complete. 1978), partly by the species concept (bino- mial classical or biological) and the sub- If we try to apply the biological species = division of ecological forms into species. concept to oak, we find that the boundaries Species delineation is difficult if popula- be readily identified. So many cannot tions intermate and gene flow attains dif- classical species hybridize in Quercus that ferent degrees of intensity even with differ- the genetically defined concept must be ent subpopulations and individuals. one of extraordinary complexity (Burger, 1975). The genetic (biological) species in If apply the biological species we con- oaks is simply too complex and too difficult cept (Mayr, 1963) with the following defini- to recognize to serve as the basis of a tion: "Species are groups of actually or po- stable and functional system of nomencla- tentially interbreeding populations, which ture. are reproductively isolated from other such groups", the number of oak species would The genus is subdivided into 3 subgen- be reduced considerably. One may even with 1-7 sections each (table I). era
In this paper we follow the species defi- ever, oak trees constituted a valuable base nition of Krüssmann (1978). Twenty-four for human life in the past. The fruits served oak species and different hybrid forms ex- as a nutritional base for their animals and ist in Europe, partly as introgression zones the wood was invaluable for construction, in the natural range. Fifteen species are tools and shipbuilding. Oak was therefore deciduous, 9 species evergreen (table II). extensively planted and Quercus silvicul- Only 8 of these are of economic impor- ture was developed at an early date tance. (Krahl-Urban, 1959; Thirgood, 1971). In for oak silviculture France, ship- Oak forests considerable per- cover a construction was given active encourage- centage of the total woodland area in most ment by Colbert as early as 1661; around European countries. They exceed 30% in 1700, Carl XII established oak stands in some countries (Greece and France), of- Sweden with seed imported from Poland ten cover 25% in others (eg UK, Romania, (Krahl-Urban, 1959). Hungary, Belgium) and only in a few cases do they comprise < 10% of the forest area Nineteen oak species are natural to the (eg Germany, Czechoslovakia, The Neth- former USSR. But more than 60 species erlands). have been introduced from different regions of the world; most have been The natural range of oak species has successful, reproduce under plantation been drastically influenced by human activ- ity. Since oak forests covered rich sites at conditions (Trofimenko, personal commu- low elevations, the majority of these have nication) and hybridize with local popula- been converted into agricultural land. How- tions.
The transfer of populations had oc- species. The variability in all characteris- curred over considerable distances. Large considerable within the tics observed was quantities of oak seed (eg, up to 4 000 species. tonnes annually) were imported to Ger- For the Quercus species with extended many mainly from southeastern Europe natural ranges, such as Q robur and Q during the last century (Lüdemann, 1962). petraea (fig 1) certain geographical trends These stands hybridized with local popula- can be observed with latitude, longitude tions. Therefore the pattern of variation we and elevation. Since all these studies of find in the economically important oak spe- phenotypic variation within and between cies today may be far from natural. Studies stands do not enable separation of genet- of phenotypic variation in ’natural’ popula- ic and environmental components, only a tions generally exhibit a surprisingly high few are discussed in more detail in this variability in all characters studied. Growth, paper. stem form, crown morphology, formation of epicorms, wood characters, flushing, bud set, lammas shoot formation and attack by INTRASPECIFIC VARIATION Microphaera alphitoides and Tortix virida- na, for example, were found to differ from The intensity of research into intraspecific one population to another and quite often variation is largely dependent upon the ec- were even more variable within stands onomic importance of the species. Variabil- (Krahl-Urban, 1959; Weiser, 1964). Leaf ity is greatly influenced by the extent of the number per branch unit and leaf size vary natural range. Extreme differences exist with location and stand age, eg, in Q cer- between the oak species. Since almost no ris, Q frainetto, Q pedunculiflora, Q pe- provenance or progeny studies have been traea, Q pubescens and Q robur (Dissescu carried out with oak species other than and Coca, 1973). The variation in leaf Q robur and Q petraea, these will be dis- characteristics decreases, however, with cussed first.I am aware that not all the age (Semerikov, 1974). Pollen size and literature can be covered by our central structure are different for species groups system andI am grateful to those col- and for species. Smit (1973) divides Quer- leagues who have provided me with addi- cus pollen grains into 3 groups according tional information on provenance and prog- to their morphology: 1) Q robur/petraea eny tests which have not yet been type; 2) Q ilex/coccifera type; and 3) Q published. suber type. Colombo et al (1983) were able to differ- entiate between the species of the Mediter- Quercus cerris ranean area on the basis of pollen mor- and Quercus dalechampii phology and size. However, individual variability is considerable and sampling has to be extensive. A comparative plantation with 9 oak spe- cies was established in 1982 in Levice For- Kissling (1977) studied the hairs on the est Enterprise in West Slovakia. Leaf area lower side of the leaves of Q pubescens, (Masarovicova and Pozgaj, 1988) was fol- Q petraea, Q robur and Q cerris and found lowed for 3 of these species. Variability in that these were a good character for differ- individual trees within the species Q cerris entiating the 4 species. The hybrid forms and Q dalechampii was larger than be- had heteromorphous hairs which were intermediate between those of the parent tween the species.
Quercus conferta the Harz mountains, 975 m in the Black Forest, 1 185 in the Central Alps and 1 600 m in the French Alps (Rameau et al, Morphological variability of Q conferta with- 1989). Q robur remains about 200-300 m in the Strandsha Mountains has been stud- lower in the mountains. According to Krahl- ied by Garilov and Stojkov (1978). Thirty- Urban (1959), Q petraea has its optimum five morphological forms have been de- in France between the Seine and the scribed and 3 ecological forms differentiat- Loire, in Germany in Spessart, the Pfälzer ed. The stem form is better in the meso- Forest, the Mosel region, the north-eastern phillous form and worse in the xerophytic plains of Lower Saxony, Mecklenburg, form. Pommern, and Brandenburg, and also in Croatia and Bosnia. Q robur grows best in the Rhine valley, in the Danube-Drau- Quercus ilex Save (Slavonia) lowlands and in northern Germany. The genetic component is influ- The growth of 2-year-old seedlings of 46 enced by the species itself. provenances was correlated with acorn a sequence of subspecies Q robur has size but not with site parameters or geo- Menitzky (1971) concluded that Q pe- and graphic variables of the location of origin dunculiflora Koch, Q longipes Stev and (Bonani et al, 1988). Local provenances Q erucifolia Stev not autonomous spe- are were slow growing. Pollination occurred cies, but form the southern subspecies of between individuals which flower synchro- Q robur. Their formation was substantially nously (Yacine and Lumaret, 1988). Vari- influenced by introgressive hybridization ability in phenology was found to be con- with xerophytic oaks, especially Q pubes- siderable both between individuals within a cens and to a lesser degree Q petraea. stand and between populations (Yacine Q virgiliana Ten is an intermediate form and Lumaret, 1989). between Q robur and Q pubescens. Q pe- dunculiflora and Q virgiliana are listed as separate species here following Krüssmann Quercus suber (1978) (table II), in spite of the fact that they could also be ranked as subspecies. The species is subdivided into 4 intraspe- For Q petraea, 4 subspecies have been cific taxa depending upon the lifespan of described (Jovanovic and Tucovic, 1975): leaves and duration of acorn development Q petraea Liebl ssp petraea Menits, Q pe- (1 or 2 years), (Globa-Mikhailenko, 1973). traea Liebl ssp iberica (Stev) Krassil, Q pe- The high variability of these characters in traea Liebl ssp dshorchensis (Koch) Men- the species is demonstrated. its and Q petraea Liebl ssp medrvediewii (Camus) Menits. Q dalechampii Ten is an intermediate form between Q pubescens Quercus petraea and Quercus robur and Q petraea in the contact zones of both species; it is however, kept separate here The variability of these species is affected too. Studies on intrapopulation variability of by environmental and genetic factors. The Q petraea show differences in leaf shape environment of the habitat of both species and size, such as platyphyllous, laciniate is very variable. Within its natural range and longifoliate (Schwarz, 1936-1939), (fig 1),Quercus petraea covers an eleva- which have been separated into different tional range from sea level up to 600 m in varieties.
For Q robur and Q pefraea, a number of be dominant. On rich, humid sites, can Q robur usually prevails and on warm, dry provenance experiments have been estab- lished since the beginning of this century sites Q petraea is dominant. On sites with mosaic pattern of dry and wet areas as (table III). The information obtained from a in the mountainous regions of Germany, these experiments is the basis of the fol- both species are sympatric and show in- lowing sections of this paper. tensive introgression. Hybridization Leaves A discussion of variability in these species Numerous studies have compared leaves is not possible without looking into their hy- of both species (eg Oelkers, 1913; Rush- bridization. Since this topic has been treat- ton, 1976, 1978; Staszkievicz, 1970). ed by Rushton and others (this volume) I Some characteristics are typical of the spe- will present here only some major results. a continuous variation ex- cies; however, Experiments with controlled pollination ists from one species to the other. Will- demonstrate that hybridization of Q robur komm (1875-1887) considered that and Q petraea is easier with Q robur as Q robur had the highest variability of all the mother (Dengler, 1941; Aas, 1988, broad-leaved tree species, mainly in the 1990). Considerable differences in crossa- size of the leaves, their shape, crenature, bility exist on an individual level, covering structure and leaf color. the whole range from infertility to full fertili- Semerikov (1974) suspected a stabiliz- ty as compared to the within-species ing selection for leaf characteristics of iso- crosses. lated populations. The first statistical analy- It is of interest that isolation seems to separation of Q robur and sis for the be more developed in Q petraea as com- Q petraea, using leaf and fruit characteris- pared to Q robur. One could speculate that tics, was made by Oelkers (1913), who re- adaptation to the more specific site condi- garded both species as subspecies. He tions of Q petraea (dry, fewer nutrients) also observed the phenological variation needs higher specialization and that this within and between the species. He found can only be maintained by better protec- considerable variation in all characters ob- tion against introgression. Q robur, on the served and an overlap in all characteris- other hand, usually grows under more opti- tics. Burger (1914) summarized the knowl- mal ecological conditions and thus can edge of morphological differences in both maintain a broader gene pool. species and established a provenance ex- Morphological studies show that in addi- periment in the nursery. tion to pure and mixed stands of both spe- cies, there are also stands with hybrid Phenology forms and stands in which the latter forms prevail (Burger, 1921; Seitz, 1923; Opper- Under the same site conditions, Q robur mann, 1932; Krahl-Urban, 1959; Cousens, has a longer vegetative period with earlier 1965; Gardiner, 1970; Olsson, 1975a; flushing and later bud set. However, ex- Rushton, 1978, 1983; Dupouey, 1983, treme provenance differences exist (Krahl- Spethmann, 1986b; Lower Saxonomy For- Urban, 1959). Provenances from regions est Research Institute, Escherode, 1986- with shorter vegetative periods flush earlier (Oppermann, 1932), but this trend is not 1991) (fig 2). Depending upon the ecologi- the other consistent (Cieslar, 1923). cal conditions of the site, one or
The differences between the results of Burger (1921) compared Q robur and Q petraea from the same region around Krahl-Urban (1959) and Burger (1921) can Zurich for flushing and bud set. Earlier easily be explained by ecotypic variation flushing at a young age was found in Q pe- and sampling. We studied 198 Q robur traea, while at advanced ages there stands and 183 Q petraea stands from were differences between the species. After northern Germany in the nursery. The fre- no germination, leaf color is red in Q robur. quency distribution for provenance mean Flowering and seed ripening were syn- flushing is given in figure 3. From this fig- chronous with considerable individual dif- ure, it is obvious that the 2 species do not ferences within species. differ significantly in flushing time. Q robur is more represented at the extremes. As Hauch (1909) found late flushing prove- early as 1923, Cieslar found considerable from Slavonia and Galicia, early nances ecotypic variation in flushing but a more cli- flushing provenances from Hungary and nal pattern with bud set with continental other southern sources.
and compared in provenance plantations. and northern provenances occurring early, Henrik (1973) found a 7-day difference in southern and oceanic provenances occur- flushing and a 3-day difference in bud set ing late. Menitsky (1971) stated that clinal in stands of Q robur in Poland. These were variation was hardly noticeable in oak. not correlated with wood characters. Krahl- Variation in flushing of mother trees is Urban (1959) described provenance differ- clearly inherited by their progenies, but ences in flushing of 3 weeks between there was no correlation between growth Slavonian (Q robur tardissima) and Ger- and flushing in Oppermann’s studies man provenances. (1932). In the Voronezh region, late flushing Different provenances from all over Eu- oaks show higher resistance to late frost rope of the same species show greater dif- and are therefore recommended for culti- ferences in flushing than between individu- vation (Verchenko, 1975). On the other als of the different species at the same hand, late flushing provenances have a location (Burger, 1949). For German prov- maximum water demand in July and trans- enances, the individual variability in pheno- piration exceeds early provenances by logical traits is larger than between prove- 31%; therefore Silina (1951) recommend- (Cieslar, 1923). nance means ed planting early flushing provenances on Bud set is earlier in Q petraea at the elevated areas where little water is availa- same location. Considerable differences in ble. leaf yellowing and fall were observed be- Southern provenances set buds later tween individuals of Q robur at the north- are more sensitive to early frost (Cies- and eastern limit of the natural range (Danilov lar, 1923). Provenances from central et al, 1972). Late yellowing leaves have France and the UK, for example, are less higher transpiration rates and lower solu- resistant to winter frost in central Germany ble carbohydrate contents. and Denmark than local provenances (Op- permann, 1932; Krahl-Urban, 1959). North- Phenological characteristics of Q robur provenances set buds earlier (Kienitz, also studied by Jevlev (1972a,b,c) ern were
1879) and are more resistant to early frost to urban conditions and selected adapted for this purpose (Thompson, 1986). Ex- (Cieslar, 1923); however, they do not use the growth potential of southern regions treme variability in form can occur, as (Oppermann, 1932). described by Oppermann (1932), like pendula, fastigiata, pyramidalis and creep- Lammas shoot formation is fre- more ing, straight or crooked trunks. Hauch quent in Q robur (Krahl-Urban, 1959; Jova- (1916-1921), Cieslar (1921), Oppermann novi&jadnr; and Tucovi&jadnr;, 1975) and thus suscep- (1932) and Krahl-Urban (1959) found a tibility to Microsphaera alphitoides is close relationship between the shape of higher (Rack, 1957). But again, prove- the crown and the trunk of the mother nance differences and individual variability trees and their progenies. The same was are considerable. Late bud-setting prove- true for stands with good and bad form. nances have a high incidence of lammas Oppermann gave the frequency distribu- shoots and thus high susceptibility to Mi- tion for progenies of good and bad stands crosphaera alphitoides and early frost. at age 17 years: good stands had 10 good, Summarizing the research on phenolo- 3 medium and 2 bad progenies; bad gy one can state: clinal and ecotypic pat- stands had 0 good, 1 medium and 5 bad terns of variation exist side by side. Bud progenies. Including stand- and single tree set shows a clearer clinal variation pattern progenies in the comparison, Opper- - than flushing. Individual variation is consid- mann found that: good mother trees had erable and exceeds variation between 61% good, 25% medium and 14% bad provenances of limited areas. progenies; bad mother trees had 15% good, 26% medium and 59% bad proge- nies. Form Within more limited geographically de- There are some general species descrip- fined regions, there is a clear indication tions showing the typical differences be- that stem form and crown form are under tween Q robur and Q petraea (Burger, strict genetic control. 1921; Krahl-Urban, 1959; Jovanovi&jadnr; and Krahl-Urban (1959) found high heritabili- Tucovi&jadnr;, 1975). Crowns of Q robur are ty for forking, multiple leader and stem usually irregularly forked with widely straightness in progeny tests. spaced, vigorous, crooked branches main- The transfer of provenances from a ly in a horizontal plane. Monoaxiality is milder climate with associated frost dam- rare. Q petraea, on the other hand, has a age is reflected in worse stem form and more regular crown with thinner branches, slower growth. But there are some broadly which are more evenly distributed and at a adaptable provenances. Early flushing can more acute angle. be associated with late frost damage and This general description excludes the the resultant form defects. Generally, Q ro- high variability within populations, between bur provenances from Lipovljana in Croatia provenances and the introgression prob- and from South Bohemia had the best lem. Both species show high variabilities in stem form (Cieslar, 1923; Oppermann, crown types, as demonstrated by Opper- 1932; Krahl-Urban, 1959) under different mann (1932) and Krahl-Urban (1959); growing conditions. these are inherited. Koloszar (1987) Krahl-Urban (1959) found 50-70% demonstrated that the excellent form of straight Q Q robur tardissima from Slavonia is main- robur trees in progenies of tained under a variety of plantation condi- stands in the Save Valley, 25-30% in tions. Narrow-crowned oaks are especially Q petraea from Slavonia, 20% in Q pe-
traea from the Spessart Mountains and 0% Northern provenances usually display a in Q roburfrom Haste in Lower Saxony. slower growth rate (Oppermann, 1932; Naidenova and Kostov, 1979). Good stem The general opinion is that crown form form is correlated with good growth over and stem form are better with acute branch limited areas (Oppermann, 1932). In Bul- angles than with broad crowns (Cieslar, garia, early flushing trees within prove- 1921; Hauch, 1916-1921), but timber quali- nances displayed better growth (Kostov, ty may be inferior with acute branches. 1983). Among German provenances, vol- Oak from The Netherlands had exception- ume production may differ by as much as ally good form in Denmark, but growth was 100% at age 35 years. Non-indigenous somewhat slower (Oppermann, 1932). provenances may be included in the com- Davidova (1970) reported that pheno- parison. Similar differences have been de- typic selection gave satisfactory results in scribed by Oppermann (1932) for Danish form among > 313 trees tested. provenance experiments. Growth potential shows more ecotypic than clinal variation. Growth Quite often, phenological characteristics seem to be correlated with growth charac- growth of oak is influenced by Juvenile teristics. weight. This may be one of the rea- acorn sons for the better early growth of Q robur Wood and bark (Burger, 1921; Krahl-Urban, 1959; Jova- novi&jadnr; and Tucovi&jadnr;, 1975). Influence of As early as 1893-1894 Hartig studied the seed weight only disappears after 12-15 variability of oak wood characters and the years (Cieslar, 1923). There is no clear effect of environmental influences on them. geographic trend in growth potential, but There are some minor differences between the climate of the location of origin is re- Q robur and Q petraea wood characteris- flected in the progenies (Cieslar, 1923). tics (Huber et al, 1941).Q robur has darker Variation in leaf size seems to be weakly and more compact heart- and sapwood as correlated with growth, with oceanic origins compared to Q petraea (Jovanovi&jadnr; and Tu- having smaller leaves (Cieslar, 1923). covid, 1975). To what degree these differ- ences are due to the variations in humidity Provenance variation can be extraordi- and nutritional levels of the respective sites nary. Krahl-Urban (1959) found > 100% remains questionable. Krahl-Urban (1959) differences in height growth at age 6 provided a good summary of the earlier years. These differences may continue at studies in oak wood characters and their later ages. Late flushing Q robur prove- variability. nances from Slavonia were not only better in stem straightness and in lack of epi- Most recent studies have been concen- corms but also 40% superior in volume trated on within-population variation. Polge growth compared with local German prove- (1984) detected considerable variability be- nances at age 69 years (Hesmer, 1958). tween trees in most wood characters stud- These provenances are not susceptible to ied and considered, like Lanier (1985), that attack by Tortrix viridana. In their natural genetic improvement was possible. Birot et range in Slavonia, considerable variation al (1980) found significant individual differ- exists in stem form over short distances. ences in spiral grain. Nepveu et al (1981) The percentage of forking is especially low studied infradensity, early wood percent- in Lipovljana and high in Otok and Krstovi age, percentage of vessels, fibers and rays in Q robur and Q petraea using clones and (Krahl-Urban, 1959).
Roots described high heritabilities for infradensity and earlywood percentage. In a study of Root systems have been studied for young Q robur grafted clonal material, Nepveu plants under comparable conditions. Bur- (1984a,b) reported high variability. Early ger (1921) and Krahl-Urban (1959) found wood percentage was under more strict more intensive root systems for Q petraea genetic control than vessel percentage. with a considerable variability in root per- Basic density showed high heritability and centage and structure within species. Jov- shrinkage low heritability. Savill (1986) anovi&jadnr; and Tucovi&jadnr; (1975) reported that studied shake in Q robur and Q petraea young plants of Q robur have a superior and described high variability. Early wood root system to that of Q petraea. Q robur vessel size was significantly correlated roots were found to be less sensitive to with shake. Nikolov et al (1981) detected waterlogging than other oak species (Col- significant correlations between wood den- in-Belgrand et al, 1991) and photosynthe- sity and flushing in Q robur. Early flushing sis is less influenced by waterlogging. trees with rough bark had the highest ba- Q robur roots can penetrate compacted sic density and widest rings with maximum soil and improve it (Oppermann, 1932). latewood percentage. Similar results were reported by Jevlev (1972a,b,c). The me- chanical properties of early flushing forms Seed and flowering of Q robur have 10-14% higher values than those of late flushing forms. On average, the seed of Q robur is bigger Bark structure of Q robur is coarser than that of Q petraea (Oelkers, 1913; Bur- than that of Q petraea (Klepac, 1957; ger, 1914, 1921),but there is a wide over- Krahl-Urban, 1959; Jovanovi&jadnr; and Tucov- lap between the species. The best charac- i&jadnr;, 1975). However, a high variability exists teristics for species differentiation are the on individual tree levels within populations. dark longitudinal strips on Q robur seed Significant influences of tree age and com- (Oelkers, 1913) and the relationship be- petition on bark structure were observed tween length and diameter of acorns, (Krahl-Urban, 1959). No correlation be- which is > 1.6 for Q robur (Burger, 1914). tween bark structure and wood character- Provenance differences in size are bigger istics could be detected by Schulz (1954). than species differences (Krahl-Urban, Jevlev (1972a,b,c) studied different bark 1959). The 1 000-seed weight ranges from types in the Voronezh reserve and de- 2 900 to 4 200 g for Q petraea and from scribed 6 different forms, 2 of which 3 000 to 5 450 g for Q robur on a prove- showed differences in wood characteristics. nance mean level. Seed size has an influ- ence on initial growth. Wood quality is negatively influenced by epicorms. Late flushing oak (Q robur tar- Seed shape inQ robur can be quite dif- dissima) forms fewer epicorms, harder and ferent even between neighboring trees durable wood and narrower sap- more (Kienitz, 1879). Southern sources were Within provenances, there also wood. found to have higher variability than north- seems to be considerable variability in the ern sources. Petrov (1975) described con- potential to form epicorms. siderable variability in all seed characters studied in 117 trees of the Alma Ata re- R Kleinschmit has selected trees free gion. Peduncle length was most variable; from epicorms for a seed orchard since length, diameter and form were the most 1960. This seed orchard is flowering and stable. But a high variability among acorns included in progeny testing.
also exists in terms of size and shape of seed at more frequent intervals (Litsharev, 1969). Flower variability was used for clas- the same tree (Oppermann, 1932; Kleinschmit, 1976), which is dependent sification into several types (Kravtsova, upon the time of ripening, the year and the 1968). Abnormal flowers occur with bisex- ual or female flowers on male catkins position. (Piatnitsky, 1954). Numbers of acorns are Q petraea seed germinates earlier. It low in relation to flowers formed with high germinate on the tree in autumn. Ger- can individual variabilities ranging from 0 to mination capacity rapidly decreases with 30%. length of storage (Szczotka, 1978; Tylkow- ski, 1982; Suszka and Tulkowski, 1983). The pollen grains of Q robur and Q pe- DISCUSSION traea have very similar exine and intine structures. The pollen grains have a regu- European oak species are not strict biolog- lar oval form with 3 longitudinal scars sym- species but hybridize naturally in the ical metrically arranged and are 30-45 μm in size (Jovanovi&jadnr; and Tucovi&jadnr;, 1975). Intra- contact zones with other species. This in- specific variation is higher than interspecif- natural variability considerably creases and makes it difficult to draw clear boun- ic variation. Pollen dimensions are greater daries. From a practical point of view, it is in Q petraea (Olsson, 1975b). Trees with abnormal leaf forms show pollen which nevertheless meaningful to maintain the may be of a hybrid nature. Rushton (1976) binomial system, as discussed by Burger compared pollen of Q robur, Q petraea (1975) for the North American oaks. Evolu- and suspected hybrid trees and described tionary differentiation parallels ecological a broad overlap in both species but with a differentiation and this is reflected in the higher variability in the putative hybrids. patterns of variation. This variation is a combination of clinal variation (eg for bud One male catkin contains > 550 000 pol- set) and ecotypic variation (eg flushing, len grains which can be transported 60-70 growth, form). Ecotypic variation prevails, km at elevations up to 3 000 m (Pianitzky, however. Adaptation to climate is not only 1954). The maximum pollen concentration reflected in phenological traits but also in is, however, within 100-200 m (Jovanovi&jadnr; survival. Southwestern provenances in and Tucovi&jadnr;, 1975). Since protandry is particular show high losses when planted common, self-fertilization is reduced. Pol- in northern or eastern countries and may len germination is better on the stigma of exhibit extreme growth depressions due to other genotypes than on that of the parent frost damage. These injuries in return are trees. reflected in the shape of the trees. There- Usually flowering starts at 50-70 years. fore phenotypic selection only makes sense Early and late flowering trees exist in popu- if the material is grown under similar climat- lations. We found fertile seed on trees as ic conditions. If environmental conditions young as age 7 years in plantations. Popu- are changed drastically, lack of pheno- lation differences in flowering are also de- logical adaptation has a series of negative scribed for Q robur var praecox Czern var consequences. However, broadly adapta- tardiflora Czern (Jovanovi&jadnr; and Tucovi&jadnr;, ble provenances exist which perform well 1975). even under different climatic conditions of is irregular with 3- to 10-year the plantation site. Q robur from certain lo- Flowering intervals between good seed years, de- cations in Slavonia seems to constitute pending upon climate. Some trees bear such provenances.
de l’angle du fil du bois chez quelques feuil- The natural pattern of variation in oaks lus : hêtre, chêne et Eucalyptus dalrymplea- superimposed by a man-made artificial is na. Ann Sci For 37, 19-36 pattern due to provenance transfer, planta- Cappelli M, Tauri L (1988) Research Bonani S, tion activities and subsequent hybridiza- interspecific variation in holm oak (Quer- on tion. This considerably complicates the de- cus ilex). Third preliminary note: growth in scription of oak variability and sometimes the nursery during the first two years. Atti Ist makes it impossible to trace what was nat- Ecol Selvicolt, Univ Studi Padova 5, 39-70 artificial. ural variation and what was Burger H (1921) Über morphologische und bio- For a practical approach, intensive test- logische Eigenschaften der Stiel- und ing and selection of populations and single Traubeneiche und ihre Erziehung im Forst- trees seems worthwhile due to the consid- garten. Mitt Schweiz Centralanstalt Forstl Versuchswesen 11, 306-376 erable variation available in growth and quality traits. The possibilities for genetic Einfluß der Herkunft des Sa- H (1949) Burger Eigenschaften der forstlichen auf die improvement of oaks have been discussed mens Holzgewächse. VII. Mitt Eiche Mitt Schweiz elsewhere (Gathy, 1969; Beuschel, 1975; Anst Forstl Versuchswesen 26, 59, 9-90 Kelinschmit et al, 1975a,b; Davidova, in Quer- Burger WC (1975) The species concept 1977; Kleinschmit and Svolba, Molotkov Taxon 24, 45-50 cus. and Davydova, 1979; Mol’chenko, 1982; (1938-1939) Monographie du genre Tishchenko, 1982; Spethman, 1986a; Mei- Camus A Quercus. Texte Paris 2, 195-378 er-Dinkel, 1987; Harmer, 1989). Cieslar A (1923) Untersuchungen über die wirts- Studies on susceptibility of European chaftliche Bedeutung der Herkunft des Saat- oaks to oak wilt disease and the relation- gutes der Stieleiche. Centralbl Ges Forstwes ship between growth and timber quality for 49, 97-149 veneer production are important topics for Colin-Belgrand M, Dreyer E, Biron P (1991) future research. Sensitivity of seedlings from different oak long rotations of oak trees make it The species to waterlogging: effects on root growth and mineral nutrition. Ann Sci For 48, particularly important to choose well- 193-204 adapted, fast-growing reproductive materi- al of high quality when plantations are es- Colombo PM, Lorenzoni FC, Grigoletto F (1983) Pollen grain morphology supports the taxon- tablished. omical discrimination of Mediterranean oaks (Quercus, Fagaceae). Plant Syst Evol 141, 273-284 REFERENCES Cousens JE (1965) The status of the peduncu- late and sessile oaks in Britain. Watsonia 6, Aas G (1988) Untersuchungen zur Trennung 161-176 und Kreuzbarkeit von Stiel- und Traube- neiche (Quercus robur L und Quercus pe- Danilov MD, Leukhina TA, Federov PN, Ku- traea (Matt) Liebl). Dissertation, Univ Munich dryavtseva MV, Petrova IN, Grigor’ev AJ (1972) The ecological and physiological char- Kreuzbarkeit und Unterscheidung Aas G (1990) acteristics of forms of Quercus robur differing Stiel- und Traubeneiche. Allg Forstz 45, von as regards time of leaf yellowing and fall. 219-221 Sb Tr Mariisk Politekhn Inst 59, 211-221 (1975) Untersuchungen über die Beuschel G Davidova NJ (1970) Ocenka pljusovih derevjev Vererbung und umweltbedingte Veränderung duba po semenomu potomstvu. Lesn Genet quantitativer und qualitativer Eigenschaften Selekc. Petrozavodsk 251-257 bei freiabgeblühten Traubeneichen. Disserta- tion, Univ München Davidova NJ (1977) Results of long-term trials of oak seed progeny. Lesovod Agrolesomeli- Birot Y, Dufour J, Ferrandes P, Teissier Du Cros E, Azouf P, Hoslin R (1980) Variabilité or 48, 10-15
A (1941) Bericht über Kreuzungsver- Hauch LA (1915) Proveniensforsög med I. Dengler Eg suche zwischen Trauben- und Stieleiche und Forstl Forsøgsvaes Dan 4 zwischen europäischer und japanischer Hauch LA (1916-1921) Proveniensforsög med Lärche. Mitt Akad Dtsch Forstwiss 1, 87-109 Eg II.Forstl Forsøgsvaes Dan 5 Densior LK (1970) O vosmoznosti selekciji duba Hauch LA (1925-1928) Proveniensforsög med na ustojcivost protiv obrazovani- cerescatovo Eg III. Forstl Forsøgsvaes Dan10 ja morozobaja. Lesn Genet Selekc Petroza- Henrik J (1973) Phenological forms of Quercus vodsk 236-242 robur. Sylwan 117, 67-69 Dissescu G, Coca C (1973) Variation in the num- Hesmer H (1958) Wald und Forstwirtschaft in ber of leaves in the genus Quercus, in relation Nordrhein-Westfalen. Hannover 1958 to species and tree age. Rev Padurilor 88, Huber B, Holzheide W, Raack K (1941) Zur 362-367 Frage der Unterscheidbarkeit des Holzes von Dreyer E, Colin-Belgrand M, Biron P (1991) Stiel- und Traubeneiche. Holz als Roh- und Photosynthesis and shoot water status of Werkstoff seedlings from different oak species submit- Jevlev VV (1972a) Phenological form and eco- ted to waterlogging. Ann Sci For 48, 205-214 types of Quercus robur in the Voronezh re- Dupouey JL (1983) Analyse multivariate de serve. Tr Voronezh Gos Zapov 18, 20-48 quelques caractères morphologiques de pop- Jevlev VV (1972b) Forms of Quercus robur dif- ulations de chênes (Quercus robur L et Quer- fering in bark. Tr Voronezh Gos Zapov 18, cus petraea (Matt) Liebl) du Hurepoix. Ann 54-64 Sci For 40, 265-282 Jevlev VV (1972c) Physical and mechanical Gardiner AS (1970) Pedunculate and sessile properties of the wood of early and late forms oak (Quercus robur L and Quercus petraea of Quercus robur. Tr Voronezh Gos Zapov (Mattuchka) Liebl). Forestry 43, 151-160 18, 129-132 Garilov T,Stojkov H (1978) Morphologische und Johnsson H (1952) Ungdomsutvecklingen hos ökologische Formen bei der ungarischen stjälek, druvek och ròdek. Sven Skogsvärds- Eiche (Q conferta Kit) im Strandsha Gebirge. foren Tidskr 2, 168-193 Gorskostop Nauka 15, 12-28 Jovanovi&jadnr; M, Tucovi&jadnr; A (1975) Genetics of com- Gathy P (1969) Contribution à l’étude de la gé- mon and sessile oak (Quercus robur L and nétique des chênes. 2nd World Consul For Q petraea Liebl). Ann For Zagreb 7, 23-48 Tree Breeding. Washington 1969. Proc FAO, Kienitz M (1879) Über Formen und Abarten hei- 2, 979-987 mischer Waldbäume. Forstl Z 241-160/297- Globa-Mikhailenko DA (1973) The development 327 of forms in the genus Quercus. Byull GI Bot P (1977) Les poils des quatre espèces Kissling Sada 88, 35-38 de chênes du Jura (Quercus pubescens, P (1987) Contribution à la Grandjean G, Sigaud Q petraea, Q robur et Q cerris). Ber Schweiz taxonomie et à l’écologie des chênes du Bot Ges 87, 18p Berry. Ann Sci For 44, 35-65 Kleinschmit J (1976) Untersuchungen über Fall- Harmer R (1989) Selection of superior oak. Res- und Eichelgewichte bei Stiel- und zeitpunkt earch information note. For Commun UK Traubeneiche. Mitt Hessischen Landesforst- 149, 1-2 verwalt 14, 52-63 Hartig R (1893) Untersuchungen über den Kleinschmit J, Svolba J (1979) Möglichkeiten Wachstumsgang und Ertrag der Eichen- der züchterischen Verbesserung von Stiel- bestande des Spessarts. Forsti Naturwiss Z und Traubeneiche (Q robur u Quercus pe- 2, 249-269 traea). III. Nachkommenschaftsprünfung von Hartig R (1894) Untersuchungen über die Ent- Eichenzuchtbäumen. Allg Forst Jagdztg 150, stehung und die Eigenschaften des Eichen- 111-120 holzes. Forstl Naturwiss Z 5, 193-203 Otto H, Sauer A (1975a) Kleinschmit J, Hauch LA (1909) Erblichkeit bei Buche und der züchterischen Verbesse- Möglichkeiten Eiche. Centralbl Ges Forstwes 35, 333-348 Stiel- und Traubeneichen (Quercus von rung
Menitsky JL (1971) Dubi kavkaza. Izv Nauka petraea). I. Inventur der robur and Quercus ANSSSR (Leningrad) 196 Eichensamenplantagen. Allg Forst Jagdztg 146, 157-166 Mitchell A (1974) Die Wald- und Parkbaüme Eu- ropas. Paul Parey Verlag, Hamburg, 233-254 Kleinschmit J, Otto H, Sauer A (1975b) Ver- suche zur Stecklingsvermehrung der Eiche. Mol’chenko LL (1982) Selection of plus trees by Allg Forst Jagdztg 146, 179-186 an integrated set of characters. Lesn Khozy 10, 33-36 D (1957) Research into Bark Thickness Klepac of Quercus robur and Quercus petraea. Su- Davydova NI (1979) The use of Molotkov PI, marski Inst Zagreb internal report properties as an indicator of differ- bioelectric ent growth and selection classes in oak Kollmann F (1955) Technologie des Holzes und trees. Lesovod Agrolesome Lior 55, 24-29 der Holzwuchs-stoffe, 2. Aufl Springer Ver- lag, Berlin Naidenova TS, Kostov KD (1979) Rate of photo- synthesis and transpiration of Quercus robur Koloszar J (1987) Die slawonische Eiche in Un- in relation to seed origin. Gorskostop Nauka garn. Forst Holzwirt 11, 293-296 16, 3-10 Kostov KD (1983) Survival and variation in Neger FT, Münch E (1950) Die Laubhölzer. height and diameter growth in some prove- Sammlung Göschen de Gruyter, Berlin Bd nances of Quercus robur. Gorskostop Nauka 718, 160 20, 4-14 p Déterminisme génotypique Nepveu G (1984a) Krahl-Urban J (1953) Rassenfragen bei Eichen de la structure anatomique du bois chez und Buchen. Allg Forstz 8, 477-480 Quercus robur. Silvae Genet 33, 91-95 Krahl-Urban J (1957) Über Eichen-Provenienz- héréditaire de la Nepveu G (1984b) Contrôle versuche. Silvae Genet 6, 15-31 densité et de la rétractibilité du bois de trois (1959) Die Eichen. Paul Parey Krahl-Urban J espèces de chêne (Quercus petraea, Quer- Verlag, Hamburg, 288 cus robur et Quercus rubra). Silvae Genet Kravtsova NV (1968) Morfologija pesticnik cvet- 33, 110-115 kov raznik form duba. Apomiks i citoembriol Lemoine M (1981) Héréd- Nepveu G, Garbaye I, rastenii Saratov, 131-135 de la forme et de la qualité ité génotypique G (1978) Handbuch der Laub- Krüssmann du bois de chêne. Reprise, développement et gehölze. Paul Parey Verlag, Berlin, 79-115 port en plantation de copies végétatives de chênes sélectionnés pour la qualité de leur Lanier L (1985) Production de bois de haute qualité technologique en futaie régulière. bois. Ann Sci For 38, 531-532 Ann Gembloux 91, 163-172 (1990) Jahresbericht der Nied- NFV ersächsischen Forstlichen Versuchsanstalt. (1969) Zakonomernnosti formirova- Litsharev IN Abt Forstpflanzenzüchtung, internal annual nija urozajev duba. Probl Gorn Lesovod Sev 16 Kaukaze Krasnodar 150-166 report Nikolov S, Denev D, Bl’Skova G, Dragozov I Lüdemann G (1962) Die Forstpflanzenanzucht (1981) Structure and properties of the wood in Kämpen und Forstbaumschulen Nord- of some forms of the Vardim oak. Gorskostop deutschlands. Dissertation, Univ Göttingen Nauka 18, 28-36 Masarovicova E, Pozgaj J (1988) Comparative Stiel- und Traubeneiche, eine Oelkers J analysis of the leaf area in three oak species (1913) variationsstatistische Untersuchung. Z Forst a methodical contribution. Biologia 43, - Jagdwes 45, 18-45 449-457 (1975a) A morphological analysis of Olsson U (1963) Animal Species and Evolution. E Mayr phenotypes in populations of Quercus (Faga- Belknap Press/Harvard Univ Press, The ceae) in Sweden. Bot Not 128, 55-68 Cambridge, MA Olsson U (1975b) On the size and microstruc- Meier-Dinkel A (1987) Propagation of Prunus ture of pollen grains of Quercus robur and and Quercus by tissue culture. In: Proc Eur Q petraea (Fagaceae). Bot Not 128, 256- Semin Wood Prod Harvesting. Bologna 2, 265 76-83
Shirnin VK, En’Kova EJ (1975) Quality of the A (1932) Egens Traeformer og Oppermann rac- wood of phenological varieties of Quercus ro- Forstl Forsøgsvaes Dan. XII, 400 p er. bur in the conditions of the fresh oak forest Patlai IN, Boiko AV (1977) Some features of type. Genet Sel Semenorodstvo i Introdukt- shoot formation in oaks of different prove- siya les Porod 2, 169-174 nances. Lesovod Agrolesomelior 48, 79-83 (1951) Transpiration of early and late Silina AA Petrov SA (1975) Variation in features of the re- flushing races of oak in the forest-steppe productive organs of Quercus robur in SE Tr Inst Lesa 41, 104-110 zone. Kazakhstan. Lesn Zh 2, 162-164 Smit A (1973) A scanning electron microscopi- SS (1939) der Eichen. Pitanitsky Hybridisierung cal study of the pollen morphology in the ge- Lesn Khoz 7, 1 nus Quercus. Acta Bot Neerl 22, 655-665 Pitanitsky SS (1954) Selkcija Duba. Golesbu- W (1986) Stecklingsvermehrung Spethmann mizdat, Moscow-Leningrad, 147 Stiel- und Traubeneiche. Schriften aus von Polge H (1984) Production de chênes de qualité der Forstl. Fakultät der Univ Göttingen und en France. Rev For Fr (No special), 34-48 der Nieders. Forstl Versuchsanstalt 86, 99 Rachwal L (1982) Provenance experiments with W (1986) Artbestimmung von zuge- Spethmann various species in Niepolomice forest. Arbor lassenen Eichen-Beständen durch Untersu- Kornickie 27, 367-389 chung von Einzelblättern und Fruchtständen. Nieders Forstl Versuchsanstalt, internal re- Rack K (1957) Untersuchungen der Anfälligkeit port, 7 p Manuscript 17 verschiedener Eichenprovenienzen ge- genüber dem Eichenmehltau. Allg Forst Staszkiewicz J (1970) Quercus robur L; Quer- cus sessilis Erb. In: Variability of Leaves and Jagdztg 128, 150-156 Fruits of Trees and Shrubs in the Forest As- Rushton BS (1976) Pollen grain size in Quercus sociations of the Bialowieza National Park. robur L and Quercus petraea (Matt) Liebl. Monogr Bot 32, 222 Watsonia 11, 137-140 Suszka B, Tylkowski T (1980) Storage of acorns Rushton BS (1978) Quercus robur L and Quer- of the English oak (Quercus robur L) over 1- cuds petraea (Matt) Liebl: a multivariate ap- 5 winters. Arbor Kornickie 25, 199-229 proach to the hybrid problem. 1. Data acqui- of respiration in the Szczotka Z (1978) Intensity sition, analysis and interpretation. Watsonia of Quercus borealis Michx and embryo axes 12, 81-101 Quercus robur L acorns during storage and Rushton BS (1983) An analysis of variation of ageing under controlled conditions. Arbor leaf characters in Quercus robur L and Quer- Kornickie 23, 145-151 cus petraea (Matt) Liebl population samples JV (1971) The historical significance of Thirgood from Northern Ireland. Ir For 40, 52-77 oak. In: Oak Symposium Proceedings. North- Savill PS (1986) Anatomical characters in the eastern For Exp Sta For Serv USDA, Upper wood of oak (Quercus robur L and Q petraea Darby, PA, 1-18 Liebl) which predispose trees to shake. Com- A of wisdom. Hortic (1986) Pillars Thompson monw For Rev 65, 109-116 Week 9, 20-21 Schulz H (1954) Untersuchungen über die Be- Tishchenko VYA (1982) The fruiting of Quercus wertung von Eichen-stammholz. Dissertation robur in a clonal seed orchard. Lesn Khoz Hann-Münden, Univ Göttingen 11, 30-32 Schwarz O (1936-1939) Monographie der T (1982) Height increment of 1-year Tylkowski Eichen Mitteleuropas und des Mittelmeerge- shoots of the English oak (Quercus robur L) bietes. Feddes Rep (Sonderbeih D) 5, 72- and the northern red oak (Quercus borealis 173 Michx Q rubra L) from 4-year-old roots of = Seitz W (1923) Fragt die Eichen, wie sie wach- seedlings raised from acorns stored over 1-5 sen! Z Forst winters. Arbor Kornickie 27, 357-365 Jagdwes 25, 321-331 (1974) Evaluation of stabilizing Semerikov LF Verchenko VG (1975) Using phenological varie- populations of oak. Ekologiya 5, ties of Quercus robur for ravine and quelley selection in stands. Lesn Zh 3, 7-10 5-10
Weiser F (1964) Probleme der Auswahl und Er- Yacine A, Lumaret R (1988) Distribution spatiale gebnisse der Erkundung autochthoner des génotypes dans une population de Herkünfte der Stiel- chêne vert (Quercus ilex L), flux génétique et und Traubeneiche regime de reproduction. Genet Sel Evol 20, (Quercus robur L und Quercus petraea (Matt) Liebl) in der Volksrepublik Rumänien 181-198 für die Anlage von Provenienzversuchen. Yacine A, Lumaret R (1989) Genetic diversity in Arch Forstwes 13, 843-864 Holm oak (Quercus ilex L): insight from sev- Willkomm M (1875, 1887) Forstliche Flora von, eral enzyme markers. Silvae Genet 36, Deutschland und Österreich. Winter, Leipzig 140-148