LEIDEN BOTANICAL No. SERIES, Variation in 3, root D.F. Jodrell Summary Variability L. and Acer generalised in the in statements of root anatomy in L. the Cutler wood of Kew reference the literature on wood anatomy Botanic Gardens, Kew, Great Britain Laboratory, Royal pseudoplatanus 1976 143-156, pp. root wood selected specimens particularly microscope slide collection excelsior Fraxinus is discussed in relation to anatomy. Introduction Although variability some detail economic root root in the to is so that structure regulate little relatively because root more difficult important know about the to necessary has been studied in work has been carried out comparable wood is it is often slight. However, it is therefore anatomy root the anatomical characters of trunk wood probably importance samples; in help by wood. This is root on in various authors, and its obtain, to be to able to identify range of variability possible identifications can be made. Use is made of variability accurate of certain fruit the final size of the trees for the selection of of the tree, crown stocks root and the age which will which at fruiting begins. In the the a above-ground position parts from which the of sample number of respects from trunk in twigs, for example, and may have scalariform describe twig (or Samples from near the in stem) even twig and junction detailed and sometimes with of The arrangement of even in vessels, adjacent Crown of the Botanic wood or with the reason, near the base of of extensive or within a study a tree for each and (1950) family. frequently compared to show samples leaning trunks experiment. and parenchyma single ring smaller usually features of reaction wood from subject is dissimilar in Metcalfe & Chalk anatomy separately fibres, tracheids, Gardens, is often related to twigs in vessels of the trunk anatomical differences when growth rings copyright; reproduced Director, Royal simple perforation plates branches, trunk. The numbers of cells of each * length juvenile anatomy wood of and diameter of cells is vessels. For this special and from branches have been the extent, the mature gross taken from the main trunk. The in wood variability is taken. The wood; species plates young tree, a from can vary opposite to some sides of a type per unit area, the size and thickness of their permission of the controller of Her Majesty's Stationary Office, and Kew. 143 D. F. CUTLER cell walls based at a on also can standard taken, only in growth rings a however, has only a be to a should anatomy samples or There may be even fall in a is found, distance, applied to counts or length occur may so hardly change of measurements or a often shorter are say tracheid in, maximum may be in very old trees. Generally, of elements. From the first age a steep that the axial elements of or of successive lengths of cells the life of the tree, relating length and increase gradual length made, continuous increase a xylem outwards there is normally short be taken and distribution frequency have been measurements between the occur rings throughout new normally from different specimens have been obtained. rings. recent curve formed secondary levels off after show number of numbers of large figures maintained, Sanio a of wood descriptions a trunk. The first formed (oldest) axially arranged cells more in successive reached and on detectable differences than those in length, when that reliable easily Less made from the trunk. The sizes of cells, their position should be stated so vary. General observations at increase in which subsequent growth rings all. Usually cell dimensions length a statistical analysis trends before such be can appreciated. addition In between the to It is inappropriate will find that the Bary (1884); Bosshard 1952); (1951); White & Novruzova Root wood It seems Spurr of there can be considerable differences obtained from samples composition of the many & & list of references Faull points trees Hyvarinen Liese & (1954); (1968); Jane varies in much the same is frequently more & here, outlined above: (1934); Fegel (1941); (1966); Philipson Mariani (1968); anatomy that cover Robards of the same but the reader Sanio (1872); Bannan (1941-2, Dadswell de 1944, Din- (1959); Butterfield(1968); Yaltirik(1968, (1970); Gottwald Baas (1971); (1973); (1974). variability of additional factors ence single tree, a comprehensive a (1930); Baily der Graaff & Baas tree. give to following Chalk woodie (1963); 1970); within xylem anatomy different environmental conditions. grown under species van variability secondary such of soil atmosphere as and soil so way as wood from the aerial parts of a extreme. This may be due to the pres- compression, waterlogging, variability on, not in the in the environment of the aerial present parts. Although aerial parts, scattered much less as cover 57 wood that these basis for in annual 144 or anatomy were usually varieties. He in higher generally are as usually the a subject own to not In content less; (d) root anatomy amount often takes the form of short able and of considerable than that of of information notes in papers observations and his literature review make a with trunk wood follows. and fibre on there is Fayle's was comparison and hardwoods is rings subjects. only generalisations discussion, content is area with other species root published the literature. This throughout dealing largely has been Fayle (1968) points out, number of anatomy. generalisations He was careful laws. The statements do roots: (a) pith lower; (c) heartwood and is form about to a state useful absent; (b) parenchyma the number of vessels tyloses are per unit infrequent; (e) contain fewer cells and the boundaries between rings are the less VARIATION well defined; (f) cells in the less are on lignified; pits the root and bibliography also relate papers Rusch observations and on an extended list to the Suss & (1973); approach to Roots removed damage are are the Muller-Stoll sent the to with compared number of different 190 species 4,500-5,000 well need as to Very of ROOTS a at range roots the so The from the wood slides. It is possible the same not be related that specimen trunk wood slides for retained with tree to one of two species slides using plant. most a these or up largely commonly in order The roots in both the reference and particular part displayed to of a root enquiries to one MATERIALS AND Tree and shrub roots and the normal way with stained in collections of known origin were of is, then, represented, in response cause to about to the damage and be better re- related are not to can root and trunk wood select representative not are same not possible variability species. normally to referable relate variability root. METHODS sectioned safranin, are compare the range of system. Consequently it is particular types (e.g. lateral, tap) are studies very extensive collection of trunk but collections, even a of many of the there anatomy tend root There is as which of total. The slides increments of trunk wood with variability in root wood anatomy from the to a in comparative growth which of each roots identification; following or roots closely as particular structural some about 900 are subsidence from the determining differences between compare the comparison, following anatomy is very variable. The root specimens there prepared way of of material available for same to The buildings showing main drawback is that the root samples trunk wood samples from here. collection, and identified those which exhibit the widest variability in their presented than others. gave con- comprehensive a essentially practical (Cutler, 1974). much wider roots. The collection has been built identify samples, pitting AT KEW Kew is this work number of also are narrow walls, and some Patel(1965); Pil'shchikov(1969); foundations of might trees from Fayle for identification. Sections represented; enquiries of these. The considerable. as varieties from resulting of species have thinner increased with unnecessary only relatively inexpensive collection contains or OF anatomy laboratory quite evident It is damage. reference root proximity the ANATOMY (1973). slides from the references This is often possible. of study is Macdonald (1960); subject: from the rows discussion. of references THE STUDY The number of WOOD these and the observations of other authors will be tangential walls. Some of sidered in the section ROOT generally wider, longer, are larger and the are IN on a sledge safranin and microtome at haematoxylin, 25-30jum, or safranin 145 D. F. CUTLER and fast was green. Sections dehydrated and mounted were not known from which part of the lateral and were Acer not and Fraxinus excelsior pseudoplatanus selected for ring system the samples were Normally, but obtained, it most tap-roots. demonstrate a wide range of and Fraxinus root in canada balsam. variability in porous in the trunk wood. and comparison, a selected for were since special study they diffuse anatomy, and the Acer is porous root slides of trunk wood Representative simple statistical analysis applied were that numbers of so vessel elements per unit area, and the radial diameter of vessel elements could be discussed for each sample element diameter in diameter was likely tudinal elements in in In ured. root. In Fraxinus B and in D, and In Acer the with very was distinct cells and fifth were worked The following applied: trunk size out were growth rings elements in root p = 0.05 and p examined were Rosa sp., three roots, wood, Populus one sp., roots some a examined were pattern of cell were meas- could be detected, elements of each group per thirty were sampled in roots sampled. were examined. The early growth rings were made on zones where of ten unit areas areas recorded was for could be taken to avoid each overlap root. were for calculated for each using set of measurements or Student's t test, and levels of confidence 0.01. = and are discussed, Quercus (robur type), but statistical no three roots, Malus analysis pumila, was and root root. all rather narrow authors. Mature root wood had regular longi- examined. each OBSERVATIONS AND The were D, where only five accounted was at of vessels categories thus avoided and observations and standard deviations Sample counts. diameterfor growth rings, sixth, seventh, and eighth growth rings regular for the root being except in Acer mean size measurements of because of the small diameter of the The vessel tangential C which had fewer rings, the fifth, sixth, and seventh number of vessel sample, two the roots third, fourth, the anatomy The where root narrow than tangential ten from each of three consecutive analysed separately, with was Radial rather manner. wood, when compared with trunk wood. root Fraxinus, meaningful to be greater in the radial than the Thirty vessel elements, each a selected for measurement, since Fayle (1968) stated that increase was size and DISCUSSION when probably frequency compared not with those reported been laid down in any of of cell types was apparent in on them, the by but areas analysed. The number of isations showed to to be some root made. samples The abnormality, demonstratethat some critical re-examination. 146 examined for each particular or roots reported represented part of the specimen of a on is rather low for were selected range of variability. The published generalisations about root general- because they object anatomy may was need VARIATION Plata C shows 1A, B, and These demonstrate part the statistical set B root in out vessels for the roots as be an ring porous in Fig. 1, which shows in mature In the trunk mature two striking ring wood and zones the the zone' 'pore overlap. excelsior. 1. —Fraxinus area for mature trunk wood p = late Plate In the 1A and the numbers are joined by of vessels per than in the stem. unit or p line = B and B and D mature of vessel 0.01 other tissues, reconcile the be the fact that in the data by plus length, p no and D). cross-hatched. relate = = diameter samples (B, C, they (p there is element radial to 0.01; is in- set out density into two distinct size wood, although in the such distinction could be made. all vessels wood; solid black Total line and the late some of Fraxinus excelsior. clearly grouped no one are narrower generalisation which three samples D was with diffuse porous wood. This the vessel elements to early wood, to indicate that area. to species, could be and three root wood and a 'pore zone' stated that if the species wood from roots roots Comparison (A) 0.05, cross-hatched, wood root between radial vessel diameter and vessel root C, vessels root feature is that in Fig. and by diffuse porous species the in the of the tissues. This is substantiated vessels of the range of the smaller with porous porous!) comparison a wood and representing root the section (T.S.) (first formed vessels) comparable with not and trunk wood. It would appear interpretation the dimensions. He diffuse (and not zone' most growth ring) complicated attempt unnecessarily than those of the classes the supplementary tissue, extra, accurate, The of cell late wood of compare the of ANATOMY diffuse timbers should be treated separately porous and porous ring comparative study to transverse represented 'pore whereas in stem normally larger (throughout considered that regarded vessels of the , He seems in which root C is 1 Fig. stated that in Fraxinus (1965) are should of.Fraxinus excelsior in WOOD of variability which exists, and they complement range smaller in the roots than in the are ROOT Plate 1C. by Patel data three roots of the IN one 0.01) significant difference and number of vessels in per unit Blocks indicate vessel diameter: In A and another. The centre fall within the size portion of C, the blocks for separate line, p = early lines indicate 0.05. 147 D. F. CUTLER size. In larger the smaller vessels C, root vessels trunk wood mature from that of the mature root compared are B, Plate 2B, and statistical unit in Fig. 2, C, root significantly four variable and Plate Plate 2D analysis, those of the late wood and the vessels of the larger roots of area but wood, Mature trunk wood and doplatanus to spring wood of the specimen. number of vessels per The similar in size are smaller than the significantly are roots a more (at p visual significantly not 0.01) = are of the diffuse porous photographs (T.S.) 2C, together give C is B and with of the trunk of the impression root D. species Acerpseuwood, additional root, an different present in not in variability Plate 2A, used in the wood root anatomy. Root B The root in Plate 2D Referring is not (Plate 2C) collections. Root C erence the average condition of Acer (Plate 2B) represents to Fig. be can seen E root are larger, different. Vessel sample C (p number in = 0.01) In each of the above are Fig. area p = significantly 2. Acer 148 per when trunk wood and building representative it = unit area. Total line can at sample seen of vessel a roots p = 0.01; mature = soil. B and D trunk 0.05, signif- are significantly are not significantly root root C radial portion mature = 0.05) sample Blocks The of as it is trunk wood. of trunk wood. In diameter and number of vessels samples (B-E). centre E exist in which vessel elements representative solid black and cross-hatched. length, roots of significantly greater (p that element wood lakeside. 0.01. with the number in the be at a heavy clay on a the confidence level p in trunk wood and from and four root 0.01 p = B and D is than those (A) p at compared examples, narrower 0.05, cross-hatched, of vessels the and taken area samples pseudoplatanus. Comparison for mature a than those of the trunk wood. The elements in The numbers of vessels per unit from the Kew ref- that vessel element radial diameterin smaller than those of the trunk wood in roots waterlogged ground from under foundations of different from that in significantly wood. The vessels in icantly larger came 2 it was taken from separate line, p per unit indicate vessel diameter: with = lines indicate the number 0.05. VARIATION Fraxinus Acer trunk wood (mature the diameter of approach (mature roots ring porous) vessels in roots as it wider than in are a trend towards and others sample. Fayle, Patel, WOOD may be exceptions wider being suggest that, generally, trunk wood. The evidence mature that the number of ANATOMY In three of the four sample. trunk wood diffuse porous) vessels show is, shows ROOT has vessel elements which roots vessels of the trunk wood spring than in the mature trunk wood limited of the none IN presented here, high enough to warrant further research into the matter. From in area hardwoods is be cast mentioned, As in have Acer water shows C with rings Fayle's are more be a to the was not of more variability again some taken into account, doubt site. The waterlogged roots presented by Pohl(1926, growing in soil. The of presence is the water were not roots growing most or com- actually in Fayle, in water elements in Acer narrow in all as be extensive. to 1927, reported the caprea, but, growth ring significance then, has been shown slide difficult are of the spaces of In a with number of samples trunk of roots rings some are many of the incomplete. growth rings. on photographs in Plate 2 shows that Acer number of the generalisations vessels per unit in difficulty detect in several of the Acer roots; this is in accordance to generalisation comparison a distinguishable are collection. area made than in the trunk wood wood. In Acer root C than in the trunk wood. These roots root B (Plate 2B) by Fayle, except (Plate 2A). (Plate 2C) number of the cells have thicker walls than in the trunk narrower and situation, of trunk wood with average of Alnus and Salix related growth rings than those of the a Evidence trees some species Probably from came but in soil. well with responds unit trunk wood. In four of the mature be the per particles. in the A visual to samples vessel elements than those In Fraxinus Growth select to frequency. 3C root might, then, between soil samples shown themselves where roots that for narrower root than in roots not trunk wood for either taken was between the growing 1968) mature care noted, in this is that the number of vessels indicates generalisation. cell size and vessel parison less usually here, also Fayle the value of the results of this is limited since the full present work range in variability width, the on Obviously, of evidence roots examined seven can collected Most cells and the root in cor- that there are larger Plate 2D and all cell types appear wood, do not follow the generalised concept in these respects. Plate 3A, are and C shows has been analysis was B, included here because sent to the from a from roots. root, since it anatomy in has or a specimen and vessel wall uniseriate shown The they attempted. Laboratory parts serve The as a of as a sucker shoot states Quercus among other rays points, but in T.S. These no quantitative in Plate 3C is of interest because this shown). Fayle identified pitting Quercus (robur type) illustrate several structure pith (not was to roots root, but could be wide, multiseriate; Plate 3C is of three things. on that in is Quercus of intermediate widths abnormal in this respect when pith generally the basis of details Rays compared sample arising adventitiously rarely with robur occur. absent of are cortex either The the trunk root wood, 149 D. F. but CUTLER many oak roots have abnormal rays apparently also are of ment, typical The since abnormal, they all are Q. reference collections. Even in the from the very The vessel elements (Riedl, 1937). do show dendriticarrange- slight a robur. shown in Plate 3A and 3B roots They narrow. small are mature vessels of the first more root like the majority (3A) growth rings shown in growth ring of oak are roots in the indistinct. Apart Plate 3B, there is little distinction between the diameters of vessels from early and late wood, and ring porousness is lost. This accords excavations made are walls. It is essential likely be the to and eter observations. Fayle's to close to be able to often encountered when since distinguish them from the roots of trees which of structural cause are buildings, roses commonly planted are near to are more damage. shown in Plate 4C and D demonstrate differences in vessel element diam- roots The with of Rosa also have variable roots. Roots of Rosa Species number of vessels per unit of fibres and Growth parenchyma. area. rings Note also in cell wall thickness variability the indistinct, are roots fitting well with gener- alisations in this respect. Malus sylvestris selected in Plate in which on an less are for variability in the root, and which at they in purposes of individual were more vessels per unit the root most found anatomy, and 1962; Shimaji, A to have can roots both 1962; heterogeneous trees. Unfortunately Populus rays of which occurrence sible to give rise The roots are of are separate to contain fewer and the cells, (1969) has reported are in and Salix are usually Patel, Plate the area and finer features which rays are homocellular in the associated with to Salix with the basis of is to exclusively root situation in 1961, Populus ID. ascribed either numbers of heterocellular rays in on many which This 1965). commonly readily or also may be heterocellular (Lebedenko, is shown in Populus, the two genera to difficulties in making identifications. One of in which rays homocellular, larger fewer The vessels are, showed variations related tendency for species in reported ray Whereas the trunk wood is example growing. apparent of these is the the trunk wood species. area. this point, the larger differences of vessel size and number per unit to in rings horticulture. Pil'shchikov the arrangement of cells have been discussed. There vary but the Mention has already been made of the selection of Malus distinct. roots anatomy, shows many of the features to differ from trunk wood. There generally supposed dwarfing variation between Up 4A) some average, wider than those of the stem, annual stocks depth root parenchyma more boundaries root in contrasted with trunk wood (Plate wood is root fibres and Mill, also shows 4B, to the buildings. majority of heterocellular rays, the Populus wood damage Populus, roots makes it impos- anatomy alone. CONCLUSIONS From the extensive collections of reference slides of small 150 sample selected serves to indicate that the root probable anatomy held extent of at Kew, the variability in root VARIATION Plate 1. —A-C, Fraxirus excelsior variability (x65); D, Populus sp., transverse radial sections longitudinal of roots, section IN to illustrate showing a ROOT part WOOD of the heterocellular range ray ANATOMY of anatomical (x 130). 151\1 D. F. CUTLER Plate 2. wood 152\2 Acer — of a pseudoplatams. A, range of specimens for transverse section comparison with of mature trunk A (all X 65). wood; B-D, transverse section of root VARIATION Plate 3. —Quercus (robur type). A-C, anatomy (all x transverse section of roots IN ROOT of different WOOD specimens for ANATOMY comparison of 80). 153\3 D. F. Plate B, CUTLER 4. —A, B, Malus transverse section the 154\4 range of sylvestris. of mature root variability (all x 65). A, transverse wood; C, D, section of roots of Rosa mature trunk sp. wood, for in transverse section comparison with demonstrating part B; of VARIATION wood is greater than in and may WOOD made between trunk are ANATOMY wood anatomy, and root wood, need revision. Additional research is required because of the economic root ROOT normally appreciated. 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