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Influence of crown exposure on the morphological needle traits of nine conifers
Utjecaj ekspozicije krošnje na morfološka svojstva iglica devet četinjača
Biljana M. Nikolić, Katarina Mladenović, Ljubinko Rakonjac, Slobodan Milanović, Marija M. Marković, Srdjan Bojović, Nevena Čule
Summary
The aim of this research was to investigate if the crown exposure of some conifers influenced to needle properties. The leaf morphological traits of sixty-six trees of nine conifers: Atlas cedar, Austrian pine, Blue spruce, Douglas fir, European spruce, European yew, Serbian spruce, Silver fir, and White fir, from six Belgrade parks, were analyzed. Five needles were measured from each of the four main crown exposures. Length, width, area and perimeter of needles were investigated. Species, parks in which they were found, as well as crown exposures, differed mostly in needle length and needle width. Correlations between measured needle traits were determined by linear regression analysis. Strong positive correlations were found between the length, perimeter, and area of needles. The differences among the species in terms of light requirement determine species for individual planting as light-loving or partial shade species (Atlas cedar, European spruce, Serbian spruce, Blue spruce, Austrian pine and Douglas fir), or for group planting as shade-loving species (Silver fir, White fir and European yew).
Key words: conifers, correlations, exposure, needle morphology, parks.
INTRODUCTION
UVOD
The influence of light exposure on the tree crown, among other ecological factors is very important for the successful development of a species. In parks, where crowns are almost open, it is relevant whether a species is light or shade-loving. Other habitat conditions such as habitat exposition, geological substratum, soil, assemblage, air pollution, etc. also influence tree growth (e.g. Hällgren and Fredriksson 1982; Donovan et al. 2005; Freer-Smith et al. 2005).
Silver fir (Abies alba Mill.) is a tall, European high-mountain species, growing on silicate and limestone. As it tolerates dry air, arid soil, and polluted air of urban city very poorly, it is rarely used in parks. White fir (Abies concolor /Gordon/ Lind. ex Hildebr.) is a tall, high-mountain species though sometimes it can be found below 1000 MSL. It

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inhabits the Western USA. It is resistant to frost, wind and drought, as well as dust and harmful gases in the air and has modest soil requirements. Atlas cedar (Cedrus atlantica /Endl./ Mann. ex Carrière) is a tall, high-mountain fast-growing conifer native to Algeria and Morocco. It is resistant to climate extremes and can grow near the sea, as well as in mountainous regions at low elevations. It is an alkalophilic and heliophilic species. European spruce (Picea abies /L./ Karst.) is a tall, European mountain species that requires plenty of humidity both in the air and the soil. It grows on silicate substrate and acidic soil, though some plant communities also grow on serpentinite or limestone substrate. It has poor tolerance for polluted urban area and industrial air. Serbian spruce (Picea omorika /Panč./ Purkyně) is a tall, high-mountain species, tertiary relict and endemite of the Balkan Peninsula (Bosnia and Herzegovina and Serbia, Europe). Grows mostly on limestone, as well as silicate and swampy soil. Because of its resistance to city conditions, it is very common in parks. Blue spruce (Picea pungens Engelm.) is a tall, North American high-mountain species, growing on humid podzolic, acidic and carbonated soils. It tolerates severe frost, as well as dry air and summer droughts, and urban area conditions. Austrian pine (Pinus nigra J. F. Arnold) is a medium-height mountain species spreading from east Spain to Asia Minor and Crimea. It is a pioneer species, heliophilic, widely used for the afforestation of arid stony areas. It grows on steep limestone, dolomite and serpentinite cliffs. It is resistant to drought, and wind, it tolerates urban area conditions. Douglas fir (Pseudotsuga menziesii /Mirb./ Franco), is a tall species, native to the Pacific zone of North America and requires enough relative air humidity and acidic soil. It is widely cultivated in Serbian parks and forest cultures. European yew (Taxus baccata L.) is slow-growing but long-living species, often shrubby and naturally distributed from North and South Europe to the Mediterranean, Algeria, Morocco, Asia and the Caucasus. It can grow in shade, as well as in open positions, on shallow and poor or deep and rich soils. It is widely cultivated in parks (Vukićević 1982).
Up to now, examinations of the interacting effects of drought and light intensity (e.g. Holmgren, 2000; Aranda et al. 2005; Dutilleul et al. 2015) were performed almost exclusively on seedling growth. Gebauer et al. (2019) examined the impact of drought stress on the growth of one-year needles of Picea abies and ascertained that drought stress was correlated with tree assemblage and needle morphological traits. They also discovered that the differences in needle shape cross-section correlated to light intensity i.e. that the cross-section of needles exposed to light was quadrangular, while for the ones in the shade was ellipsoid.
To date, research has focused on the influence of crown light exposure on tree growth i.e. tree height and diameter (Wyckoff and Clarck 2005; Osada 2012; etc.), as well as tree mortality (Shenkin et al. 2018). The correlation of air polution with morphoanatomical traits of the needles of conifers living in urban areas had been previously examined (Nikolić et al. 2019 and refs. cited therein).
The aim of present paper is to examine the influence of crown exposure on the needle morphology of nine conifer species from different genera found in the parks of Belgrade, Serbia. To our knowledge, there has been no similar research conducted in Serbia with adult trees published to date.
MATERIALS AND METHODS
MATERIJALI I METODE
Geographic and geologic characteristics of the study parks and their position in town were presented in Table 1 and Figure 1, respectively). Sixty-six trees of nine conifer species: Abies alba (3 trees), Abies concolor (5 trees), Cedrus atlantica (14 trees), Picea abies (3 trees), P. omorika (7 rees), P. pungens (6 trees), Pinus nigra (10 trees), Pseudotsuga menziesii (6 trees), and Taxus baccata (12 trees) (Table 2) were analyzed. One to four trees of every species per park were analyzed, as was indicated in Table 2. One-year old needles were collected from the tips of lower third of the crown of solitary trees. Five needles (ca. 20 needles per tree) were analyzed from each of the four crown exposures (E, N, S, W). Four morphological needle properties (length, width, area, and perimeter) were measured using SigmaScan Pro

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5.0 image analysis software (Systat Software, Inc., San Jose, CA, USA).
Mean values and differences between the species’ needle morphology were examined using the Cluster Analysis (Nearest Neighbor Method, Squared Euclidean Distance, Figure 2), while differences between parks and exposures were examined by ANOVA and LSD test at 95% level, using Statgraphics Centurion XVI, Version 16.1.11., (USA) statistical program (Figures 3 and 4; Table 2; Table 3). Correlations between the four measured needle traits of nine conifer species were determined by linear regression analyses (presented as text in Results section).
RESULTS
REZULTATI
Geographic and geologic characteristics of the study parks were presented in Table 1. For TP park slight air pollution was found. AC and BB parks had moderate, but MM, PS and PP parks excessive air pollution (according to distance from trafic). Geologic substrata also differ between parks, but pedologic substrata were almost same (limestones), with the exception of BB park (serpentinite).
Mean values of four morphological needle traits (length, width, area and perimeter) influenced by the crown exposure of all nine investigated species were presented in Tables 2,3 and 4 and Figures 2,3 and 4. The mean values of all quoted morphological traits were almost published before (Nikolić et al. 2019) (all histograms which were marked as “Mean” in Figures 3 and 4, and in Table 2).
Pinus nigra possessed the highest needle trait values (Table 2, Figure 2). Abies concolor, Taxus baccata and Abies alba also had larger needles in comparison to other investigated conifers.
Strong positive linear correlations were found between length and perimeter (r = 0.94 – 1.00), length and area (r = 0.82 – 0.96), and area and perimeter (r = 0.84 – 0.97) (results were not presented). Predominantly weak positive correlations were found between width and area (r = 0.10 – 0.80). Negative correlations between width and length and width and perimeter (r = -0.37 and r = -0.29, resp.) were found for C. atlantica. There were strong positive correlations (r > 0.75) between almost all four traits of P. pungens.
Means for each analyzed park were presented in Table 2. Mean values for four main exposures (E, N, S, W) had been presented in Table 2 too, as well as for every of analyzed park (Figure 3 a-i and Figure 4 a-i). Statistical signification was presented in Table 3.
In Abies alba needles, the mean values of the four measured traits (length, width, area and perimeter) were presented (Table 2, Figure 3a, Figure 4a), with statistically approved differences between parks BB and PP for needle length

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(Table 3a). Northern crown exposure was the best for needle length, too (Table 3a). In A. concolor, needle mean values for BB and PP were nearly identical. Northern exposure had almost the best needle length and width (Figure 3b, Figure 4b), but it was not statistically approved in both properties (only in needle width,Table 3b). In Cedrus atlantica needles, great differences in the analyzed traits between parks were found only in needle width (Figure 3c, Figure 4c, Table 3c), where parks AK and BB had statistically the best values. At the same time, we can not conclude that some exposures gave the best results (Table 3c). In Picea abies needles (Figure 3d, Figure 4d), needle length and width were significantly higher in BB than in PP (Table 3d). The best results were found on northern and western exposure (needle length and width, resp.). In P. omorika needles the highest values were statistically approved in park PS (Figure 3e, Figure 4e, Table 3e). The longest needles were found on eastern exposure, but it was not statistically significant. For P. pungens needles (Figure 3f, Figure 4f, Table 3f), the highest and statistically approved length and width were found in park PS, while eastern exposure was the best (Table 3f). In Pinus nigra needles (Figure 3g, h;

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Figure 4g, h), the highest needle length was found in MM and PS (Table 3g). Southern exposure was found to be also statistically the best. In needles of Pseudotsuga menziesii, the highest values of length and width were exibited in PP (Table 2, Figure 3h, Figure 4h, Table 3h). The largest needles (statistically approved) according to needle length and width were found on eastern and northern exposures, resp. In needle length of T. baccata, park BB was the best (Table 2, Figure 3i, Figure 4i, Table 3i), while in needle width PP was the best. Western and northern exposures were significantly the best, resp.
DISCUSSION
RASPRAVA
Abies alba, according to Jovanović (1967) and Janković (1973), is a sciophilic species (thrives in the shade), which agrees with presented results where the maximum values of needle length on northern exposure was shown, even though, according to Robakowski et al. (2004), young seedlings require somewhat more light. A. concolor is also a shade-loving species, however, it copes well with sunny locations as well and is a desirable park species (Jovanović 1967; Vukićević 1982). Presented results, since no statistical differences in crown exposure were found, confirmed statements of both authors. Furthermore, Mori and Takeda (2004) reported that branches of alpine species, A. mariesi and A. veitchii, developed more slowly in the shade. Needle masses per area of these species (results of current-year needles!) were lower in the shade. Cedrus atlantica is favorite park species which in the examinations presented therein exhibited variability concerning exposure. Its both heliophilic (Vukićević 1982) and xerotherm characters (Jovanović 1967), were not statistically approved in presented results, although on southern exposure needle length was the highest. Picea abies is a species that copes better in the shade (Jovanović 1967) or in partial shade (Janković 1973). The both opinions were approved in presented examination where higher needle length was found on northern, and higher needle width on western exposure. P. omorika, though a sciophilic species, it can thrive in the light as well (Jovanović 1967). In presented results its needle length had the highest mean value on the eastern side, but its needle width was the best on the northern side. It is important to notice that both findings didn’t have statistical support. P. pungens is also considered a heliophilic species (Jovanović 1967), which coincides with the maximum values found mostly on eastern exposure (presented results, statistically approved). Furthermore, needle masses per area (of current-year needles) of P. jezoensis (Mori and Takeda, 2004) was lower in the shade. Pinus nigra is a distinct heliophilous (Jovanović 1967; Janković 1973), as confirmed in presented results where maximum values were found on southern exposure, but it was not statistically confirmed. Pseudotsuga menziesii is considered a partial shade species and is successfully grown in forest cultures (Vukićević 1982). It was in accordance with presented evidence where the longest needles were found on southern exposure, but the thickest needles were found on northern exposure. Taxus baccata is a species that tolerates deep shade (Jovanović 1967; Janković 1973; Robakowski et al. 2004), and in presented results it achieved the best results on eastern or northern exposures (needle length and needle width, resp.).
The significant impact between light and shade on leaf mass per area had also been established in some tropical species (Martin et al. 2020) and other leaf physiological traits. It was founded that differences in needle morphology between parks could be consequence of tree maturation, too (quoted in the case of Douglas-fir needles, where length, width, thickness, and roundness of needles grew through the needle age, Apple et al. 2002).
Increased dryness and the poverty of soil led to decreased length and area of needles (Tyukavina et al. 2019a). Needle area of Scots pine forests in taiga was reduced in conditions far from optimal water regime, so consequences in changing the width and thickness of needles occured (Tyukavina et al. 2019b).
In previous study (Nikolić et al. 2019), for all investigated species significant differences in needle morphology (and anatomy) between species as well as between individuals were found, too. Our results is in accordance to general data which was well known (Vukićević et al. 1982).

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BB is the only park on serpentinite ground that has most likely favoured the development of the needles of species A. alba, P. abies and T. baccata (primarily needle length). The remaining six studied species demonstrated better results on limestone surfaces. According to Vukićević (1982), P. pungens is not too demanding in terms of soil characteristics but we couldn’t approve this since all investigated trees of this species were on same substrat, loam. Speaking of soil properties, this paper’s findings and conclusions should be taken with reservation because some of the parks implemented cultivation measures, primarily fertilization (but, unfortunatelly, it was known for sure only for parks AC and BB).

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CONCLUSIONS
ZAKLJUČCI
According to obtained results the crown exposure influenced to needle properties in many of analyzed conifers. Species, parks in which they were found, as well as crown exposures, differed mostly in needle length and needle width.
The differences among the species in terms of light requirement found in the present research determined species for individual cooperation in parks (i.e. as solitary trees) as

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light-loving or partial shade species (C. atlantica, P. abies, P. omorika, P. pungens, P. nigra and P. menziesii), or group cooperation as shade-loving species (A. alba, A. concolor and T. baccata), which is something to be taken into consideration when setting up parks in the future.
ACKNOWLEDGEMENTS
ZAHVALE
This study was supported by the Ministry of Science, Technological Development and Innovation of the Republic of Serbia, grant numbers: 451-03-47/2023-01/200027 and 451-03-47/2023-01/200169.
REFERENCES
LITERATURA
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Donovan, R.G., H.E. Stewart, S.M. Owen, A.R. MacKenzie, C.N. Hewitt, 2005: Development and application of an urban tree air quality score for photochemical pollution episodes using the Birmingham, United Kingdom, area as a case study. Environ Sci Technol, 39(17):6730-6738.
Dutilleul, P., L. Han, F. Valladares, C. Messier, 2015: Crown traits of coniferous trees and their relation to shade tolerance can differ with leaf type: a biophysical demonstration using computed tomography scanning data. Front Plant Sci, 6:172.
Freer-Smith, P. H., K. P. Beckett, G. Taylor 2005: Deposition velocities to Sorbus aria, Acer campestre, ‘Beaupre’, Pinus nigra and x Cupressocyparis leylandii for coarse, fine and ultra-fine particles in the urban environment. Environ Pollut, 133(1):157-167.
Gebauer, R., D. Volařík, J. Urban, I. Børja, N.E. Nagy, T.D. Eldhuset, P. Krokene, 2019: Effects of mild drought on the morphology of sun and shade needles in 20- year-old Norway spruce trees. IForest, 12(1):27-34.
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Janković, M., 1973: Enciklopedijski leksikon mozaik znanja – Biologija, Interpres, Tom 19, Beograd.
Jovanović, B., 1967: Dendrologija sa osnovama fitocenologije. Naučna knjiga, 576 str., Beograd.
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Mori A., H. Takeda, 2004: Functional relationships between crown morphology and within-crown characteristics of understory saplings of three codominant conifers in a subalpine forest in central Japan. Tree Physiol, 24(6):661-670.
Nikolić, M.B., A.M. Stefanović, M.M. Veselinović, D.S. Milanović, D.K. Mladenović, Ž.S. Mitrović, M.S. Eremija, B.Lj Rakonjac, 2019: Needle morpho-anatomy and pollen morpho-physiology of selected conifers in urban conditions. Appl Ecol Env Res, 17(2):2831-2848.
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Sažetak
Analizirano je 66 stabala devet vrsta četinjača: atlaskog cedra, crnog bora, bodljikave smreke, duglazije, obične smreke, šumske tise, Pančićeve omorike, koloradske jele i obične jele, iz šest beogradskih parkova. Analizirano je pet iglica sa svake od četiri glavne ekspozicije krošnje. Ispitivana je duljina, širina, površina i opseg iglica. Vrste, parkovi u kojima su pronađene, kao i ekspozicije njihovih kruna, razlikovali su se po duljini i širini iglica. Korelacije između izmjerenih svojstava iglica određene su linearnom regresijskom analizom. Utvrđene su jake pozitivne korelacije između duljine, opsega i površine iglica. Razlike među vrstama u zahtjevima za svjetlom određuju vrste za pojedinačnu sadnju kao vrste koje vole svjetlo ili polusjenu (atlaski cedar, obična smreka, Pančićeva omorika, bodljikava smreka, crni bor i duglazija) ili za grupnu sadnju kao sjenoljubne vrste (obična jela, dugoigličava jela i šumska tisa).
Ključne riječi: četinjače, korelacije, izloženost, morfologija iglica, parkovi