Urban tree avenues - morphological characteristics as a factor in tree selection

M. Dobrilovič: Urban tree avenues - morphological characteristics as
a factor in trees selection

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URBAN TREE AVENUES – MORPHOLOGICAL
CHARACTERISTICS AS A FACTOR IN TREES SELECTION
URBANI DRVOREDI – ODABIR DRVOREDNIH VRSTA NA
OSNOVI MORFOLOŠKIH MJERA
M. Dobrilovič
ABSTRACT

The knowledge about plant material and its applicability is an important
factor in achieving quality in landscape design. It manifests itself in the
appropriate selection of plant species for individual design solutions.

Nowadays we can observe the selection of plants based on fashion;
measures defined by creation of a certain space characteristic are omitted. The
main problem is that in the literature the plant species are frequently treated
individually and arranged according to the visual effect of each plant. In order
to satisfy the needs of design practice for this research, it was necessary to
ascertain the hidden designer potential of each plant, which is not identical with
the pleasing effect of the plant. This is how the central problem is formulated in
this research: to determine the selection criteria (of plant species), arising from
the visual characteristics of the plants and, based on these criteria to determine
the suitability of plants to create visual effects of a tree avenue. In this sense we
can stress the importance of proving the connection between morphological
properties of the plants and the characteristics of tree avenues, which are formed
by these very plants. The subject of this research is plant material, 193 trees,
systematized according to their size, shape, habitus, texture, colour, as well as
the seasonal appearance of individual characteristics. We established criteria for
the selection of morphological properties of the trees that allowed us to achieve
a certain visual character of a tree avenue. The process of plants selection is
shown on a concrete example, whereby certain design demands determine the
choice of adequate plants. For the final selection of plants it is necessary to add
criteria arising from eco-physiological needs of the plants and from
technological demands.
Key words: landscape architecture, tree avenues, urban places, tree
morphology, selecting species

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SAŽETAK

Odabir biljnih vrsta često se temelji na ukusu i modnom dobu, više nego na
skupu karakteristika odabranih s ciljem stvaranja određene prostorne važnosti.
U literaturi biljne vrste se često obrađuju individualno i raspoređuju prema
vidljivim učincima pojedinih biljaka. Kako bi se zadovoljile potrebe
projektanata u praksi, bilo je potrebno istraživanje biljnih vrsta s ciljem
identifikacije skrivenog potencijala biljnih vrsta za design. U tom smislu ljepota
pojedinačne biljke podređena je prostornim potencijalom biljne vrste. Osnovni
razlog iztraživanja bio je zato: utvrditi kriterije za odabir biljnih vrsta, na osnovi
vidljivih obilježja biljaka i na osnovi toga odrediti prikladnosti vrsta za
stvaranje vizualnih učinaka drvoreda u širem urbanom prostoru. U tom
kontekstu važno je pokazati dosljednost između morfoloških karakteristika
biljaka i vidljivog značenja drvoreda kao krajobraznog elementa. Predmet
istraživanja je biljni materijal: 193 različite vrste stablašica raspoređene su
prema veličini, obliku, habitusu, teksturi, boji i razdoblju pojavljivanja
pojedinih svojstava. Predstavljene su mjere za odabir morfoloških karakteristika
biljnih vrsta s kojima možemo postići točno odabrani prostorni učinak drvoreda.
Proces odabira biljaka prikazan je na određenom slučaju. Na bazi određenih
zahtjeva dizajna odabrane su odgovarajuće drvoredne vrste (stablašice). Za
konačni odabir biljnih vrsta potrebno je dodati dodatne daljne kriterije na bazi
ekofizioloških potreba biljaka i tehnoloških kriterija.
Ključne riječi: oblikovanje krajobraza, urbani prostor, drvored, morfologija
drveća, odabir biljnih vrsta

INTRODUCTION

The knowledge about plant material and its applicability is an important
factor in achieving quality in landscape design. It manifests itself in the
appropriate selection of plant species for individual design solutions.

In nature the distribution and expansion of plants is determined by biotic
factors (soil, climate and fauna) whereas in a designed landscape people
represent the main factor. Here the plants become material, and people choose,
arrange and remodel according to their needs and wishes.
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In the past – and it is often so even nowadays - the selection of plants
usually rested upon individual taste and fashion dictates of the period, rather
than starting from and following directives determined by the objective of
creating a certain space characteristic.

The primary elements or rather ‘building blocks’ of a designed landscape
are called landscape elements. They can be vegetation (lawn area, tree avenue)
or non-vegetation (water and sand areas, buildings) (Ogrin, 1996). Examining
the relevant literature (Zion, 1970, Robinson, 1992, UCONN Plant Data Base of
trees, shrubs and vines, 2005, Wyman, 1956) it was ascertained that the research
on plant suitability as regards the construction of vegetational area phenoena
had been fairly fragmentary. Merely a few basic elements have been treated,
such as hedges and ground-cover plants. The procedures of selection are non-
transparent and quite inadequate for the use in a serious design.

Therefore the main research problem can be seen as determining criteria for
the selection of plant material, arising from the visual properties of the plants
and, based on these criteria, subsequently determining the suitability of plants as
to their capacity to create visual effects of vegetation areas. Rather than dealing
with all the vegetation elements this paper focuses on vegetation areas. It is in
this light that the objective is set - examining the morphological properties of
plant species with the aim to establish a means of selection of plant species
suitable for the construction of vegetation areas.

THE TREE AVENUES – THE LINE VEGETATION ELEMENT

The line vegetation elements behave in a cultivated region as zones of
riverine vegetation, boundaries and are a consequence of differences in the use
of space. In a formed region belong to the line vegetation structures tree
avenues, cut palisades, free-grown and cut hedges, plantings in the form of
zones etc.

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Figure 1, 2. Linear structure of vegetation (avenue,  clipped avenue, palisade)
(Park Citroen, Pariz, Jardin Atlantique, Pariz)
Slika 1, 2. Linijske vegetacijske strukture (drvored, šišan drvored i linijski
potezi (Park Citroen, Pariz, Jardin Atlantique, Pariz)

If we examine in detail the procedure of the formation of tree avenues we
realize, that it is distinguished from natural line structures by a regular
arrangement of trees in a fixed direction. As one of the earliest acts of the
regional formation the procedure is evident from illustrations of the first tree
avenues in Egypt, when the line of regularly arranged trees was created by a
larger number of tree species. The procedure of the regular arrangement was
followed by a reduction of species that mostly led to the use of one single tree
species (Ogrin,1993). The development of the formation procedure is evident
from the illustrations of the tree avenues in the Garden of high officials of the
sovereign Amenhotep III. in Thebes around 1400 B.C.

Tree avenues determine the direction and the frame of space, define the
areas, divide the surface elements, and direct the views. As spatial and view
guidance they condition the orientation and the feeling of safety in space, even
more, they are the carriers of messages (such as, the lines of tree avenues in the
baroque gardens). In the history of the regional formation they first appeared in
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Figure 3.  The garden of High Official at Thebes, 1400 b.c. (Cliford, 1962: 33)
Slika 3. Vrt u Tebi, 1400 p.n.e. (Cliford, 1962: 33)

the Egyptian culture, where water canals were bordered by trees. As regional
elements tree avenues have been preserved until today. Besides space forming
and the symbolic role their ecological role is also important (creating
microclimate conditions, the passage of animals...), that is why they are an
important element for the construction of the green system of cities. Among all
the other line structures they are most appropriate for creating rhythm and
sequence in space.

Deriving from the Butina definition of the line (Butina, 2000: 44) as a series
of oriented points we can come to a conclusion that the suitability of vegetation
species for the creation of a tree avenue derives from the suitability of the
species for the creation of the vegetation point. If a certain vegetal species is
suitable for the creation of the point elements, then it can also create the line
structure – the tree avenue. It has to be pointed out, that plants are not abstract
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points, as in dense arrangements the reciprocal effects can be produced, that can
be seen in the strong shading and in the destruction of certain plants. That is
why besides the appearance of the plant it is important to take into consideration
its ecological limitations or needs.

The characteristics of species, used for the construction of tree avenues are:
• Good levelling within the species (established genotype),
• Do not spread with root offshoots and other vegetative  reproductive modes,
• The regularity of the shape of the treetop, bush (is not as important
characteristic as the selection of the plants appropriate for vegetation point,
because of the line characteristics the identification of rhythm and repetition
is more important as the identification of the individual unit... The line can
also be created with the trees that have irregular habitus. In this case we
create a compact and levelled line structure by dense planting.),
• homogeneity, levelling of the core of the vegetation volume (this
characteristi is especially important when selecting the species for a tree
avenue),
• clear division of fenophases of the development of the plant (flowers are
equally arranged on the plant, the plant flowers at the same time on all parts
and deflowers quickly),
• tolerate well different intensity of lighting (an important characteristic when
selecting species for the free-grown and cut hedges, palisades),
• tolerate well the trimming (an important characteristic when selecting
species for cut hedges and palisades),
• tolerate well the trimming (an important characteristic for creating
palisades),
• tolerate well city climate, salt in the soil, dust, hard ground, temperature
oscillation, drought (for public parks in the cities and roadside plantings).

In a wider space the tree avenue makes a defined spatial structure, and at the
same time forms a proper (sub) space, that is very important in terms of
experience. By a negative space we understand the space between the ground
and treetops (the interior of the tree avenue). The form of this space depends on
the tree species (size, shape, and habitus), density and the way of planting
(Walker, 1991). Further on only some forms of the negative space of tree
avenues are shown.
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• the space profile in the rectangular shape

Treetops are dense, the rhythm of trunks is well expressed, and the visual
field is wide. The tree species that are appropriate for such tree avenues have
tall, levelled and smooth trunks and ball-shaped, wide ball-shaped, dome-
shaped and compact treetops. Appropriate habitus are irregular, upright and
wide spread.

• the space profile in the shape of letter 'T'
Treetops are dense, the rhythm of trunks is well expressed, and the visual
field is wide. The tree species that are appropriate for such tree avenues have
tall, levelled and smooth trunks and egg-shaped treetops. Habitus is regular,
upright and spread or irregular, upright and spread.

• the space profile in the shape of letter 'T' with the emphasised central side
Treetops are not dense; the rhythm of trunks is not expressed well. The
visual field is limited to the ending of the tree avenue. The tree species that
are appropriate for such tree avenues have short trunks, ball-shaped, dome-
shaped or cone-shaped treetops, regular and upright or irregular, upright and
widely spread habitus.
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• the space profile in a rectangular shape with emphasised width
Treetops are dense, the rhythm of trunks is expressed, and the visual field is
limited by low treetops. The tree species that are appropriate for such tree
avenues have lower trunks, long framing branches, the shape of the treetop
stands out, bow-shaped, habitus is irregular, upright and sloping
(descending).

• the space profile in the rectangular shape with the emphasised height
Treetops are not dense, the rhythm of trunks is not expressed very well, and
the visual field is limited to the real ending of the tree avenue. The tree
species that are appropriate for such tree avenues have low trunks, short
branches and offshoots that are equally arranged on the extension of the
trunk. The treetop is slightly pointed (the cone-shape of the treetop) or flat
(cylinder-shape of the treetop). Habitus is regular or upright.

MATERIAL AND WORK METHOD

The subjects of this research project are plant species designed for greening
public spaces.  They are chosen on the basis of assessing their hardiness, their
resistance to urban climate, to diseases and pests, the cost of maintenance and
their design potential.
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The research encompasses autochthonous species (Kotar and Brus, 1999),
suitable for the use in public places and foreign species or sorts of trees which
are wide-spread in our country and fulfil the user’s demands (Šiftar, 2001,
Bruns Pflanzen,1997).

The principal goal of the research is to define appropriate criteria for plant
species selection in order to achieve the desired effect of the tree avenue. In
order to achieve this it was first necessary to establish the connection between
the morphological and the visual properties of the plants in the tree avenue
(height, form, texture, colour and similar) and to determine which
morphological properties of the plants affected the desired character of the
avenue.

As far as the work method is concerned it is imperative to emphasize that
the aspect here treated as far as the knowledge of plants is concerned, covers a
fairly unknown ground and does not provide well-established methods of
research. Therefore further on the work process is introduced in some detail. It
was necessary to systematize all plant species (the research subject matter)
according to their visual properties (size, shape, habitus, texture, and colour)
and the properties determined by their seasonal appearance (e.g. foliage colour
– autumn, colour of blossoms – spring …). All plants were classified on the
basis of data obtained by observing visual properties in situ, and subsequently
comparing them with findings of other authors.  This was followed by a
meticulous analysis in terms of visual properties (size, shape, colour, texture,
durability of foliage). Next step was the setting of criteria for the selection of
morphological plants by means of which we could achieve the desired visible
characteristic feature of the tree avenue. In order to help establish the afore-
mentioned connections, a scheme is demonstrated in which the grey colour
designates those morphological plant properties by which the character of tree
avenues is defined.
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Figure 4. Morphological characteristics of plants used for tree avenues
Slika 4. Morfološke karakteristike stablašica prikladnih za izgradnju drvoreda

The next step is the selection of plant species according to the desired visual
effects of the avenue. The selection process is introduced by means of a
concrete example and represents one of the results of this research. The
suitability of plant species for the construction of tree avenues is determined
according to the following criteria: size, foliage durability, shape, habitus, and
colour (leaf, blossom, shoot). The criteria are introduced gradually, each added
criteria representing a new level or rather combination of morphological
properties. We consider the most suitable those species that conform most
optimally to all levels of selection and all combinations of criteria.

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RESULTS AND DISCUSSION

Results of this research are presented in two condensed paragraphs:
1. Systematizing the plant species according to their morphological properties;
this comprehensive part of the results is not shown in the article. It is,
however, of great importance for the process of selection itself. By the same
token it represents an autonomously applicable outcome, since the elaborate
complex classification enables us to look for adequate plants with regard to
their visual properties (size, shape, habitus, texture, and colour) and seasonal
appearance.
2. The process of plant selection itself in terms of the effect that a plant species
is capable of expressing when appearing as a constituent part of a certain
vegetational element.

The selection of plant species in terms of desired visual effects of tree
avenues will now be demonstrated on a concrete example. Prior to the selection
of certain plant species a distinct design goal (or respectively a pre-formulated
design request) for a certain visible effect of the landscape element is presented.
Thus, for example, by making a certain choice we wish to find the best possible
plant species for the construction of a tree avenue whose visual effects are
defined by its height, form, texture, colour and the time when a certain effect
will be visible.

Example:  tree avenue

The expected properties (design requirements):  tree avenue height 5-10 m,
deciduous, treetops are dense, the interior of the tree avenue (negative space) in
the tube-shape, dense arc, the texture of leaves medium, colour green, uncut
treetops.

Stipulating morphological properties of adequate plant species:
• size:
tree(5-10 m )
• foliage durability:   deciduous
• shape:
rounded to oval, high domed, closed crown
• habitus:
informal, upright, widely spreading, hanging slightly
• texture - leaf:
medium
• colour - leaf:
green
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Definition of criteria for selection and a list of species that conform to these
criteria:
1.Size - foliage durability
Acer platanoides
Acer negundo
Acer platanoides 'Cleveland'
Abies nordmanniana
Acer platanoides 'Faassen's Black'
Acer platanoides 'Schwedleri'
Acer pseudoplatanus
Acer saccharinum
Acer rubrum
Ailanthus altissima
Acer saccharinum 'Pyramidale'
Betula ermanii
Aesculus hippocastanum
Carpinus betulus
Alnus cordata
Castanea sativa
Betula papyrifera
Fagus sylvatica
Betula pendula 'Fastigiata'
Acer platanoides 'Royal Red'
Carpinus betulus 'Fastigiata'
Fagus sylvatica 'Atropunicea'
Corylus colurna
Fagus sylvatica 'Dawyck Gold'
Fagus sylvatica 'Asplenifolia'
Fagus sylvatica 'Pendula'
Fagus sylvatica 'Dawyck'
Fraxinus excelsior 'Altena'
Fagus sylvatica 'Dawyck Purple'
Gleditsia triacanthos 'Shademaster'
Fraxinus excelsior
Liriodendron tulipifera
Gleditsia triacanthos 'Skyline'
Malus tschonoskii
Juglans regia
Paulownia tomentosa
Malus baccata
Populus balsamifera
Malus triloba
Populus alba f. pyramidalis
Populus alba
Populus x berolinensis
Populus alba 'Raket'
Prunus avium
Populus nigra 'Italica'
Quercus frainetto
Populus simonii
Quercus petrea 'Columna'
Pterocarya fraxinifolia
Quercus robur 'Fastigiata'
Quercus palustris
Robinia pseudoacacia 'Bessoniana'
Quercus robur
Robinia pseudoacacia 'Unifoliola'
Quercus rubra
Salix alba 'Tristis'
Robinia pseudoacacia 'Pyramidalis'
Tila x euchlora
Salix alba
Tilia cordata 'Erecta'
Sophora japonica
Tilia tomentosa
Tilia cordata

Tilia platyphyllos

Tilia x intermedia 'Pallida'

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2. Size - foliage durability  - shape

3.   Size - foliage durability  - habitus – shape (crown)
Acer negundo
Acer platanoides 'Faassen's Black'
Acer pseudoplatanus
Acer saccharinum
Acer saccharum
Aesculus hippocastanum
Ailanthus atissima
Carpinus betulus
Fagus sylvatica
Fagus sylvatica 'Asplenifolia'
Fagus sylvatica 'Atropunicea'
Fraxinus excelsior
Juglans regia
Malus baccata
Pterocarya fraxinifolia
Quercus robur
Quercus rubra
Salix alba
Sophora japonica
Tila x euchlora
Tilia cordata
Tilia tomentosa

4.  Size - foliage durability  - habitus – shape (crown and trunk)
Acer negundo
Acer saccharum
Carpinus betulus
Fagus sylvatica
Fagus sylvatica 'Asplenifolia'
Fagus sylvatica 'Atropunicea'
Juglans regia
Malus baccata
Quercus robur
 Sophora japonica

Acer negundo
Acer platanoides 'Emerald Queen'
Acer platanoides 'Faassen's Black'
Acer platanoides 'Royal Red'
Acer pseudoplatanus
Acer saccharinum
Acer saccharum
Aesculus hippocastanum
Ailanthus altissima
Carpinus betulus
Fagus sylvatica
Fagus sylvatica 'Asplenifolia'
Fagus sylvatica 'Atropunicea'
Fraxinus excelsior
Juglans regia
Malus baccata
Pterocarya fraxinifolia
Quercus robur
Quercus rubra
Salix alba
Salix alba 'Tristis'
Sophora japonica
Tila x euchlora
Tilia cordata

Tilia tomentosa
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5.  Size - foliage durability - habitus - shape (crown and trunk) – texture (leaves)
Acer negundo
Acer saccharum
Carpinus betulus
Fagus sylvatica
Fagus sylvatica 'Asplenifolia'
Fagus sylvatica 'Atropunicea'
Malus baccata
Quercus robur

6. Size - foliage durability  - habitus- shape (crown and trunk)- texture (leaves) -
color (leaves)

The dense, medium height tree avenue with the tube-shaped interior, is best
completed by the species Acer saccharum and Carpinus betulus. Both species
have low trunks, dense and compact treetops and tolerate well dense planting
the condition in the tree avenue. They tolerate well the salt in the soil, which is
an important characteristic of tree species. Branches on the exterior side of the
tree avenue with older plants are slightly curved and create a visual protection.
We get a similar effect if we use a beech tree (Fagus sylvatica, Fagus sylvatica
'Asplenifolia'), but because of the overshadowing of the first framing branch
they may develop higher and that is why the effect of 'tubes' is smaller. In
communal planting oak develops (Quercus robur) a flat, tall trunk and a small
treetop (Erker, 1957: 82), which eliminates it from the species appropriate for
creating the required spatial effect. Due to the eco- physiological unsuitability
we can also eliminate Acer negundo (it does not tolerate well the
overshadowing).
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CONCLUSION

The method used and described here for the selection of plant species
represents a novelty in the field of landscape design. By means of this research
it was ascertained that the manner of selecting plants according to their visual
properties is appropriate. For a final selection it is necessary to add to the
morphological criteria some other criteria, which are formulated on the basis of
eco-physiological needs of each plant (lighting, temperature, soil conditions)
and technological demands (cultivation requirements).

Therefore it is imperative to emphasize the relativity of such plant selection,
as evident from the above commentary, relating to a concrete example of
choosing appropriate plants. The results of the research are applicable on
different levels. By using the data gained in the research (systemizing the plant
species according to their visual properties) we can find species that belong to
certain size classes, shapes, colours, and textures. The data can be combined
arbitrarily in order to search for cross-section multitudes that in essence
represent combinations of morphological properties. The data and the above
mentioned systematizations are not presented in this paper, due to the
extensiveness of the material.

Alongside the selection criteria it is also important in which order or rather
sequence the criteria, upon which the selection of plants is based, are applied.
Which criteria are to be included and in what order, largely depends on the
design goal. The paper demonstrates the process of selection of plant species
suitable for tree avenues. In a similar way suitable plants for other landscape
elements can be chosen.

The presented process of selection is not applicable only in landscape
architecture. It can be equally useful in selective breeding, since it can help
determine the need for a particular shape, habitus, colour … In terms of its
informative quality it can be applied by tree planters, horticultural experts and
may also be seen as a welcome aid in the education of landscape architects and
horticultural experts.
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LITERATURE
Bruns Pflanzen: Sortimentskatalog 97-98. 1997. Bad  Zwischenahn, J. Bruns:
704 s.
Butina M. 2000. Mala likovna teorija. DEBORA, Ljubljana, 173 s.
Clifford D.1962. A History of garden design. London, Faber and Faber: 232 str.
Erker R. 1957. Opis gozdnega drevja in grmovja. Ljubljana, Državna založba
Slovenije: 210 str.
Kotar M., Brus R. 1999.Naše drevesne vrste. Ljubljana, Slovenska matica v
Ljubljani: 320 s.
Ogrin D. 1993. Vrtna umetnost sveta. Ljubljana, Pudon, EWO: 400 s.
Ogrin D. 1996. Teorija krajinskega oblikovanja: sinopsis predavanj. Ljubljana,
Biotehniška fakulteta (gradivo razdeljeno na predavanjih)
Robinson N. 1992. The Planting Design Handbook. Cambridge, The University
         Press: 271 str. Slovenije, 290 s.
Šiftar A. 2001. Izbor in uporaba drevnine za javne nasade. Ljubljana, Zavod za
         tehnično izobraževanje: 193 s.
Šiftar A., Kravanja, N. 1974. Vrtno drevje in grmovnice. DZS, Ljubljana, 290 s.
UCONN Plant Data Base of trees, shrubs and vines. The Univerity of
          Connecticut. Connecticut: 1-3-2005.
Walker T.D.1991. Planting design, second edition. Van Nostrand Reinhold, New
          York, 196 s.
Wyman D. 1956. Ground Cover Plants.The Macmillan Company, New York, 175s.
Zion R.L. 1970.Trees for architecture and the landscape. Reinhold Book
          Coprporation, NY, Amsterdam, London, 284 s.

Adresa autora - author´s address:

Primljeno – Received
Marko Dobrilovič, Ass. Ph.D.B.Sc

23. April, 2009
Univ. of Ljubljana,
Biotehnical Faculty,
Landscape Architecture Department,
Jamnikarjeva 101,
Ljubljana, SI-1111


Analiza rodoslovlja dinarske populacije risa (Lynx lynx)

Sindičić, Magda;Gomerčić, Tomislav;Polanc, Primož;Krofel, Miha;Slijepčević, Vedran;Gembarovski, Nina;Đurčević, Martina;Huber, Đuro

UDK 630*135+14 (Lynx lynx) (001)
Izvorni znanstveni članci – Original scientific papers
Šumarski list, 1–2 (2013): 43–49
ANALIZA RODOSLOVLJA DINARSKE
POPULACIJE RISA (Lynx lynx)
KINSHIP ANALYSIS OF DINARIC LYNX (Lynx lynx)
POPULATION
Magda SINDIČIĆ1, Tomislav GOMERČIĆ1, Primož POLANC2, Miha KROFEL3, Vedran SLIJEPČEVIĆ4,
Nina GEMBAROVSKI1, Martina ĐURČEVIĆ1, Đuro HUBER1
Sažetak:
Genetski markeri omogućuju nam uvid u srodstvene odnose divljih životinja kod kojih je takve podatke gotovo
nemoguće prikupiti izravnim promatranjem. Podaci o srodstvenim odnosima neophodni su za analize reproduk-
tivne uspješnosti, parenja u srodstvu, selekcije i protoka gena. Dinarska populacija euroazijskog risa (Lynx lynx)
uključuje risove u Sloveniji, Hrvatskoj te Bosni i Hercegovini. Vrsta je zakonom zaštićena u sve tri zemlje, te pop-
ulaciju ugrožavaju nedostatak genske raznolikosti i krivolov. Analizom srodnosti 91 genotipa, na 19 mikrosatelit-
skih lokusa, dinarske populacije risa dobili smo podatke o obiteljskim skupinama u kojima potomci dijele jednog
ili oba roditelja. Ukupno 21 od 29 analiziranih potomaka (72%) čiji su uzorci prikupljeni u razdoblju od 2000 do
2010. g. međusobno su povezani srodstvenim odnosima, potvrđujući nisku brojnost populacije i prisutnost parenja
u srodstvu.
KLJUČNE RIJEČI: mikrosateliti, rodoslovlje, euroazijski ris, Lynx lynx, Dinaridi
Uvod
Introduction
Razvojem analiza rodoslovnih odnosa na temelju DNK
otvorena su nova područja istraživanja etologije, ekologije,
evolucije i zaštite divljih životinja (Blouin 2003, Pemberton
2008, Van Horn i sur. 2008, Wang i Sature 2009, Jones i
Wang 2010a). Podaci o srodnosti daju nam uvid u odnos
jedinki u populaciji, objašnjavaju njihovo ponašanje i soci-
jalnu strukturu te sustav razmnožavanja (Ralls i sur. 2001,
Wang i Sature 2009). Međusobni odnosi životinja u srod-
stvu imaju velik utjecaj na teritorijalnost, migracije i raspro-
stranjenost populacije (Gompper i sur. 1998, Chapman i
sur. 2003). Osim toga, podaci o srodstvenim odnosima ne-
ophodni su za analize reproduktivne uspješnosti, parenja u
srodstvu, selekcije i protoka gena (Morgan i Conner 2001,
Nielsen i sur. 2001, Wang i Sature 2009, Jones i Wang 2010a).
Genetski markeri omogućuju nam uvid u srodstvene od-
nose divljih životinja kod kojih je takve podatke gotovo ne-
moguće prikupiti izravnim promatranjem (Blouin 2003,
Pemberton 2008, Jones i Wang 2010a). Mikrosatelitni lokusi
1  Dr. sc. Magda Sindičić dr. med. vet., doc. dr. sc. Tomislav Gomerčić dr. med. vet., Nina Gembarovski, Martina Đurčević, prof. dr. sc Đuro Huber dr. med.
vet.,Veterinarski fakultet Sveučilišta u Zagrebu, Heinzelova 55, 10 000 Zagreb, magda.sindicic@vef.hr, tomislav.gomercic@vef.hr, ngembarovski@vef.hr,
mdjurcevic@vef.hr, huber@vef.hr
2  Dr. sc. Primož Polanc, Odjel za biologiju Biotehnološki fakultet Sveučilišta u Ljubljani, Večna pot 111, 1000 Ljubljana, Slovenija, primozpolanc@yahoo.com
3  Dr. sc. Miha Krofel, Odjel za šumarstvo i obnovljive šumske izvore Biotehnološki fakultet Sveučilišta u Ljubljani, Večna pot 83, 1000 Ljubljana, Slovenija,
miha.krofel@gmail.com
4  Vedran Slijepčević dr. med. vet. Odjel lovstva i zaštite prirode Veleučilište u Karlovcu, J. J. Strossmayera 9, 47 000 Karlovac, vedran.slijepcevic@gmail.com
44
Šumarski list, 1–2, CXXXVII (2013), 43–49
smatraju se najpouzdanijim genetskim markerima za analizu
srodnosti, no analize se mogu raditi pomoću svih ko-domi-
nantnih markera (Jones i Ardren 2003). Osnova analize
srodnosti su podaci o frekvencijama alela, koji se statistički
obrađuju na temelju Mendelovih zakona o nasljeđivanju.
Jedinke svrstavamo u kategorije srodnosti, poput braće i
sestara koji dijele jednog ili oba roditelja, tako da na teme-
lju frekvencije alela unutar populacije računamo vjerojat-
nost pripadnosti određenoj kategoriji (Blouin 2003). Vje-
rojatnost se računa na temelju toga da li istraživane jedinke
dijele 0, 1 ili 2 alela koji imaju istovjetno podrijetlo (Blouin
2003). Razvijeno je više statističkih metoda za analize ro-
doslovlja na temelju DNK, te je izbor odgovarajuće metode
i kompjuterskog programa jedan od ključnih koraka u istra-
živanju (Blouin 2003). Prilikom izbora metode i programa
moramo uzeti u obzir sve parametre koji utječu na pouz-
danost i preciznost metode – broj i međusoban odnos uzor-
kovanih jedinki, sustav razmnožavanja istraživane vrste,
obilježja markera, broj lokusa i raznolikosti alela (Jones i
Ardren 2003, Van Horn i sur. 2008). Za analize na temelju
mikrosatelita važno je koristiti program čija metoda u obzir
uzima mutacije te pogreške prilikom genotipiziranja (poput
nul alela). Nul aleli znatno utječu na pouzdanost analiza, te
su najpouzdaniji modeli koji prilikom analiza u obzir uzi-
maju grešku genotipiziranja za svaki lokus pojedinačno (Jo-
nes i Ardren 2003, Wang 2004, Dakin i Avise 2004, Wagner
i sur. 2006).
Euroazijski ris (Lynx lynx) je zbog pretjeranog lova, gubitka
staništa i nedostatka plijena nestao s područja Dinarida kra-
jem 19. i početku 20. stoljeća (Kos 1928, Kos i sur. 2004,
Schmidt i sur. 2011). Godine 1973. naseljeno je šest jedinki
risa iz slovačkih Karpata u Sloveniju, odakle se populacija
proširila na Hrvatsku te Bosnu i Hercegovinu (Čop 1987,
Frković 2001). Danas dinarska populacija uključuje risove
u Sloveniji, južno od autoceste Ljubljana – Trst, risove u Hr-
vatskoj te Bosni i Hercegovini. Ne postoje pouzdani podaci
o brojnosti dinarske populacije, no veličina se procjenjuje
na oko 130 jedinki (Sindičić i sur. 2010a). Vrsta je zakonom
zaštićena u sve tri zemlje, te se smatra da populaciju ugro-
žavaju nedostatak genske raznolikosti i krivolov (Sindičić i
sur. 2010a, S
i
ndičić i sur. 2010b). Istraživanjem genetskih
markera potvrđen je pad brojnosti populacije u posljednjih
deset godina, niska heterozigotnost, niska efektivna veličina
populacije te prisutnost parenja u srodstvu (Sindičić 2011,
Polanc 2012, Sindičić i sur. 2012).
Budući parenje u srodstvu utječe na sposobnost preživlja-
vanja, podaci o rodoslovnim odnosima izuzetno su važni
za bolje poznavanje socijalne strukture i dinamike popula-
cije te zaštitu. Euroazijski ris je teritorijalna vrsta, živi na
velikom prostoru (Breitenmoser i sur. 2000), te je podatke
o srodstvenim odnosima gotovo nemoguće dobiti izravnim
promatranjem. Središnji dijelovi teritorija jedinki istoga
spola u pravilu se isključuju, no dolazi do preklapanja teri-
torija mužjaka s jednom ili više ženki (Breitenmoser i sur.
1993, Schmidt i sur. 1997). Ženke risa su spolno zrele s 10
do 20 mjeseci, a mužjaci s 30 mjeseci života, te su oba spola
poligamna (Kvam 1991). Parenje se odvija od veljače do
travnja, a nakon 69 dana gravidnosti ženka okoti od jedan
do četiri slijepa mladunca (Kvam 1991). Mladunci prvu go-
dinu života provode s majkom na njenom teritoriju, te se
na taj način mogu dobiti podaci o majčinstvu, no genetičke
metode su jedini izvor podataka o reproduktivnom uspjehu
mužjaka. Cilj ovoga rada bio je istražiti rodoslovne odnose
unutar sjevernog dijela dinarske populacije euroazijskog
risa uporabom mikrosatelitskih lokusa, te utvrditi učinko-
vitost korištenih lokusa za takve analize.
Materijali i metode
Materials and methods
U istraživanju smo koristili genotipove 19 mikrosatelitskih
lokusa 91 jedinke euroazijskog risa iz dinarske populacije.
Jedinke su genotipizirane iz uzoraka trofejnih krzna i lubanja
(58), krvi prikupljene od životinja obilježenih radioteleme-
trijskim ogrlicama (10), tkiva stradalih životinja (17) te uzo-
raka izmeta i dlaka prikupljenih neinvazivnim metodama
(6). Uzorci su prikupljeni u Hrvatskoj i Sloveniji (Slika 1).
Način prikupljanja uzoraka, genotipizirani lokusi, labora-
torijski postupci i analize opisani su u Polanc i sur. (2012).
Na temelju podataka dobivenih praćenjem jedinki pomoću
radio-telemetrijskih ogrlica bile su nam poznate četiri obi-
teljske skupine. Jedinke nazvane Ivan i Ana brat i sestra,
uhvaćeni su živi 2005. godine, no roditelji nisu bili poznati
(Š
trbenac i sur. 2007). Jedinka Luna obilježena je 2007. go-
Slika 1. Sivom bojom označeno je područje rasprostranjenosti dinarske
populacije euroazijskog risa (prema Linnell i sur. 2007), a crnim točkama
su označene lokacije podrijetla uzoraka koriš
tenih u istraživanju.
Figure 1. The shaded area indicates the Dinaric lynx population range (ac-
cording to Linnell et al. 2007). Dots represent sample locations.
45
SINDIČIĆ, M. et al.: ANALIZA RODOSLOVLJA DINARSKE POPULACIJE RISA (Lynx lynx)
dine kao mladunče, dok je bila uz majku nazvanu Tisa, koja
je također uhvaćena i obilježena 2008. godine (Slijepčević
i sur. 2009). Ženka Dina, obilježena 2006. godine (Krofel i
Kos. 2007), majka je dvoje mladunaca (Puhi i Burja), dok
je jedinka Snežka obilježena 2008. godine majka jedinkama
Copko i Točka (Krofel i sur. 2008).
Za analizu rodoslovnih odnosa korišten je program Colony
(Jones i Wang 2010b), u kojemu su implementirani metoda
vjerojatnosti za cijeli pedigre (Wang 2004, Wang i Sature
2009) te algoritam simuliranog prekaljivanja (Kirkpatrick i
sur. 1983). Programski algoritam istovremeno analizira vje-
rojatnost za cjelokupni pedigre te svrstava jedinke u obitelj-
ske skupine. Međusobni odnosi jedinki unutar obiteljske
skupine defi
 nirani su kao roditelj–potomak, braća i sestre
kojima su oba roditelja zajednički, te braća i sestre koji ima
ju
jednog zajedničkoj roditelja. Izračun vjerojatnosti odnosa
jedinki na temelju cjelokupnog pedigrea je precizniji od
vjerojatnosti na temelju odnosa samo te dvije jedinke ko-
jega koristi većina ostalih programa, te je ova metoda pri-
kladnija za uporabu kod poligamnih vrsta (Wang i Santure
2009).
Kao ulazne podatke smo koristili genotipove jedinki podi-
jeljene u tri skupine – potencijalni potomci (29 životinja čiji
su uzorci prikupljeni u razdoblju od 2000. do 2010. godine),
potencijalne majke (33 jedinke) te potencijalni očevi (58
jedinki). Ista jedinka se može nalaziti u skupini potencijal-
nih potomaka i potencijalnih roditelja, te smo s obzirom na
veličinu istraživane populacije, sustav razmnožavanja i broj
prikupljenih uzoraka, naveli 10% mogućnost da se traženi
roditelj nalazi u skupini potencijalnih roditelja. Kao pro-
gramske postavke izabrali smo poligamni sustav parenja za
oba spola, s prisutnim parenjem u srodstvu. Za svaki mi-
krosatelitni lokus naveli smo učestalost nul alela. U pro-
gramske postavke unijeli smo podatke o poznatim paro-
vima roditelj – potomak, brat – sestra, te za svakog potomka
popis jedinki koje na temelju podataka o godini rođenja i/
ili smrti isključujemo kao potencijalne roditelje. Za daljnju
analizu u obzir smo uzeli samo obiteljske skupine s vjero-
jatnošću iznad 0,95.
Rezultati
Results
Analiza srodnosti 91 genotipa dinarske populacije risa po-
moću programa Colony (Jones i Wang 2010b) je s vjerojat-
nošću >0,95 potvrdila srodstvene odnose koji su bili poznati
na temelju radio-telemetrijskih istraživanja, te smo dobili
podatke o obiteljskim skupinama u kojima potomci dijele
jednog ili oba roditelja (Tablica 1 i 2, Slika 2).
Analiza je potvrdila srodnost risova Ivana i Ane, koji su
uzorkovani 25. 10. 2005. g. u Gorskom kotaru kada su
pronađeni kao mladunci koji su došli u naselje Mrkopalj
Tablica 1. Obiteljske skupine s potomcima koji imaju bar jednog pozna-
tog roditelja ili dijele oba roditelja.
Table 1. Family groups with descendants sharing at least one identified
parent or both parents.
Obiteljska
skupina
Potomci
Majka
Vjerojatnost
majčinstva
Otac
Vjerojatnost
očinstva
1
Luna
Tisa
1.000
Nepoznat (*1)
2
Burja
Dina
1.000
H004
0.976
Puhi
1.000
0.976
3
LCRO9
Snežka
1.000
Nepoznat (*2)
Copko
1.000
Točka
1.000
Mihec
1.000
Nepoznat (*3)
R19
1.000
Nepoznat (*4)
4
Ivan
Nepoznata (#1)
Nepoznat (*5)
Ana
5
037Y
Nepoznata (#2)
Nepoznat (*6)
Lojzka

Tablica 2. Potomci koji dijele jednog roditelja.
Table 2. Descendants that share one parent.
Potomci
Vjerojatnost
1
R44
R19
1.000
2
R44
037Y
1.000
3
R44
Lojzka
1.000
4
R19
LCRO9
1.000
5
R19
Mihec
1.000
6
R19
Copko
1.000
7
R19
Točka
1.000
8
EJE
Copko
1.000
9
EJE
Točka
1.000
10
Dina
Dinko
1.000
11
Luna
Dinko
1.000
12
LCRO9
Mihec
1.000
13
Mihec
Copko
1.000
14
Mihec
Točka
1.000
15
EJE
LCRO9
0.999
16
EJE
H004
0.984
17
R19
037Y
0.981
18
R19
Lojzka
0.981
19
Ana
LCRO11
0.981
20
Ivan
LCRO11
0.981
21
039Y
Luna
0.968
22
039Y
Dinko
0.968
23
H004
Copko
0.963
24
H004
Točka
0.963
25
H004
LCRO9
0.962
26
EJE
Snežka
0.958
27
039Y
LCRO5
0.951
46
Šumarski list, 1–2, CXXXVII (2013), 43–49
(Štrbenac i sur. 2007). Ivan i Ana dijele oba roditelja, te je
utvrđeno da imaju istu majku kao i ženka LCRO11, čija je
lešina pronađena 17.6.2009. godine u Gorskom kotaru.
Analizom srodnosti potvrdili smo da je ženka Dina koja je
obilježena radiotelemetrijskom ogrlicom 28.12.2006. go-
dine na Snežniku u Sloveniji (Krofel i Kos 2007, Krofel i sur.
2008), majka dva risa (Puhi i Burja) čiji su uzorci dlake pri-
kupljeni iz njenog legla 30.6.2007. godine u Gorskom ko-
taru u Hrvatskoj. Mužjak Puhi je zatim 14.3.2008. uhvaćen
i obilježen ogrlicom na Snežniku u Sloveniji (Krofel i sur.
2009). Mužjak koji je genotipiziran iz uzorka dlake oznake
H004, prikupljene 14.12.2006. g. na Snežniku, otac je risova
Puhi i Burja. Utvrdili smo da risovica Dina ima istu majku
kao i mužjak Dinko, koji je 12.2.2008. g. uhvaćen i obilježen
ogrlicom u Gorskom kotaru. Dok Dinko ima istog oca kao
i Luna (za koju smo potvrdili da je kćer od Tise), te ženka
pod oznakom 039Y, koja je obilježena radio-telemetrijskom
ogrlicom na jugozapa
du Slovenije 10.3.2000. g. Ženka 039Y
ima istu majku kao i mužjak LCRO5 odstrijeljen 19. 5. 2004.
g. kod Senja.
Risovica Snežka, obilježena 9.2.2008. g. na Snežniku (Kro-
fel i sur. 2009) majka je petero potomaka (Slika 3). Od toga
tri potomka ima s istim mužjakom, no iz dva različita legla.
Tijekom radio-telemetrijskog praćenja ove ženke utvrđena
je lokacija njenog legla na Snežniku, te su 14.5.2008. g. pri-
kupljeni uzorci dlake mladunaca nazvanih Copko i Točka.
Utvrđeno je da ti mladunci imaju zajedničke roditelje s žen-
kom LCRO9, koja je 1.2.2009. g. godine pronađena mrtva
u Gorskom kotaru u Hrvatskoj. Dakle isti nepoznati muž-
jak (označen *2) je otac risovima
 označenim LCRO9, Copko
i Točka, mužjaku H004 (za kojega je utvrđeno da je s riso-
vicom Dinom roditelj potomcima Puhi i Burja), te jedinki
koja je genotipizirana iz uzorka izmeta označen EJE, koji
je prikupljen 16.2.2004. g. u Sloveniji na Kočevju. Risovica
Snežka je također majka mužjaku Mihecu, koji je obilje-
žen radiotelemetrijskom ogrlicom 23.12.2012. g. na Gu-
mancu na hrvatsko-slovenskoj granici (Krofel i sur. 2011),
te mužjaku R19 koji je u jugozapadnoj Sloveniji odstrijeljen
18.9.2001. g. Dakle mužjak R19 dijeli istu majku s LCRO9,
Copkom, Točkom i Mihecom, dok dijeli istog oca s jedin-
kama 037Y, Lojzkom i R44. Jedinka 037Y pronađena je mr-
tva 1.12.2002. g. godine u Loškom Potoku u Sloveniji. Ženka
Lojzka uhvaćena je i obilježena radiotelemetrijskom ogrli-
com na Ljubljanskom vrhu 7.3.2005. godine (Krofel i sur.
2006). Jedinka označena kao R44 genotipizirana je iz uzorka
mekuši, prikupljenog s trofejnog krzna mužjaka risa koji je
odstrijeljen u studenome 2001. g. u Notranjsko-kraškoj regiji
u Sloveniji. Zanimljivo je da jedinka EJE i Snežka imaju istu
majku, dok je nepoznati mužjak označen kao *2 otac EJE, te
je zajedno sa Snežkom roditelj LCRO9, Copku i Točki.
Rasprava
Discussion
U ovom istraživanju otvrdili smo da se analizom mikrosa-
telitskih lokusa pomoću programa Colony (Jones i Wang
2010b) može dobiti uvid u rodoslovne odnose unutar di-
narske populacije euroazijskog risa. Pritom smo koristili
genotipove 19 mikrosatelitskih lokusa, te smo dobili po-
datke s vjerojatnošću >0,95. Učinkovitost ovih 19 lokusa za
razlučivanje bliskih srodnika je visoka, te već kod uporabe
šest lokusa vjerojatnost identifi
 kacije jedinke iznosi 99,9%
Slika 2. Obiteljske skupine i potomci koji dijele jednog ili oba roditelja. Nepoznati očevi označeni su * i brojkom, a nepoznate majke # i brojkom.
Plavom bojom označene su očinske linije, a crvenom majčinske.
Figure 2. Family groups and descendants that share one or both parents. Unidentified fathers are marked with asterisk (*) and number, while unidenti-
fied mothers are marked with number sign (#) and number. Paternal lines are marked with blue and maternal with red color.
47
SINDIČIĆ, M. et al.: ANALIZA RODOSLOVLJA DINARSKE POPULACIJE RISA (Lynx lynx)
(Polanc i sur. 2012). Uporabom velikog broja lokusa, visoke
učinkovitosti razlučivanja, osigurali smo pouzdanost ana-
lize srodstvenih odnosa kod populacije koja se pari u srod-
stvu. Slična istraživanja na crvenom risu (Lynx rufus) kori-
stila su manji broj lokusa – osam (Janečka i sur. 2006) i 12
(Janečka i sur. 2007). U preliminarnom istraživanju (Đur-
čević i Gembarovski 2012) program Colony (Jones i Wang
2010b) pokazao se kao najpouzdaniji program za analizu
rodoslovnih odnosa kod dinarske populacije risa. Zbog ni-
ske raznolikosti alela u populaciji za pouzdanost analize
ključnim se pokazalo korištenje dodatnih parametara po-
put demografskih podataka, godina rođenja i smrti jedinki,
spol jedinke te uključivanje podataka o postotku greške pri
genotipizaciji (Gembarovski i Đurčević 2012).
Uzorci uključeni u analizu srodnosti su prikupljani u Hr-
vatskoj i Sloveniji. Ris je u Hrvatskoj prisutan na oko 11300
km2 (Sindičić i sur. 2010b), a u Sloveniji na 6300 km2 (Ko-
ren i sur. 2006), a prikupljeni uzorci većinom potječu iz sre-
dišnjeg dijela rasprostranjenosti populacije. U Europi se ras-
pon životnih prostora mužjaka risa kreće od 180 do 2780
km
2
, a ženki od 98 do 759 km
2
(Breitenmoser i sur. 2000).
Huber i sur. (1995) su u Sloveniji istraživali životni prostor
četiri risa, pri čemu su vrijednosti za mužjake bile 156 i 200
km
2
, te 132 i 222 km
2
 za ženke. Područje kretanja ženke risa
praćene u Sloveniji 2005. godine iznosilo je 109 km2 (Kro-
fel i sur. 2006). Odrasli mužjak risa čije je kretanje praćeno
u Gorskom kotaru pomoću radio-telemetrijske ogrlice, u
83 dana praćenja kretanjem je pokrio površinu od 116 km2,
dok je odrasla ženka u 54 dana praćenja svojim kretanjem
pokrila površinu od 62 km2 (Slijepčević i sur. 2009). Genet-
ska istraživanja pokazala su da unutar dinarske populacije
nije došlo do podjele na subpopulacije, tj. da fragmentira-
nost staništa ne utječe na kretanje životinja i potragu za
partnerima (Sindičić 2011, Polanc 2012). S obzirom na ve-
ličinu područja s kojeg su prikupljeni uzorci te područje
kretanja jedinki risa, u populacijama kod kojih se ne javlja
parenje u srodstvu i nema zapreka kretanju jedinki, kod
ovakve analize srodnosti očekuje se srodstvena povezanost
malog broja životinja. Ovim istraživanjem utvrdili smo da
od 29 životinja čiji uzorci su prikupljeni u razdoblju 2000–
2010. godine, njih 72 % je međusobno povezano srodstve-
nim odnosima. S obzirom na površinu s koje su prikupljeni
uzorci, podatak da fragmentacija staništa ne utječe na pro-
tok gena na istraživanom području, te površinu kretanja
jedinki euroazijskog risa u Sloveniji i Hrvatskoj, rodoslovna
povezanost tako velikog broja životinja nije očekivana i
smatramo da je posljedica parenja u srodstvu.
Istraživanja mikrosatelitskih lokusa pokazala su da je zbog
niske brojnosti kod risova iz dinarske populacije prisutno
parenje u srodstvu, te da se javlja parenje između polubraće
i polusestara, baka i djedova s potomcima, te srodnika u
prvom koljenu (Sindičić 2011, Polanc 2012). Među životi-
njama čije smo genotipove analizirali u ovom istraživanju
nismo pronašli primjere tako bliskog parenja u srodstvu,
no čak 21 od 29 analiziranih potomaka (72 %) čiji su uzorci
prikupljeni u razdoblju od 2000. do 2010. g. međusobno su
povezani srodstvenim odnosima. Smatramo da bi analizom
većeg broja uzoraka prikupljenih u kraćem razdoblju utvr-
dili prisutnost parenja među srodnicima u prvom ili dru-
gom koljenu. U svakom slučaju rezultati ovog istraživanja
su dodatna potvrda potrebe za unosom novih jedinki u po-
pulaciju.
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Summary:
Genetically identifi
 ed kinship relationships enable us valuable insight into wild animal pedigrees, which are
almost impossible to reconstruct by direct observations. Data on pedigrees are of fundamental importance for
analysis of reproductive success, inbreeding, selection and gene fl
 ow. Dinaric Eurasian lynx (Lynx lynx) in-
cludes animals from Slovenia, Croatia and Bosnia and Herzegovina. Th
 e species is strictly protected in all three
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microsatellite loci, using program Colony provided data about family groups with off
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both parents. A total of 21 (72 %) out of 29 analyzed off
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period, were related to each other, confi
 rming low population number and presence of inbreeding.
KEY WORDS: microsatellites, pedigree, Eurasian lynx, Lynx lynx, Dinaric Mountains