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

<s>The knowledge about plant material and its applicability is an important factor in achieving quality in landscape design.</s><s>It manifests itself in the appropriate selection of plant species for individual design solutions.</s><s>Nowadays we can observe the selection of plants based on fashion; measures defined by creation of a certain space characteristic are omitted.</s><s>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.</s><s>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.</s><s>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.</s><s>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.</s><s>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.</s><s>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.</s><s>The process of plants selection is shown on a concrete example, whereby certain design demands determine the choice of adequate plants.</s><s>For the final selection of plants it is necessary to add criteria arising from eco-physiological needs of the plants and from technological demands.</s>
<s>Odabir biljnih vrsta često se temelji na ukusu i modnom dobu, više nego na skupu karakteristika odabranih s ciljem stvaranja određene prostorne važnosti.</s><s>U literaturi biljne vrste se često obrađuju individualno i raspoređuju prema vidljivim učincima pojedinih biljaka.</s><s>Kako bi se zadovoljile potrebe projektanata u praksi, bilo je potrebno istraživanje biljnih vrsta s ciljem identifikacije skrivenog potencijala biljnih vrsta za design.</s><s>U tom smislu ljepota pojedinačne biljke podređena je prostornim potencijalom biljne vrste.</s><s>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.</s><s>U tom kontekstu važno je pokazati dosljednost između morfoloških karakteristika biljaka i vidljivog značenja drvoreda kao krajobraznog elementa.</s><s>Predmet istraživanja je biljni materijal: 193 različite vrste stablašica raspoređene su prema veličini, obliku, habitusu, teksturi, boji i razdoblju pojavljivanja pojedinih svojstava.</s><s>Predstavljene su mjere za odabir morfoloških karakteristika biljnih vrsta s kojima možemo postići točno odabrani prostorni učinak drvoreda.</s><s>Proces odabira biljaka prikazan je na određenom slučaju.</s><s>Na bazi određenih zahtjeva dizajna odabrane su odgovarajuće drvoredne vrste (stablašice).</s><s>Za konačni odabir biljnih vrsta potrebno je dodati dodatne daljne kriterije na bazi ekofizioloških potreba biljaka i tehnoloških kriterija.</s><s>Ključne riječi: oblikovanje krajobraza, urbani prostor, drvored, morfologija drveća, odabir biljnih vrsta</s>
<s>The knowledge about plant material and its applicability is an important factor in achieving quality in landscape design.</s><s>It manifests itself in the appropriate selection of plant species for individual design solutions.</s><s>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.</s><s>Here the plants become material, and people choose, arrange and remodel according to their needs and wishes.</s><s>M. Dobrilovič: Urban tree avenues - morphological characteristics as a factor in trees selection</s>
<s>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.</s><s>The primary elements or rather ‘building blocks’ of a designed landscape are called landscape elements.</s><s>They can be vegetation (lawn area, tree avenue) or non-vegetation (water and sand areas, buildings) (Ogrin, 1996).</s><s>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.</s><s>Merely a few basic elements have been treated, such as hedges and ground-cover plants.</s><s>The procedures of selection are nontransparent and quite inadequate for the use in a serious design.</s><s>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.</s><s>Rather than dealing with all the vegetation elements this paper focuses on vegetation areas.</s><s>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.</s><s>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.</s><s>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.</s>
<s>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.</s><s>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.</s><s>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).</s><s>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.</s><s>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).</s><s>In the history of the regional formation they first appeared in M. Dobrilovič: Urban tree avenues - morphological characteristics as a factor in trees selection</s>
<s>Figure 3. The garden of High Official at Thebes, 1400 b.c. (Cliford, 1962: 33) the Egyptian culture, where water canals were bordered by trees.</s><s>As regional elements tree avenues have been preserved until today.</s><s>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.</s><s>Among all the other line structures they are most appropriate for creating rhythm and sequence in space.</s>
<s>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.</s><s>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.</s><s>It has to be pointed out, that plants are not abstract M. Dobrilovič: Urban tree avenues - morphological characteristics as a factor in trees selection</s>
<s>116 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.</s><s>That is why besides the appearance of the plant it is important to take into consideration its ecological limitations or needs.</s>
<s>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...</s><s>The line can also be created with the trees that have irregular habitus.</s><s>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 • 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),</s>
<s>• 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),</s>
<s>• tolerate well city climate, salt in the soil, dust, hard ground, temperature oscillation, drought (for public parks in the cities and roadside plantings).</s><s>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.</s><s>By a negative space we understand the space between the ground and treetops (the interior of the tree avenue).</s><s>The form of this space depends on the tree species (size, shape, and habitus), density and the way of planting (Walker, 1991).</s><s>Further on only some forms of the negative space of tree avenues are shown.</s>
<s>• the space profile in the rectangular shape Treetops are dense, the rhythm of trunks is well expressed, and the visual field is wide.</s><s>The tree species that are appropriate for such tree avenues have tall, levelled and smooth trunks and ball-shaped, wide ball-shaped, domeshaped and compact treetops.</s><s>Appropriate habitus are irregular, upright and wide spread.</s>
<s>• 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.</s><s>The tree species that are appropriate for such tree avenues have tall, levelled and smooth trunks and egg-shaped treetops.</s><s>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.</s><s>The visual field is limited to the ending of the tree avenue.</s><s>The tree species that are appropriate for such tree avenues have short trunks, ball-shaped, domeshaped or cone-shaped treetops, regular and upright or irregular, upright and widely spread habitus.</s>
<s>• 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.</s><s>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).</s>
<s>• 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.</s><s>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.</s><s>The treetop is slightly pointed (the cone-shape of the treetop) or flat (cylinder-shape of the treetop).</s><s>Habitus is regular or upright.</s>
<s>The subjects of this research project are plant species designed for greening public spaces.</s><s>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.</s>
<s>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).</s>
<s>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.</s><s>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.</s>
<s>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.</s><s>Therefore further on the work process is introduced in some detail.</s><s>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 …).</s><s>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.</s><s>This was followed by a meticulous analysis in terms of visual properties (size, shape, colour, texture, durability of foliage).</s><s>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.</s><s>In order to help establish the aforementioned connections, a scheme is demonstrated in which the grey colour designates those morphological plant properties by which the character of tree avenues is defined.</s>
<s>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.</s><s>The selection process is introduced by means of a concrete example and represents one of the results of this research.</s><s>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).</s><s>The criteria are introduced gradually, each added criteria representing a new level or rather combination of morphological properties.</s><s>We consider the most suitable those species that conform most optimally to all levels of selection and all combinations of criteria.</s><s>M. Dobrilovič: Urban tree avenues - morphological characteristics as a factor in trees selection</s>
<s>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.</s><s>It is, however, of great importance for the process of selection itself.</s><s>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.</s>
<s>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.</s>
<s>The selection of plant species in terms of desired visual effects of tree avenues will now be demonstrated on a concrete example.</s><s>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.</s><s>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.</s>
<s>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.</s>
<s>• foliage durability: deciduous rounded to oval, high domed, closed crown informal, upright, widely spreading, hanging slightly medium</s>
<s>The dense, medium height tree avenue with the tube-shaped interior, is best completed by the species Acer saccharum and Carpinus betulus.</s><s>Both species have low trunks, dense and compact treetops and tolerate well dense planting the condition in the tree avenue.</s><s>They tolerate well the salt in the soil, which is an important characteristic of tree species.</s><s>Branches on the exterior side of the tree avenue with older plants are slightly curved and create a visual protection.</s><s>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.</s><s>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.</s><s>Due to the eco- physiological unsuitability we can also eliminate Acer negundo (it does not tolerate well the overshadowing).</s>
<s>The method used and described here for the selection of plant species represents a novelty in the field of landscape design.</s><s>By means of this research it was ascertained that the manner of selecting plants according to their visual properties is appropriate.</s><s>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).</s><s>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.</s><s>The results of the research are applicable on different levels.</s><s>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.</s><s>The data can be combined arbitrarily in order to search for cross-section multitudes that in essence represent combinations of morphological properties.</s><s>The data and the above mentioned systematizations are not presented in this paper, due to the extensiveness of the material.</s>
<s>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.</s><s>Which criteria are to be included and in what order, largely depends on the design goal.</s><s>The paper demonstrates the process of selection of plant species suitable for tree avenues.</s><s>In a similar way suitable plants for other landscape elements can be chosen.</s>
<s>The presented process of selection is not applicable only in landscape architecture.</s><s>It can be equally useful in selective breeding, since it can help determine the need for a particular shape, habitus, colour …</s><s>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.</s>


Analiza rodoslovlja dinarske populacije risa (Lynx lynx)

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

<s>Genetski markeri omogućuju nam uvid u srodstvene odnose divljih životinja kod kojih je takve podatke gotovo nemoguće prikupiti izravnim promatranjem.</s><s>Podaci o srodstvenim odnosima neophodni su za analize reproduktivne uspješnosti, parenja u srodstvu, selekcije i protoka gena.</s><s>Dinarska populacija euroazijskog risa (Lynx lynx) uključuje risove u Sloveniji, Hrvatskoj te Bosni i Hercegovini.</s><s>Vrsta je zakonom zaštićena u sve tri zemlje, te populaciju ugrožavaju nedostatak genske raznolikosti i krivolov.</s><s>Analizom srodnosti 91 genotipa, na 19 mikrosatelitskih lokusa, dinarske populacije risa dobili smo podatke o obiteljskim skupinama u kojima potomci dijele jednog ili oba roditelja.</s><s>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.</s>
<s>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.</s><s>2008, Wang i Sature 2009, Jones i Wang 2010a).</s><s>Podaci o srodnosti daju nam uvid u odnos jedinki u populaciji, objašnjavaju njihovo ponašanje i socijalnu strukturu te sustav razmnožavanja (Ralls i sur.</s><s>2001, Wang i Sature 2009).</s><s>Međusobni odnosi životinja u srodstvu imaju velik utjecaj na teritorijalnost, migracije i rasprostranjenost populacije (Gompper i sur.</s><s>1998, Chapman i sur.</s><s>2003).</s><s>Osim toga, podaci o srodstvenim odnosima neophodni su za analize reproduktivne uspješnosti, parenja u srodstvu, selekcije i protoka gena (Morgan i Conner 2001, Nielsen i sur.</s><s>2001, Wang i Sature 2009, Jones i Wang 2010a).</s><s>Genetski markeri omogućuju nam uvid u srodstvene odnose divljih životinja kod kojih je takve podatke gotovo nemoguće prikupiti izravnim promatranjem (Blouin 2003, Pemberton 2008, Jones i Wang 2010a).</s><s>Mikrosatelitni lokusi vet.,Veterinarski fakultet Sveučilišta u Zagrebu, Heinzelova 55, 10 000 Zagreb, magda.sindicicŽvef.hr, tomislav.gomercicŽvef.hr, ngembarovskiŽvef.hr, mdjurcevicŽvef.hr, huberŽvef.hr</s>
<s>4 Vedran Slijepčević dr. med. vet.</s><s>Odjel lovstva i zaštite prirode Veleučilište u Karlovcu, J. J. Strossmayera 9, 47 000 Karlovac, vedran.slijepcevicŽgmail.com smatraju se najpouzdanijim genetskim markerima za analizu srodnosti, no analize se mogu raditi pomoću svih ko-dominantnih markera (Jones i Ardren 2003).</s><s>Osnova analize srodnosti su podaci o frekvencijama alela, koji se statistički obrađuju na temelju Mendelovih zakona o nasljeđivanju.</s><s>Jedinke svrstavamo u kategorije srodnosti, poput braće i sestara koji dijele jednog ili oba roditelja, tako da na temelju frekvencije alela unutar populacije računamo vjerojatnost pripadnosti određenoj kategoriji (Blouin 2003).</s><s>Vjerojatnost se računa na temelju toga da li istraživane jedinke dijele 0, 1 ili 2 alela koji imaju istovjetno podrijetlo (Blouin 2003).</s><s>Razvijeno je više statističkih metoda za analize rodoslovlja na temelju DNK, te je izbor odgovarajuće metode i kompjuterskog programa jedan od ključnih koraka u istraživanju (Blouin 2003).</s><s>Prilikom izbora metode i programa moramo uzeti u obzir sve parametre koji utječu na pouzdanost i preciznost metode – broj i međusoban odnos uzorkovanih jedinki, sustav razmnožavanja istraživane vrste, obilježja markera, broj lokusa i raznolikosti alela (Jones i Ardren 2003, Van Horn i sur.</s><s>2008).</s><s>Za analize na temelju mikrosatelita važno je koristiti program čija metoda u obzir uzima mutacije te pogreške prilikom genotipiziranja (poput nul alela).</s><s>Nul aleli znatno utječu na pouzdanost analiza, te su najpouzdaniji modeli koji prilikom analiza u obzir uzimaju grešku genotipiziranja za svaki lokus pojedinačno (Jones i Ardren 2003, Wang 2004, Dakin i Avise 2004, Wagner Euroazijski ris (Lynx lynx) je zbog pretjeranog lova, gubitka staništa i nedostatka plijena nestao s područja Dinarida kraSchmidt i sur.</s><s>2011).</s><s>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).</s><s>Danas dinarska populacija uključuje risove u Sloveniji, južno od autoceste Ljubljana – Trst, risove u Hrvatskoj te Bosni i Hercegovini.</s><s>Ne postoje pouzdani podaci o brojnosti dinarske populacije, no veličina se procjenjuje na oko 130 jedinki (Sindičić i sur.</s><s>2010a).</s><s>Vrsta je zakonom zaštićena u sve tri zemlje, te se smatra da populaciju ugrožavaju nedostatak genske raznolikosti i krivolov (Sindičić i ndičić i sur.</s><s>2010b).</s><s>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.</s><s>2012).</s>
<s>Budući parenje u srodstvu utječe na sposobnost preživljavanja, podaci o rodoslovnim odnosima izuzetno su važni za bolje poznavanje socijalne strukture i dinamike populacije te zaštitu.</s><s>Euroazijski ris je teritorijalna vrsta, živi na velikom prostoru (Breitenmoser i sur.</s><s>2000), te je podatke o srodstvenim odnosima gotovo nemoguće dobiti izravnim promatranjem.</s><s>Središnji dijelovi teritorija jedinki istoga spola u pravilu se isključuju, no dolazi do preklapanja teritorija mužjaka s jednom ili više ženki (Breitenmoser i sur.</s><s>1993, Schmidt i sur.</s><s>1997).</s><s>Ženke risa su spolno zrele s 10 do 20 mjeseci, a mužjaci s 30 mjeseci života, te su oba spola poligamna (Kvam 1991).</s><s>Parenje se odvija od veljače do travnja, a nakon 69 dana gravidnosti ženka okoti od jedan do četiri slijepa mladunca (Kvam 1991).</s><s>Mladunci prvu godinu 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.</s><s>Cilj ovoga rada bio je istražiti rodoslovne odnose unutar sjevernog dijela dinarske populacije euroazijskog risa uporabom mikrosatelitskih lokusa, te utvrditi učinkovitost korištenih lokusa za takve analize.</s>
<s>U istraživanju smo koristili genotipove 19 mikrosatelitskih lokusa 91 jedinke euroazijskog risa iz dinarske populacije.</s><s>Jedinke su genotipizirane iz uzoraka trofejnih krzna i lubanja (58), krvi prikupljene od životinja obilježenih radiotelemetrijskim ogrlicama (10), tkiva stradalih životinja (17) te uzoraka izmeta i dlaka prikupljenih neinvazivnim metodama (6).</s><s>Uzorci su prikupljeni u Hrvatskoj i Sloveniji (Slika 1).</s><s>Način prikupljanja uzoraka, genotipizirani lokusi, laboratorijski postupci i analize opisani su u Polanc i sur. (2012).</s><s>Na temelju podataka dobivenih praćenjem jedinki pomoću radio-telemetrijskih ogrlica bile su nam poznate četiri obiteljske skupine.</s><s>Jedinke nazvane Ivan i Ana brat i sestra, uhvaćeni su živi 2005. godine, no roditelji nisu bili poznati trbenac i sur.</s><s>2007).</s><s>Jedinka Luna obilježena je 2007. goSlika 1. Sivom bojom označeno je područje rasprostranjenosti dinarske populacije euroazijskog risa (prema Linnell i sur.</s><s>2007), a crnim točkama su označene lokacije podrijetla uzoraka koriš tenih u istraživanju.</s>
<s>Figure 1. The shaded area indicates the Dinaric lynx population range (according to Linnell et al. 2007).</s><s>Dots represent sample locations.</s><s>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.</s><s>2009).</s><s>Ženka Dina, obilježena 2006. godine (Krofel i Kos.</s><s>2007), majka je dvoje mladunaca (Puhi i Burja), dok je jedinka Snežka obilježena 2008. godine majka jedinkama 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.</s><s>1983).</s><s>Programski algoritam istovremeno analizira vjerojatnost za cjelokupni pedigre te svrstava jedinke u obiteljske skupine.</s><s>Međusobni odnosi jedinki unutar obiteljske skupine defi</s>
<s>Materials and methods nirani su kao roditelj–potomak, braća i sestre kojima su oba roditelja zajednički, te braća i sestre koji ima jednog zajedničkoj roditelja.</s><s>Izračun vjerojatnosti odnosa jedinki na temelju cjelokupnog pedigrea je precizniji od vjerojatnosti na temelju odnosa samo te dvije jedinke kojega koristi većina ostalih programa, te je ova metoda prikladnija za uporabu kod poligamnih vrsta (Wang i Santure Kao ulazne podatke smo koristili genotipove jedinki podijeljene 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).</s><s>Ista jedinka se može nalaziti u skupini potencijalnih 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.</s><s>Kao programske postavke izabrali smo poligamni sustav parenja za oba spola, s prisutnim parenjem u srodstvu.</s><s>Za svaki mikrosatelitni lokus naveli smo učestalost nul alela.</s><s>U programske postavke unijeli smo podatke o poznatim parovima roditelj – potomak, brat – sestra, te za svakog potomka popis jedinki koje na temelju podataka o godini rođenja i/ ili smrti isključujemo kao potencijalne roditelje.</s><s>Za daljnju analizu u obzir smo uzeli samo obiteljske skupine s vjerojatnošću iznad 0,95.</s>
<s>Analiza srodnosti 91 genotipa dinarske populacije risa pomoću programa Colony (Jones i Wang 2010b) je s vjerojatnošću >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).</s><s>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 poznatog roditelja ili dijele oba roditelja.</s>
<s>(Štrbenac i sur.</s><s>2007).</s><s>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.</s><s>Analizom srodnosti potvrdili smo da je ženka Dina koja je obilježena radiotelemetrijskom ogrlicom 28.12.2006. godine na Snežniku u Sloveniji (Krofel i Kos 2007, Krofel i sur.</s><s>2008), majka dva risa (Puhi i Burja) čiji su uzorci dlake prikupljeni iz njenog legla 30.6.2007. godine u Gorskom kotaru u Hrvatskoj.</s><s>Mužjak Puhi je zatim 14.3.2008. uhvaćen i obilježen ogrlicom na Snežniku u Sloveniji (Krofel i sur.</s><s>2009).</s><s>Mužjak koji je genotipiziran iz uzorka dlake oznake H004, prikupljene 14.12.2006. g. na Snežniku, otac je risova Puhi i Burja.</s><s>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.</s><s>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</s>
<s>du Slovenije 10.3.2000. g. Ženka 039Y ima istu majku kao i mužjak LCRO5 odstrijeljen 19. 5. 2004.</s><s>Risovica Snežka, obilježena 9.2.2008. g. na Snežniku (Krofel i sur.</s><s>2009) majka je petero potomaka (Slika 3).</s><s>Od toga tri potomka ima s istim mužjakom, no iz dva različita legla.</s><s>Tijekom radio-telemetrijskog praćenja ove ženke utvrđena je lokacija njenog legla na Snežniku, te su 14.5.2008. g. prikupljeni uzorci dlake mladunaca nazvanih Copko i Točka.</s><s>Utvrđeno je da ti mladunci imaju zajedničke roditelje s ženkom LCRO9, koja je 1.2.2009. g. godine pronađena mrtva u Gorskom kotaru u Hrvatskoj.</s><s>Dakle isti nepoznati mužjak (označen *2) je otac risovima</s>
<s>Rasprava i Točka, mužjaku H004 (za kojega je utvrđeno da je s risovicom 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.</s><s>Risovica Snežka je također majka mužjaku Mihecu, koji je obilježen radiotelemetrijskom ogrlicom 23.12.2012. g. na Gumancu na hrvatsko-slovenskoj granici (Krofel i sur.</s><s>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 jedinkama 037Y, Lojzkom i R44.</s><s>Jedinka 037Y pronađena je mrtva 1.12.2002. g. godine u Loškom Potoku u Sloveniji.</s><s>Ženka Lojzka uhvaćena je i obilježena radiotelemetrijskom ogrlicom na Ljubljanskom vrhu 7.3.2005. godine (Krofel i sur.</s><s>2006).</s><s>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.</s><s>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.</s>
<s>U ovom istraživanju otvrdili smo da se analizom mikrosatelitskih lokusa pomoću programa Colony (Jones i Wang 2010b) može dobiti uvid u rodoslovne odnose unutar dinarske populacije euroazijskog risa.</s><s>Pritom smo koristili genotipove 19 mikrosatelitskih lokusa, te smo dobili podatke s vjerojatnošću >0,95.</s><s>Učinkovitost ovih 19 lokusa za razlučivanje bliskih srodnika je visoka, te već kod uporabe šest lokusa vjerojatnost identifi</s>
<s>Slika 2. Obiteljske skupine i potomci koji dijele jednog ili oba roditelja.</s><s>Nepoznati očevi označeni su * i brojkom, a nepoznate majke # i brojkom.</s><s>Plavom bojom označene su očinske linije, a crvenom majčinske.</s><s>Figure 2. Family groups and descendants that share one or both parents.</s><s>Unidentified fathers are marked with asterisk (*) and number, while unidentified mothers are marked with number sign (#) and number.</s><s>Paternal lines are marked with blue and maternal with red color.</s><s>SINDIČIĆ, M. et al.: ANALIZA RODOSLOVLJA DINARSKE POPULACIJE RISA (Lynx lynx) (Polanc i sur.</s><s>2012).</s><s>Uporabom velikog broja lokusa, visoke učinkovitosti razlučivanja, osigurali smo pouzdanost analize srodstvenih odnosa kod populacije koja se pari u srodstvu.</s><s>Slična istraživanja na crvenom risu (Lynx rufus) koristila su manji broj lokusa – osam (Janečka i sur.</s><s>2006) i 12 (Janečka i sur.</s><s>2007).</s><s>U preliminarnom istraživanju (Đurčević i Gembarovski 2012) program Colony (Jones i Wang 2010b) pokazao se kao najpouzdaniji program za analizu rodoslovnih odnosa kod dinarske populacije risa.</s><s>Zbog niske raznolikosti alela u populaciji za pouzdanost analize ključnim se pokazalo korištenje dodatnih parametara poput demografskih podataka, godina rođenja i smrti jedinki, spol jedinke te uključivanje podataka o postotku greške pri genotipizaciji (Gembarovski i Đurčević 2012).</s><s>Uzorci uključeni u analizu srodnosti su prikupljani u Hrvatskoj i Sloveniji.</s><s>Ris je u Hrvatskoj prisutan na oko 11300 km2 (Sindičić i sur.</s><s>2010b), a u Sloveniji na 6300 km2 (Koren i sur.</s><s>2006), a prikupljeni uzorci većinom potječu iz središnjeg dijela rasprostranjenosti populacije.</s><s>U Europi se raspon životnih prostora mužjaka risa kreće od 180 do 2780 (Breitenmoser i sur.</s><s>2000).</s>
<s>Huber i sur. (1995) su u Sloveniji istraživali životni prostor četiri risa, pri čemu su vrijednosti za mužjake bile 156 i 200 za ženke.</s><s>Područje kretanja ženke risa</s>
<s>kacije jedinke iznosi 99,9% praćene u Sloveniji 2005. godine iznosilo je 109 km2 (Krofel i sur.</s><s>2006).</s><s>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.</s><s>2009).</s><s>Genetska istraživanja pokazala su da unutar dinarske populacije nije došlo do podjele na subpopulacije, tj. da fragmentiranost staništa ne utječe na kretanje životinja i potragu za partnerima (Sindičić 2011, Polanc 2012).</s><s>S obzirom na veličinu 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.</s><s>Ovim istraživanjem utvrdili smo da od 29 životinja čiji uzorci su prikupljeni u razdoblju 20002010. godine, njih 72 % je međusobno povezano srodstvenim odnosima.</s><s>S obzirom na površinu s koje su prikupljeni uzorci, podatak da fragmentacija staništa ne utječe na protok 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.</s><s>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).</s><s>Među životinjama 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.</s><s>Smatramo da bi analizom većeg broja uzoraka prikupljenih u kraćem razdoblju utvrdili prisutnost parenja među srodnicima u prvom ili drugom koljenu.</s><s>U svakom slučaju rezultati ovog istraživanja su dodatna potvrda potrebe za unosom novih jedinki u populaciju.</s>
<s>• Blouin, M. S., 2003: DNA-based methods for pedigree reconstruction and kinship analysis in natural populations.</s><s>Trends in Ecology and Evolution 18: 503–511.</s>
<s>• Breitenmoser, U., C. Breitenmoser-Würsten, H. Okarma, T. Kaphegyi, U. Kaphegyi-Wallmann, U. M. Müller, 2000: Action plan for the conservation of the Eurasian lynx in Europe (Lynx lynx), Council of Europe Publishing, 69 str., Strasbourg.</s><s>Slika 3. Prikaz lokacija prikupljenih uzoraka, žen Snežke (puni plavi krug), njenih petero potomaka (plava kružnica), te njihove polubraće i polusestara s kojima imaju zajedničkog oca (žuto/crvenom zvjezdicom su označeni potomci istog oca).</s><s>Plavim linijama su označeni teritoriji Snežke i MiFigure 3. Sampling locations for female Snež (blue circle), her five descendants (blue circular), and their half siblings (yellow/blue asterisk indicates offspring of the same father).</s><s>Territories of Mihec and Snežka are marked with blue lines.</s>
<s>R. C. Fleischer, 2001: Kinship, social relations, and den sharing in kit foxes.</s><s>Journal of Mammalogy 82: 858–866. e im</s>
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<s>Genetically identifi ed kinship relationships enable us valuable insight into wild animal pedigrees, which are almost impossible to reconstruct by direct observations.</s><s>Data on pedigrees are of fundamental importance for analysis of reproductive success, inbreeding, selection and gene fl ow.</s><s>Dinaric Eurasian lynx (Lynx lynx) in</s>
<s>Genetically identifi cludes animals from Slovenia, Croatia and Bosnia and Herzegovina.</s><s>Th e species is strictly protected in all three countries, being threatened by loss of genetic diversity and poaching.</s><s>Kinship analysis of 91 genotypes, on 19 microsatellite loci, using program Colony provided data about family groups with off spring sharing one or</s>