Research


EVOLUTION, SYSTEMATICS AND BIOGEOGRAPHY OF THE MEDITERRANEAN MOUNTAIN FLORA



The floristically rich and biogeographically interesting Dinaric Alps have attracted me from the beginning of my botanical studies. My main topics of interests are the flora, vegetation, biogeography, taxonomy and conservation of polymorphic (e.g. taxa of the genera Edraianthus, Moehringia, Wulfenia, Athamanta, Astragalus sections Incani, Trachycercis and Dissitiflori, Drypis, Festuca varia agg.), rare, and endemic taxa of the South-eastern Europe and the Mediterranean.


Systematics of the genus Edraianthus (Campanulaceae)

New species of the genus Edraianthus and a change in taxonomic status for Edraianthus serpyllifolius f. pilosulus from the Balkan Peninsula

We described and illustrated a new species, Edraianthus pulevicii (Campanulaceae), endemic to the Durmitor mountain range (SE Dinaric Alps, Montenegro, Balkan Peninsula). Additionally, we proposed a specific rank for E. serpyllifolius f. pilosulus from the Komovi Mts. (NW Prokletije mountain range, Montenegro, Balkan Peninsula), E. pilosulus . Both are morphologically similar to E. serpyllifolius s. s., all having spathulate leaves. Edraianthus pulevicii differs from E. serpyllifolius s. s. in having distinctly crenate as well as much longer and broader basal and cauline leaves with indumentum on the upper side, hairs on the leaf margin and leaf surface oriented towards the leaf base, broader and longer bracts, and revolute calyx lobes reflexed at the apex. Edraianthus pilosulus is morphologically similar to both E. serpyllifolius and E. pulevicii but differs from the former in having more or less dense hairs on the upper side of the leaves, and from the latter in having considerably smaller basal and cauline leaves with no or sparse crenation, hairs on the leaf margin and the leaf surface oriented towards the point, and narrower bracts. A new systematic treatment is supported by chloroplast DNA sequence data and AFLP fingerprinting data.
  • Contributors: Gerald M. Schneeweiss, Peter Schönswetter (both Universiy of Vienna, Austria), Tamara Rakić, Vlado Stevanović, Dmitar Lakušić, (all University of Belgrade, Serbia), Saša Stefanović (University of Toronto at Mississauga, Canada).
  • Funding: European Community (MEIF-CT-2005024315)
  • Reference: Systematic Botany, 34 (3): 602-608, 2009
 




    
     Sepals of E. pulevicii, E. serpyllifolius and E. pilosulus (photo: Dmitar Lakušić)
 
   Bracts of the relevant Edraianthus taxa with spatulate leaves  (photo: Dmitar Lakušić)



                      
Edraianthus x lakusicii a new intersectional natural hybrid: morphological and molecular evidence

During our recent fieldtrips to Mt Lovćen (Dinaric Alps, Montenegro), we encountered putative hybrid individuals, morphologically intermediate between two sympatric taxa, E. tenuifolius (E. sect. Edraianthus) and E. wettsteinii subsp. lovcenicus (E. sect. Uniflori). Multivariate morphometric and molecular analyses were carried out to investigate the occurrence of hybridization between these two species. As a result, a new nothospecies is described here, Edraianthus x lakusici V. Stevanović & D. Lakusić, a natural hybrid between E. tenuifolius and E. wettsteinii subsp. lovcenicus. At present, this hybrid is known only from the single locality of Mt. Lovćen. Its population size is estimated to be <50 mature individuals and the estimated ‘‘area of occupancy’’ is smaller than 1 km2.


Differences between a–c Edraianthus x lakusicii nothosp. nova, Holotypus (a habitus, b inflorescence with involucral bracts, c flowers with calyx teeth), d–f E. wettsteinii subsp. lovcenicus (d habitus, e inflorescens with involucral bracts, f flowers with calyx teeth), g–i Edraianthus tenuifolius (g habitus, h inflorescens with involucral bracts, i flowers with calyx teeth). All from locus classicus, Montenegro, Mt. Lovćen, Branjevine above village Mirac (photo: Dmitar Lakušić).


















  • Contributors: Tamara Rakić, Vlado Stevanović, Dmitar Lakušić, (all University of Belgrade, Serbia), Saša Stefanović (University of Toronto at Mississauga, Canada), Peter Schönswetter, Gerald M. Schneeweiss, (both Universiy of Vienna, Austria)
  • Funding: European Community (MEIF-CT-2005024315)
  • Reference: Plant Systematics and Evolution,  280: 77-88, 2009



Validation of the name Edraianthus sutjeskae (Campanulaceae), a narrow endemic of southeastern Dinaric Alps (eastern Bosnia and Herzegovina)


Edraianthus sutjeskae R. Lakušić, originally described in 1974, was an invalidly published name because in its type citation the collector’s name and collection date were omitted. We validated the name by indicating the holotype kept at the National Museum of Bosnia and Herzegovina (SARA).

Herbarium label of the holotype of
Edraianthus sutjeskae
, a narrow endemic of the Sutjeska canyon (Bosnia and Herzegovina), collected by Karlo Maly (a renowned botanist and connoisseur of the Bosnian flora) on 11th of August 1925 and kept at the National Museum of Bosnia and Herzegovina in Sarajevo (Photo: Dejan Kulier).











  • Contributors: Dmitar Lakušić (University of Belgrade, Serbia), Gerald M. Schneeweiss, Peter Schönswetter (both University of Vienna, Austria), Sabaheta Abadžić (Sarajevo, National Museum of Bosnia and Herzegovina)
  • Funding: European Community (MEIF-CT-2005024315)
  • Reference: Nordic Journal of Botany, 28: 603-604, 2010

Evolution on the Balkan Peninsula: Phylogeny and phylogeography of the genera Edraianthus and Heliosperma

[Edraianthus glisicii] Many surveys confirm that the Balkan Peninsula in general is a hotspot of European biodiversity, but the extremely high level of endemism in the Balkans is probably the most striking element.In order to contribute to an explanation of this phenomenon, we use two model plant genera (Edraianthus, Heliosperma) to address the following objectives within the project: establishment of sound molecular phylogenetic hypotheses on the relationships of the study groups both to its closest relatives as well as within the respective genera; testing the Pleistocene speciation hypothesis; establishment of sound comparative phylogeographic hypotheses within species and species groups; pinpointing the putative centres of origin of the respective groups based on the phylogenetic and phylogeographic pattern; determining the migration paths from the Balkans to the Apennines; determining frequency and timing of the colonisation of the Apennines and pinpointing secondary centres of origin in the Apennines. Research was carried out applying molecular methods (DNA - sequencing, cpDNA haplotype analysis, DNA-fingerprinting (AFLP), molecular dating), morphometrics, and biogeographic analyses at the Institute of Botany, Department of Biogeography and Botanical Garden, University of Vienna, Vienna.


E
draianthus glisicii, a narrow endemic of the Tara canyon.


  • Contributors: Harald Niklfeld, Peter Schönswetter, Gerald M. Schneeweiss (all University of Vienna, Austria)
  • Funding: European Community (MEIF-CT-2005024315)


Quaternary range dynamics of ecologically contrasting species (Edraianthus serpyllifolius and E. tenuifolius, Campanulaceae) within the Balkan refugium


We investigated Quaternary range dynamics of two closely related, but ecologically divergent species (cold-tolerant Edraianthus serpyllifolius and thermophilic Edraianthus tenuifolius) with overlapping distribution ranges endemic to the western Balkan Peninsula, an important yet understudied Pleistocene refugium. Our aims were to test predictions of the ‘refugia-within-refugia’ model of strong genetic subdivisions due to population isolation in separate refugia; to explore whether two ecologically divergent species reacted differently to Pleistocene climatic fluctuations; and to test predictions of the displacement refugia model of stronger differentiation among populations in the thermophilic E. tenuifolius compared with the cold-tolerant E. serpyllifolius. We gathered AFLP data and plastid DNA (cpDNA) sequences from two to five individuals from 10 populations of E. serpyllifolius and 22 populations of E. tenuifolius, spanning their entire respective distribution areas.

Distribution ranges and sampled populations of Edraianthus serpyllifolius (solid line and black circles) and E. tenuifolius (hatched
line and white circles) on the western Balkan Peninsula. The insert shows the position of the sampling area in south-eastern Europe.
















AFLP data were analysed using a Bayesian clustering approach and a distance-based network approach. Plastid sequences were used to depict relationships among haplotypes in a statistical parsimony network, and to obtain age estimates in a Bayesian framework. In E. serpyllifolius, both AFLP and plastid sequence data showed clear geographic structure. Western populations showed high AFLP diversity and a high number of rare fragments. In E. tenuifolius, both markers congruently identified a major phylogeographic split along the lower Neretva valley in central Dalmatia. The most distinct and earliest diverging cpDNA haplotypes were found further south in the south-easternmost populations. North-western populations, identified as a separate cluster by Bayesian clustering, were characterized by low genetic diversity and a low number of rare AFLP markers.

Sampled populations and patterns of amplified fragment length polymorphism (AFLP) and plastid DNA variation of Edraianthus
serpyllifolius on the western Balkan Peninsula. (a) Sampled populations (see Table 1 for further details). (b) NeighborNet diagram based on (a) Nei–Li distance matrix (Nei & Li, 1979). Bootstrap values above 50% derived from a neighbour-joining analysis are given for the main
branches. (c) Nei’s (1987) gene diversity. (d) Frequency down-weighted marker values (Scho¨nswetter & Tribsch, 2005; Winkler et al., 2010). In (c) and (d), the size of the dots is directly proportional to the depicted values. (e) Bayesian clustering of AFLP data using the software structure. The three gene pools obtained in the optimal solution are colour-coded (white, grey and black). Populations may be composed of one or several gene-pools. (f) Geographic distribution of plastid DNA haplotypes derived from concatenated sequences of trnGUCC–trnRUCU, rpl32–trnLUAG and trnTUGU–trnLUAA–trnFGAA intergenic spacers. (g) Statistical parsimony network of plastid DNA haplotypes. (h) Majority rule consensus tree from strict clock Bayesian analysis with the software beast. Node heights correspond to median ages. Numbers along branches are Bayesian posterior probabilities; identical haplotypes or unresolved polytomies are collapsed as triangles, their vertical extension being proportional to the number of individuals.










Sampled populations and patterns of amplified fragment length polymorphism (AFLP) and plastid DNA variation of Edraianthus
tenuifolius on the western Balkan Peninsula. (a) Sampled populations (see Table 1 for further details). (b) NeighborNet diagram based on (a) Nei–Li distance matrix (Nei & Li, 1979). Bootstrap values above 50% derived from a neighbour-joining analysis of the same matrix are given for the main branches. (c) Nei’s (1987) gene diversity. (d) Frequency down-weighted marker values (Scho¨nswetter & Tribsch, 2005; Winkler et al., 2010). In (c) and (d), the size of the dots is directly proportional to the depicted values. (e) Bayesian clustering of AFLP data using the software structure. The three gene pools obtained in the optimal solution are colour-coded (white, grey and black). Populations may be composed of one or several gene-pools. (f) Geographic distribution of plastid DNA haplotypes derived from concatenated sequences of trnGUCC–trnRUCU, rpl32–trnLUAG and trnTUGU–trnLUAA–trnFGAA intergenic spacers. (g) Statistical parsimony network of plastid DNA haplotypes. (h) Majority rule consensus tree from strict clock Bayesian analysis with the software beast. Node heights correspond to median ages. Numbers along branches are Bayesian posterior probabilities; identical haplotypes or unresolved polytomies are collapsed as triangles, their vertical extension being proportional to the number of individuals.










Clear evidence for multiple Pleistocene refugia is found not only in the high-elevation E. serpyllifolius, but also in the lowland E. tenuifolius, despite the lack of obvious dispersal barriers, in line with the refugia-within-refugia model. Genealogical relationships and genetic diversity patterns support the hypothesis that cold-adapted E. serpyllifolius responded to climatic oscillations mostly by elevational range shifts, whereas thermophilic E. tenuifolius did so mainly by latitudinal range shifts, with different phases (and probably extents) ofrange expansion. In contrast to the displacement refugia hypothesis, the two elevationally differentiated species do not differ in their genetic diversity.
  • Funding: European Community (MEIF-CT-2005024315)
  • Contributors: Harald Niklfeld, Gerald M. Schneeweiss, Peter Schönswetter, Michael Barfuss, Chris Dixon (all University of Vienna, Austria), Božo Frajman (University of Ljubljana, Slovenia), Darko Mihelj (University of Zagreb, Croatia)
  • Reference: Journal of Biogeography, 38 (7): 1381-1393.


Biogeographic Relationships Among the Mountain Ranges of Europe, Middle East and Asia: Evolution, Phylogeny and Phylogeography of the Genus Wulfenia


Both the high, geographically structured species richness in Southern Europe and the Middle East and the presence of disjunctions both within the Balkans and between the Balkans, Alps and the Middle East render the genus Wulfenia highly suitable for investigation questions concerning evolutionary patterns and speciation in Southern Europe, at the interface between phylogeny and phylogeography. Hence, our objectives are: establishment of sound molecular phylogenetic hypotheses on the relationships of the study group both to their respective closest relatives as well as within the genus; based on obtained phylogenetic hypotheses, the Pleistocene speciation hypothesis will be tested; establishment of sound comparative phylogeographic hypotheses within species and species groups; pinpointing the putative centers of origin, based on the phylogenetic and determining frequency and timing of the colonization of the Alps.

   Wulfenia orientalis var. glanduligera    Wulfenia baldaccii (photo T. Wraber).


  • Contributors: Dirk Albach (University of Oldenburg, Germany), Halil Çakan (University of Adana, Turkey), Dmitar Lakušić (University of Belgrade, Serbia), Mićo Praščević (Brezojevice, Montenegro), Marash Rakaj (University of Shkodra, Albania), Fadil Milaku (University of Prishtina, Kosovo), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: European Community (MERG-CT-2007-201204)
  • Reference: Taxon 63 (4): 843-858



Systematics and phylogeography of adriatidae


Systematics and phylogeography of Campanula pyramidalis agg. (Campanulaceae)

The Campanula pyramidalis complex is a group of closely related taxa with a distribution across the Balkans, from
the Gulf of Trieste in the north to the Peloponnese Peninsula in the south, with small disjunct parts of the range in the south
Apennines. Although 21 taxa were described within this complex, only three, C. pyramidalis, C. versicolor, and C. secundiflora,
have been generally accepted in recent synoptical taxonomic treatments. Our molecular phylogenetic analyses based
on sequences of three non-coding chloroplast regions (psbA-trnH, psbZ-trnfM, trnG-trnS) as well as of nuclear ribosomal
internal transcribed spacers (nrITS), lend strong support to the recognition of several lineages which only partially correspond
to generally accepted taxonomic concepts. Molecular data presented in this study showed that C. pyramidalis is a polyphyletic
assemblage that segregates into three distinct lineages, one of which is described here as a new species, C. austroadriatica sp.
nov. The lectotype of C. pyramidalis, redefined in a strict sense, is designated. Neither C. versicolor nor C. secundiflora were
found to be strictly monophyletic, but their monophyly could not be rejected. Morphological and biogeographical implications
are discussed.

  • Contributors: Dmitar Lakušić (University of Belgrade, Serbia), Sanja Kovačić, Zlatko Liber, Toni Nikolić (all University of Zagreb, Croatia), Saša Stefanović (University of Toronto Mississauga, Canada)
  • Funding: Serbian Ministry of Science and Technological Development (project no. 173030 to D. Lakušić), by the Ministry of Science, Education and Sport, Republic of Croatia (projects no. 119-1191193-1227 to T. Nikolić and 119-1191193-1232 to Z. Liber), and by the NSERC of Canada Discovery grants to S. Stefanović (326439).
  • Reference: Taxon 62 (3): 505-524.



Biogeography of the Dinaric tussock grasslands

Delimitation of the alliances
Caricion firmae (Seslerietalia albicantis) and Seslerion juncifoliae (Seslerietalia juncifoliae) in the Southeastern Alps and Dinaric Mountains

We performed a syntaxonomic and phytogeographic delimitation of the calcareous open sedge swards in the alpine belt of the Alps (Caricion firmae) and subalpine and alpine tussock grasslands in wind-exposed habitats (Seslerion juncifoliae) in the area of the south-eastern Alps and he Dinaric Alps. Analyses based on hierarchical classification, ordination and chorology clearly show the distinction between the syntaxa: stands from the Liburnian karst (Mt Snežnik–SW Slovenia, Mts Risnjak and Snježnik–NW Croatia), Lička Plješivica and the Velebit Mts belong to the Dinaric alliance Seslerion juncifoliae, whereas stands from the Trnovski gozd plateau (W Slovenia, north-westernmost part of the Dinaric Alps), although somewhat transitional between the two alliances, and stands from the Alps, were classified in the alliance Caricion firmae. The alliance Seslerion juncifoliae of the Dinaric Alps vicariates Caricion firmae of the Alps.
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia).
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Plant Biosystems,  139 (3): 399-410, 2005



FLORA AND VEGETATION OF THE MEDITERRANEAN


Flora and vegetation of the Kvarner Islands
(NW Adriatic)

Flora of the islands of Susak


  • Contributors: Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Branko Vreš, Valerija Babij (both Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Culture of Republic of Croatia


Flora of the island of Prvić


  • Contributors: Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Andrej Radalj (The Jezero Society, Croatia), Andrija Željko Lovrić (Sesvete, Croatia)
  • Funding: Public Institution for Managing Protected Nature Areas in County of Primorje & Gorski Kotar



Flora and vegetation of the Liburnian karst


Chasmophytes on screes? A rule and not an exception in the vegetation of the Karst (south-west Slovenia)

We investigated the community ecology and autecology of chasmophytes in the Karst (SW Slovenia). An unusually massive occurrence of chasmophytes on settled screes and rock falls was detected in some of the collapse dolines of Škocjan Caves and surroundings, building floristically and physiognomically homogenous plant communities. At first sight, no significant floristic distinctions in vascular plants were observed between scree and rock crevice stands, despite striking differences in habitats. A detailed floristic survey of stands, including vascular plants and bryophytes, as well as ecological parameters, revealed four distinct vegetation types. Floristically, bryophytes proved to differentiate groups of stands excellently but, in contrast to vascular plants, their presence and abundance only poorly reflected site conditions. Among chasmophytes, which, in terms of the Braun-Blanquet approach in phytosociology, are assigned to the class Asplenietea trichomanes (sensu Braun-Blanquet and Oberdorfer), only Saxifraga petraea and S. tridactylites prefer rock crevices and artificial walls, respectively, while others, e.g., Saxifraga crustata, Primula auricula and Athamanta turbith, prefer screes and their habitat selection is governed by different ecological parameters, shaped primarily by the phenomenon of temperature inversion in collapse dolines.
  • Contributors: Andrej Martinčič (Ljubljana, Slovenia), Igor Dakskobler (Science and Research Centre of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia).
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Plant Biosystems 146 (4): 1078-1091, 2012.


Heaths with dwarf ericaceous shrubs and Alpine juniper (Juniperus alpina) in the Dinaric Alps: A nomenclatorial and synsystematic re-appraisal

Ecology and phytosociology of north-western Dinaric heaths of the association Rhododendro hirsuti-Juniperetum alpinae Horvat ex Horvat et al. 1974 nom. corr. prop. as well as the syndynamics and synsystematics of heaths in the Dinaric Alps are discussed. While the structure (physiognomy) of these stands is very homogenous and dominated by few species, the flora is heterogeneous since ecotonal areas, where heaths are most frequently developed, represent a contact zone of elements of different syntaxa. Due to abrupt reduction in pasture activities strong encroachments of shrubs and trees have become common, which additionally contribute to floristic heterogeneity of heaths. Although the identification and circumscription together with synecology and synchorology of heaths in general are more or less easily understood and strait-forward, their floristic affinities, in relation to structure homogeneity and syndynamics, are complicated, which led to the proposal of several synsystematic schemes depending on interpretation of the relationship between flora and structure of stands. Dinaric heaths are classified into three classes, Erico-Pinetea, Vaccinio-Piceetea and Festuco-Brometea and a classification scheme is proposed together with nomenclatorial revision of the analyzed heaths with dwarf ericaceous shrubs and Alpine juniper (Juniperus alpina) in the Dinaric Alps.

  • Contributor: Željka Modrić Surina (Natural History Museum Rijeka, Croatia)
  • Funding: Public institution Priroda, Rijeka, Croatia (project no. 112-07/11-02/01–2170-52-01/1-11-19)
  • Reference: Acta Botanica Croatica 72 (1): 113-132, 2013.

Phytosociology and ecology of Dinaric fir-beech forest (Omphalodo-Fagetum) in the Trnovski gozd plateau (W Slovenia, NW Dinaric Alps)


We studied the phytosociology, ecology and biogeography of the Dinaric fir-beech stands (Omphalodo-Fagetum) in the Trnovski gozd plateau, at the north-westernmost part of the Illyrian floral province. We identified and/or confirmed two geographical variants (var. geogr. Saxifraga cuneifolia – central and western part of the plateau, and var. geogr. Calamintha grandiflora – eastern part of the plateau), and 10 floristically and ecologically well differentiated subassociations (-rhododendretosum hirsuti, -saxifragetosum cuneifoliae, -adenostyletosum glabrae, -festucetosum altissimae, -calamagrostietosum arundinaceae, -stellarietosum montanae, -seslerietosum autumnalis, -calamagrostietosum variae, -sambucetosum nigrae and -asaretosum europei). The most frequent stands belong to subassociation –festucetosum and -calamagrostietosum arundinaceae, which, in terms of site ecology and floristic composition, represent central forest types in the research area. They are floristically impoverished and lack majority of association's characteristic taxa which is in line with the biogeographic peculiarites of the research area.
  • Contributors: Igor Dakskobler, Branko Vreš (both Institute of Biology, Science and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Mitja Zupančič (Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Hacquetia 12 (1): 11-85, 2013


Forest vegetation of tectonic dolines Pihlja and Vitra above the Vinodol valley (Liburnian karst, NW Croatia)


We studied floristic composition, structure and topology of forest stands in tectonic dolines Pihlja and Vitra above Vinodol valley (Liburnian karst, NW Adriatic). Floristically and structurally homogenous stands represent zonal forests of the association Aristolochio luteae-Quercetum pubescentis (=Ostryo carpinifoliae-Quercetum pubescentis, Ostryo-Carpinion orientalis) and cover 1,75 and 2,73 ha, each (85 and 70 % of tectonic dolines, respectively). Preliminary multivariate analyses revealed several incongruences in current synsystematics and forest topology within the alliance Ostryo-Carpinion orientalis and raised a need for a thorough revision. Unsettled synsystematics makes addressing the forest vegetation zonation of the area uncertain. We assume that various stands with Carpinus orientalis in northwestern Adriatic represent only secondary succession stages in several thermophytic vegetation types. Studied forests in tectonic dolines Pihlja and Vitra represent well preserved stands without any visible traces of wood cutting and are valuable in giving insights into patterns, processes and dynamics of northern-Adriatic vegetation. As such they are in need of a special protection.
  • Funding: Public institution "Priroda"
  • Reference: Šumarski list 138 (5-6): 259-269, 2014


Flora and vegetation of the Obruč mountain range
  • Contributors: Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Andrej Radalj (The Jezero Society, Croatia)
  • Funding: Public Institution for Managing Protected Nature Areas in County of Primorje & Gorski Kotar


Flora and vegetation of Mt Učka

  • Contributors: Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Marko Randić (Public Institution for Managing Protected Nature Areas in County of Primorje & Gorski Kotar, Croatia), Marta Blažević (Učka Nature park, Croatia)
  • Funding: Učka Nature Park


Distribution area, habitat preferences and (meta)population density estimates of Campanula tommasiniana, a narrow endemic of Mt Učka (Liburnian karst, north-western Adriatic)

Campanula tommasiniana is a narrow endemic of Mt Učka occupying 6-7 km2 between 50-1396 m a.s.l. Within its distribution range, 5 more or less homogenously inhabited micro-sites were identified with the highest (meta)population density estimates on higher altitudes of the summit ridge, and lowest on lower altitudes. Majority of the population occupies rock crevices of boulders, cliffs and rocky outcrops above 900 m a.s.l., within the zonal thermophytic and altimontane and subalpine beech forests, although specimens were frequently recorded in various secondary habitats across its distribution range. Due to the lack of the data on species biology, genetic population structure and potential gene flow within and among metapopulations, it is hard to assume extents of particular threats to the species. Nevertheless, destruction of already occupied and potential habitats may represent a real threat for long-term survival of this narrow endemic with pronounced patchy distribution. Hence, every established and potential growth site, especially rocky outcrops and cliffs should be strictly preserved.

  • Contributors: Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Marko Randić (Public Institution for Managing Protected Nature Areas in County of Primorje & Gorski Kotar, Croatia)
  • Funding: Učka Nature Park
  • Reference: Natura Croatica, 22 (1), 213-222, 2013.



Phytosociology and ecology of Campanula tommasiniana (Campanulaceae), a narrow endemic of Mt Učka (Liburnian karst, NW Dinaric Alps)

Although Campanula tommasiniana is a well known narrow endemic of Mt Učka (Liburnian karst, NW Dinaric Alps), it is surprising that we know very little regarding its chorology, ecology and phytosociology. Our investigation is primarily focused on filling the gaps with elementary data in order to properly manage and protect its actual and potential growth sites.


Campanula tommasiniana


























Campanula tommasiniana
is a typical chasmophyte occupying calcareous rock crevices and cracks along the wide range of various ecological gradients demonstrating high degree of ecological plasticity and stress tolerance with regards to abiotic factors. Generally, three ecologically and floristically distinct groups of stands were recognized and typified according to sigmatistic approach: (a) Seslerio juncifoliae-Campanuletum tommasinianae ass. nov., with stands occupying higher elevated sites fully exposed to sun and strong winds; (b) Seslerio autumnalis-Campanuletum tommasinianae ass. nov., representing stands predominantly developed within the thermophytic beech stands, semi- to fully-shaded by the nearby tree canopy; (c) Cystopteri fragilis-Campanuletum tommasinianae, sciophytic, moist and cold adapted stands with high frequency and coverage of bryophytes. Results of DCA analyses using unimodal model suggest that Campanula tommasiniana is primarily a plant of open and exposed sites of higher elevation despite being most frequently found in rock crevices within the thermophytic and altimontane beech forests.
  • Contributors: Andrej Martinčič (Ljubljana, Slovenia), Željka Modrić Surina (Natural History Museum Rijeka, Croatia), Marko Randić (Public Institution for Managing Protected Nature Areas in County of Primorje and Gorski Kotar, Rijeka, Croatia)
  • Funding: Učka Nature Park
  • Reference: Acta Botanica Croatica, accepted.


Subalpine beech forest with Hairy Alpenrose (Polysticho lonchitis-Fagetum rhododendretosum hirsuti subass. nova) on Mt. Snežnik (Liburnian karst, Dinaric Alps)

We studied subalpine beech stands with Hairy Alpenrose (Rhododendron hirsutum) on Mt Snežnik (NW Dinaric Alps). They thrive on stony and steep slopes of northern exposure. Comparisons with other subalpine Beech stands (Polysticho lonchitis-Fagetum s. lat.), Fir-Beech stands with Hairy Alpenrose (Omphalodo-Fagetum s. lat. rhododendretosum hirsuti), and prealpine fir-beech stands with Hairy Alpenrose (Homogyno sylvestris-Fagetum s. lat. rhododendretosum hirsuti), stands of Hairy Alpenrose and Beech (Rhododendro hirsuti-Fagetum s. lat.), as well as Austrian subalpine beech stands (Saxifrago rotundifoliae-Fagetum s. lat.) show their unique floristical composition due to ecological conditions, and thus distinct syntaxonomical position within the association Polysticho-Fagetum. Therefore, a new subassociation Polysticho-Fagetum rhododendretosum hirsuti subass. nova was described, and – as differential species for the subassociation – we chose Rhododendron hirsutum, Rubus saxatilis, Rosa pendulina, and Clematis alpina.
  • Contributors: Marash Rakaj (University of Shkodra, Albania), Igor Dakskobler (Institute of Biology, Science and Research Center of the Slovenian Academy of Sciences and Arts)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Hacquetia, 6 (2): 89-102, 2008


Campanula waldsteiniana Shult. (Campanulaceae): a new species in the flora of Slovenia

Campanula waldsteiniana, an endemic hasmophyte of the NW Dinaric Alps, formerly known from the Velebit Mts (Croatia) and Mt Osječenica (Bosnia and Herzegovina), was discovered on Mt Snežnik (Liburnian karst) for the first time (new record for Slovenia). We gave an overview on the taxonomy and chorology of the "isophyloid" taxa of the genus Campanula and a phytosociological remark on stands from the Mt Snežnik.
  • Contributor: Sanja Kovačić (University of Zagreb, Croatia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Razprave IV. Razreda SAZU, 47 (1): 193-199, 2006


Phytosociology and ecology of Carex mucronata on the Mt Snežnik (SW Slovenia, Liburnian Karst)

We discussed the phytosociology and ecology of Carex mucronata stands on the Snežnik plateau. Results are based on numerical analyses and a comprehensive synoptic table indicating an unique syntaxonomical position of Dinaric stands in comparison to Alpine ones. The differentiation of syntaxa is due to a significant number of Illyrian species in stands from the Liburnian karst. As a result, a new association Scabioso silenifoliae-Caricetum mucronatae is described. The association is subdivided into two floristically and ecologically distinct subassociations: -edraianthetosum graminifolii, restricted to the summit of Mt Snežnik, and -rhododendretosum hirsuti on lower altitudes. The association is assigned to the Dinaric alliance Seslerion juncifoliae and it is restricted to the north-western part of the Liburnian karst (Mts Snežnik - SW Slovenia, Snježnik and Risnjak - NW Croatia).
Southern slopes of Mt Snežnik (Liburnian karst, NW Dinaric Alps) as seen from Mirin above the Velika Padežnica doline.

  • Contributor: Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Wulfenia, 12: 97-112


Paeonio officinalis-Tilietum platyphylli - new association of broad leaf forest stands in the Čičarija (SW Slovenia)

  • Contributors: Petra Košir, Igor Dakskobler (both Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Meje in konfini, (Rožac Darovec, V., ed.), 345-366, Koper, Založba Annales, 2005


Some novelties in the flora and vegetation of Mt Snežnik (SW Slovenia, Liburnian karst)

The first records for Scabiosa graminifolia, Potentilla clusiana and Seseli gouanii were presented for the Snežnik Plateau. One new association, Scabioso silenifoliae-Dryadetum octopetalae ass. nova, and a phytocoenon with Scabiosa silenifolia were described.

Due to the distinct geographical position of Mt Snežnik (1796 m), the highest peak of the Liburnian karst and NW Dinaric Alps, its flora, which is composed of Alpine and Dinaric/Illyrian species, is comparatively well known. The first botanist who climbed the mountain and pointed out the floristic and phytogeographyic peculiarities of the area was Henrik Freyer in 1827. As an extremely good mountaineer, he walked the distance between the cities of Idrija and Ilirska Bistrica (at that time known as Trnovo; Slovenia), round 80 km, in one day (16.6.1827). The day after, as the first botanist ever, he climbed the summit of Mt Snežnik and found several peculiar plant taxa. However, it was his finding of Campanula graminifolia (now known as Edraianthus graminifolius), a typical Balkan endemic with a disjunction on the Apennines and Sicily, which raised a lot of attention to the botanists afterwards and since then the biogeography of the flora became a trademark of this floristically renowned mountain.



Edraianthus graminifolius, a Balkan endemic with a disjunct distribution on the Apennines, Sicily and Carpathian Mts.

  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Acta Botanica Croatica, 64 (2): 341-356, 2005


The association Doronico austriaci-Adenostyletum alliariae Horvat ex Horvat et al. 1974 on the Mt. Snežnik (Liburnian karst, NW Dinaric Alps)

We studied phytosociology and ecology of the tall-herb vegetation with Adenostyles alliariae and Doronicum austriacum on Mt Snežnik and adjecent area. Respective stands occurred on moist, nutrient rich soils and cooler growth sites, shady slopes, and moist depressions. In the subalpine belt they thirve just above the tree line, within the stands of dwarf mountain pine (Hyperico grisebachii-Pinetum mugo var. geogr. Arabis scopoliana)., subalpine beech forests (Polysticho-Fagetum var. geogr. Allium victorialis), Dinaric fir-beech (Omphalodo-Fagetum var. geogr. Calamintha grandiflora), and spruce stands (Lonicero caeruleae-Piceetum). On altimontane belt they most frequently throve in freezing ravines as a consequence of vegetation inversion. Stands were placed into the association Doronico austriaci-Adenostyletum alliariae.
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Razprave IV. Razreda SAZU, 46 (2):145-160, 2005


Phytogeographical differentiation in the Dinaric fir-beech forest (Omphalodo-Fagetum s.lat.) of the western part of the Illyrian floral province

A phytogeographical assessment of the west-Dinaric fir-beech association Omphalodo-Fagetum s. lat. (Treg. 1957 corr. Puncer 1980) Marinček et al. 1993 (=Abieti-Fagetum dinaricum Tregubov 1957 p.p.) in the western part of the Illyrian floral province was provided. The synoptic phytosociological table and the results of hierarchical classification and ordination show a quite distinct floristic and phytogeographical pattern of differentiation in a northwest-southeast direction. The proportion of Southeast – European-Illyrian (Illyricoid) species decreases towards the northwest while Alpine species decrease in the opposite direction. Stands from the Trnovski gozd plateau form a geographical variant Omphalodo-Fagetum var. geogr. Saxifraga cuneifolia, further divided into two geographical sub-variants, western – subvar. geogr. Anemone trifolia and central-eastern – subvar. geogr. Omphalodes verna. All other stands belong to the geographical variant Omphalodo- Fagetum var. geogr. Calamintha grandiflora, also further divided into two geographical sub-variants, subvar. geogr. Dentaria pentaphyllos and subvar. geogr. Dentaria polyphylla. Although the distribution area of the association is well defined in its northwest part, the question of its south-eastern border is still open. The preliminary results of analyses indicate that the stands southeast of Velebit and Lička Plješivica should be treated as a different association, distinct from Omphalodo-Fagetum. Further phytosociological and phytogeographical research into the entire Illyrian floral province will therefore be needed to resolve the problem of the southern border of the distribution area of the association Omphalodo-Fagetum and the syntaxonomy of south-eastern Dinaric fir-beech stands.
  • Contributors: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Mitja Zupančič (Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Acta Botanica Croatica, 61 (2): 145-178, 2002


Two Pre-pannonian beech associations in northeastern Slovenia

  • Contributors: Mitja Zupančič (Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Vinko Žagar (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Razprave IV. Razreda SAZU, 41 (2): 179-248, 2000




Vegetation patterns and ecology of dolines in Dinaric Alps - a phenomenon of vegetation inversion


Studies on the ecology and vegetation of freezing ravines in the Dinaric Alps started already in the 19th century. Krašan was the first who pointed out interesting vegetation patterns (namely, dwarf pine Pinus mugo at the bottom of the karst dolina within the Dinaric fir-beech stand) in Smrekova draga valley. Subsequently, Beck tried to explain this phenomenon on the case of the Paradana icehollow and Smrekova draga doline (north-western part of the Trnovski gozd plateau, NW Dinaric Alps), but the most detailed research on ecology and vegetation of freezing ravines was performed by Martinčič. He studied the ecology and vegetation patterns of some dolines in the Trnovski gozd plateau: Paradana ice-hollow, Smrekova draga, dolina in Poslušanje, dolina SW from Mt Bukov vrh (north-western part of the plateau), and Ožgani grič valley (southern part of the plateau on the border between the sub-Mediterranean and Dinaric phytogeographical region); while on Mt Snežnik he thoroughly studied the ecology and vegetation patterns in the doline of Stanišče. Actually, vegetation studies in Ožgani grič had already been done by Piskernik, who recognised three “climo-coenoses”: Hylocomio splendentis-Moehringietum muscosae (the coldest growth sites with no woody species), Salico appendiculatae-Moehringietum muscosae (shrub stands with Salix appendiculata), and Piceo excelsae-Moehringietum muscosae (forest stands with Picea abies). Martinčič classified freezing ravines in the Trnovski gozd as follows: freezing ravines with screes with natural spruce (Picea abies) stands (Ožgani grič, Poslušanje), with stands of dwarf-pine (Smrekova draga, dolina SW from Mt. Bukov vrh), and reezing ravines with permanent temperature inversion (Paradana ice-hollow). There were also extensive phytosociological studies of spruce forests in the Dinaric Alps which were related to the research topic. On the Trnovski gozd plateau and Mt Snežnik, Zupančič studied spruce stands in Smrekova draga (Lonicero caeruleae-Piceetum), Smrečje, Stanišče, Gregorjev dolec, Velika Padežnica, Mala Padežnica, Kujavič, Velika and Mala Lazna valleys (Stellario montanae-Piceetum). Spruce forests occur in the Dinaric Alps only azonally, since they are more or less restricted to freezing ravines or cold, moist, and shaded sites. During our research on the ecology of Heliosperma pusillum in the Dinaric Alps (Surina & Vreš 2004), as well as flora and vegetation of the Trnovski gozd plateau, we observed very distinct and homogeneous stands with dominating Heliosperma pusillum and Sanionia uncinata (=Drepanocladus uncinatus) in freezing ravines on moist, cold and shadowy screes of boulders of periglacial origin, most commonly, with long-lasting snow cover. Ecologically and/or floristically very similar stands were already observed and thoroughly studied on Mt Snežnik, as well as the Trnovski gozd plateau. However, detailed phytosociological observations were still missing. Our studies are focused in filling the gap in understanding the ecology and vegetation patterns of this phenomenon.

Vegetation inversion in the karst doline Velika Kolobarnica (bellow the Karolinov vrh peak) as seen from the summit of Mt Snežnik (1796 m), the highest mountain of the Liburnian karst (NW Dinaric Alps)
. The highest vegetation belt of the ridge represents stands of the association Hyperico grisebachii-Pinetum mugo, which occur zonally above the subalpine beech stands (Polysticho lonchitis-Fagetum). However, bellow the subalpine beech stands, stands with Pinus mugo occur again, this time extrazonally, as a consequence of temperature inversion. Further towards the bottom, Dinaric stands with Festuca bosniaca prevail and at the very bottom (in shade), fragmented stands of the association Edraiantho graminifolii-Caricetum firmae cover the lowest and thus the coldest parts of the doline.



Snowbed vegetation in Croatia: phytosociology, ecology and conservation status


We studied phytosociology, ecology and chorology of snowbed vegetation in Croatia. Snowbed stands, found only in freezing ravines and dolines of the Liburnian karst and Velebit Mountains (NW Dinaric Alps, NW Croatia), we classified into the association Drepanoclado-Heliospermetum (Salicion retusae, Arabidetalia caeruleae, Thlaspietea rotundifolii). Those stands, exposed mostly to the north and shaded for the majority of the growing season, were developed on more or less settled periglacial screes and boulders in the coldest parts (bottoms, smaller ditches) of the dolines with long-lasting snow cover. Due to lower altitude of Croatian mountains, snowbed vegetation could be found only azonally, in freezing ravines surrounded by altimontane and subalpine (fir-)beech and spruce forests, where these stands manage to thrive due to specific microclimatic conditions. Snowbeds host some rare, endangered and/or protected plant species in Croatia. We briefly disscussed vulnerability of the flora and vegetation of snowbeds in Croatia.
  • Contributor: Željka Modrić Surina (Natural History Museum Rijeka, Croatia)
  • Ministry of Higher Education, Science and Technology of the Republic of Slovenia, Ministry of Culture of Republic of Croatia
  • Reference: Plant Biosystems, 144 (4): 747-768.


Cyrtomnium hymenophylloides (Huebener) T.J.Kop.

During the botanical trip to notrhern Velebit Mts, Rožanski kukovi area, while investigating snow beds and hunting the rare Saxifraga prenja (S. sedoides agg.), we encountered a rather rare moss species - Cyrtomnium hymenophylloides (the first record for Croatia), in a stand belonging to the association Drepanoclado uncinati-Heliospermetum pusilli.
  • Contributors: Marko Sabovljević, Dmitar Lakušić (both University of Belgrade, Serbia), Antun Alegro, Vedran Šegota (both University of Zagreb)
  • Funding: Ministry of Culture of Republic of Croatia
  • Reference: Journal of Bryology, 31: 203, 2009


The association Drepanoclado uncinati-Heliospermetum pusilli (Arabidetalia caeruleae, Thlaspietea rotundifolii) in the Trnovski gozd plateau (Slovenia, NW Dinaric Alps)

We studied the phytosociology and ecology of Heliosperma pusillum in freezing ravines of the Trnovski gozd plateau (Slovenia, NW Dinaric Alps). The species thrives on shadowy, moist, cold and stable screes of boulders with long-lasting snow cover. The stands belong to the association Drepanoclado uncinati-Heliospermetum pusilliPaederota lutea was described. Differential species for the geographical variant are Phyteuma scheuchzericolumnae, Valeriana saxatilis, Rhodothamnus chamaecistus, and Saxifraga cuneifolia. For less stable screes with smaller rocky particles a new subassociation salicetosum retusae was described, and the differential species for the subassociation are Salix retusa and Poa alpina. Stands of the association Drepanoclado-Heliospermetum var. geogr. Paederota lutea from the Trnovski gozd plateau are on the north-westernmost part of the distribution area of the Dinaric alliance Salicion retusae.
(Salicion retusae, Arabidetalia caeruleae, Thlaspietea rotundifolii).


Due to close proximity of the Julian Alps, many SE Alpine geoelements with high biogeographic diagnostic value occur in the various vegetation types in the NW Dinaric Alps (the Trnovski gozd plateau), e.g., Rhodothamnus chamaecistus, Phyteuma scheuchzeri subsp. columnae, and Campanula zoysii.
  • Contributors: Branko Vreš, Igor Dakskobler (both Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Hacquetia, 8 (1): 31-40, 2007


Phytosociological characteristics of sites of Heliosperma pusillum (=Silene pusilla, Caryophyllaceae) in freezing ravines on the Snežnik plateau (SW Slovenia)


We studied the phytosociological and environmental conditions of the stands of Heliosperma pusillumDrepanoclado uncinati-Heliospermetum pusilli. It's stands thrive on moist screes in the coldest parts of freezing ravines. The key environmental factor which conditions both the growth of the stands of this association and the vegetation inversion is a very low temperature of soil and the air layer just above it, which is confirmed by the temperature measurements on selected profile of the freezing ravine.
Stands of the association Drepanoclado uncinati-Heliospermetum pusilli at the bottom of the the freezing ravine (doline) Stanišče (Mt Snežnik, Liburnian karst, NW Dinaric Alps). Heliosperma pusillum (Caryophyllaceae) in full flower and a moss species Drepanocladus uncinatus (=Sanionia uncinata; light green) completely dominate in the stands usually developed on the periglacial material. Other frequent phanerogams are Viola biflora, Carex capillaris, Chrysosplenium alternifolium, Carex atrata, Polygonum viviparum and Festuca nitida, while among mosses the most common species are Oncophorus virens, Campylium stellatum, Polytrichum alpinum, Pohlia elongata subsp. elongata and Orthothecium rufescens. Shrub layer (e.g., Picea abies, Rhododendron hirsutum, Salix appendiculata) is due to extreme environmental conditions not (or only scarcely) developed.
  • Contributor: Branko Vreš (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Razprave IV. Razreda SAZU, 45 (2): 147-183, 2004



Flora and Vegetation of the Julian Alps


Phytogeography and syntaxonomy of snow-bed vegetation on calcareous substrates in the south-eastern Alps: a numerical approach

Based on cluster analysis and the phytogeographic peculiarities of the area, we provided an assessment of the vegetation of snow-beds on calcareous soils (Arabidetalia caeruleae) in the South-eastern Alps. Eight distinct associations belonging to three alliances were recognised: Saxifragetum stellaro-sedoidis, Saxifragetum hohenwartii and Ranunculo traunfellneri-Festucetum nitidae (alliance Arabidion caeruleae), Salici herbaceae-Arabidetum caeruleae (alliance Salici herbaceae-Arabidion caeruleae), Salicetum retuso-reticulatae, Homogyno discoloris-Salicetum retusae, Salici retusae-Geranietum argentei and Potentillo brauneanae-Homogynetum discoloris (alliance Soldanello alpinae-Salicion retusae). Saxifragetum stellaro-sedoidis was further subdivided into three geographical variants, such as variants of Achillea oxyloba, Ranunculus traunfellneri and Campanula pulla. The Drepanoclado uncinati-Heliospermetum pusilli from the Liburnian Karst (Dinaric Alps) show clear floristic and phytogeographic distinctions and its placement into the Dinaric alliance Salicion retusae was confirmed.
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Annales, Series Historia Naturalis, 15 (2): 223-238, 2005


Subalpine and alpine vegetation in the Krn area in the Julian Alps

Applying the sigmatistic (Central European) phytosociological method, we researched the subalpine and alpine vegetation of the Krn Mts in the Julian Alps. The following associations from the class Asplenietea trichomanis and order Potentilletalia caulescentis were identified: Valeriano elongatae-Asplenietum viridis, Ranunculo traunfellneri-Paederotetum luteae ass. nova (alliance Cystopterydion), Potentillo clusianae-Campanuletum zoysii, Campanulo carnicae-Moehringietum villosae, Paederoto luteae-Minuartietum rupestris, Saxifragetum squarroso-crustatae ass. nova and Potentilletum nitidae (Androsaci-Drabion tomentosae). Scree vegetation (Thlaspietea rotundifolii) belongs to orders Arabidetalia caeruleae and Thlaspietalia rotundifolii. The associations from the first order are Saxifragetum stellaro-sedoidisHomogyno discoloris-Salicetum retusae, phytocoenon with Salix alpina and Salici retusae-Geranietum argentei ass. nova (Soldanello alpinae-Salicion retusae), while the second order includes Papaveri kerneri- Thlaspietum kerneri, Doronicetum grandiflori (Thlaspion rotundifolii), Festucetum laxae, Dryopteridetum villarii and Aconito ranunculifolii-Adenostyletum glabrae ass. nova (Petasition paradoxi). Most of the syntaxa from the alpine and subalpine calcareous grasslands (Elyno-Seslerietea) belong to the order Seslerietalia caeruleae, e.g. Avenastro parlatorei-Festucetum calvae, Ranunculo hybridi-Caricetum sempervirentis (Caricion austroalpinae), Caricetum ferrugineae (Caricion ferrugineae), Gentiano terglouensis-Caricetum firmae, Dryadetum octopetalae and Caricetum mucronataeRhododendretum hirsuti (Rhododendro-Ericion, Rhododendro hirsuti-Ericetalia carneae), whereas those occurring on more acid soils were placed in the association Empetro-Vaccinietum gaultherioidisSalicetum waldsteinianae (Alnion viridis, Adenostyletalia alliariae), while in the vegetation of cold calcareous springs (Montio-Cardaminetea, Montio-Cardaminetalia, Cratoneurion commutati) stands of the association Cratoneuretum falcati were identified. Due to the floristic peculiarities of the research area, a phytogeographic assessment of the syntaxa was also made, where several new geographical variants were described. (Loiseleurio-Vaccinion, Loiseleurio-Vaccinietalia, Loiseleurio-Vaccinietea). From the class Mulgedio-Aconitetea we identified stands of the association (Arabidon caeruleae), (Caricion firmae). Dwarf shrub-communities on calcareous soils were classified into the association Rhododendretum hirsuti (Rhododendro-Ericion, Rhododendro hirsuti-Ericetalia carneae), whereas those occurring on more acid soils were placed in the association Empetro-Vaccinietum gaultherioidis (Loiseleurio-Vaccinion, Loiseleurio-Vaccinietalia, Loiseleurio-Vaccinietea). From the class Mulgedio-Aconitetea we identified stands of the association Salicetum waldsteinianae (Alnion viridis, Adenostyletalia alliariae), while in the vegetation of cold calcareous springs (Montio-Cardaminetea, Montio-Cardaminetalia, Cratoneurion commutati) stands of the association Cratoneuretum falcati were identified. Due to the floristic peculiarities of the research area, a phytogeographic assessment of the syntaxa was also made, where several new geographical variants were described.
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Scopolia, 57: 1-222, 2005


Ranunculo traunfellneri-Paederotetum luteae: new rock crevices association from the Julian Alps (South-Eastern calcareous Alps)

Vegetation of rock crevices in the Krn Mts. (the Julian Alps) is briefly presented. The association Ranunculo traunfellneri-Paederotetum luteaeValeriano elongatae-Asplenietum viridis var. geogr. Campanula zoysiiPotentillo clusianae-Campanuletum zoysii, Campanulo carnicae-Moehringietum villosae, Paederoto luteae-Minuartietum rupestris, Saxifragetum squarroso-crustatae and Potentilletum nitidae (Androsaci-Drabion tomentosae). ass. nova was newly described. We classified it into the alliance Cystopteridion fragilis.

Ranunculus traunfellneri, an endemic restricted to SE Calcareous Alps with a disjunction on Mt Snežnik (Liburnian karst, Dinaric Alps).
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Acta Biologica Slovenica, 48 (2): 3-13, 2005


Contribution to the spring vegetation of the Julian Alps: The association Cratoneuretum falcati Gams 1927

We studied phytosociological characteristics of stands with predominating Cratoneron commutatum var. falcatum (=Palustriella commutata var. falcata) in the Krn Mts and Mt Mangart (the Julian Alps). After the comparison with similar stands from the Karavanke Mts we ranged studied stands into the association Cratoneuretum falcati (Cratoneurion commutati, Montio-Cardaminetea).
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Hacquetia, 4 (1): 53-59, 2005


Carex austroalpina Becherer, a new southeastern-Alpine taxon for the flora of Slovenia, and Viola pyrenaica Ramond ex DC., second record for the flora of the Julian Alps

Carex austroalpina (Carex ferruginea subsp. austroalpina) and Viola pyrenaica are reported from the Krn Mts in the Julian Alps (Southeastern Calcareous Alps). The two localities for C. austroalpina in the Krn Mts are at the easternmost range of species distribution area. The record of Viola pyrenaica in the Krn Mts is the second for the flora of Slovenia (the Julian Alps) after nearly half a century. The phytosociological characteristics of the sites are given. Both findings are of significant importance for further phytogeographical assessment of the Southeastern Calcareous Alps.

  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Annales, Series Historia Naturalis, 14 (1): 231-236, 2004


The association Gentiano terglouensis-Caricetum firmae T. Wraber 1970 in the Krn mountains (the Julian Alps)

The syntaxonomy and ecology of the Carex firma stands in the Krn Mts (eastern Julian Alps) are presented by applying the sigmatistic method and cluster analysis. The reserached stands are classified in the association Gentiano terglouensis-Caricetum firmae T. Wraber 1970, and are further divided into three subassociations: -dryadetosum octopetalae Poldini & Feoli 1976, -drepanocladetosum uncinati subass. nova, and -potentilletosum nitidae subass. nova.
  • Contributor: Igor Dakskobler (Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Annales, Series Historia Naturalis, 14 (1): 99-112, 2004



New localities of the spur pansy (Viola cornuta L.) in the Julian Alps

In 2002 three new localities of the Spur Pansy (Viola cornuta L.) in the Julian Alps were found. They are all located in the vicinity of the lake Krnsko jezero, not far from the Mt Veliki Lemež, the only already known and presumable indigenous locality of Spur Pansy in Slovenia. After the surprising find of Spur Pansy on the Mt Veliki Lemež eleven years before the findings of our's, T. Wraber was the first who discussed in detail about the possibility of its indigenous occurence in the Julian Alps. After thorough reflection he left the problem still open. New localities are situated east of the Mt. Veliki Lemež, namely about 100 m northeast of the lake Krnsko jezero, 1400 m a.s.l. beneath the gentle rise named Glava, on the southern slope of the Škrbina pass between the Mt Debeljak and Mt Veliki Lemež, 1450 m a.s.l., and on the western slope of the Mt Debeljak above the Škrbina saddle, 1710 m a.s.l. The occurance of the Spur Pansy on the locality 2 suits well to the already known facts about its growth site ecology, since the species prefers nitrophilous situation. Spur Pansy, growing in the slight depression, totaly prevails together with accompanied flora, e.g. Urtica dioica, Deschampsia caespitosa and Athyrium filix-femina. Therefore the occurance of the Spur Pansy on the locality 2 show clear evidence of mountain sheep and goat pasture activity and is though, as we assume, a result of a human activity. On the other hand, the occurances of the Spur Pansy on localities 3 & 4 give the impression of completely natural occurance. On the locality 3 it is homogeneously disposed within the Festuca calva stand in the association Avenastro parlatorei-Festucetum calvae. The growth site No 4, assuming its indigenous status in the Julian Alps, looks even more convincing, since the plant thrives very well and it is homogeneously disposed in the stands of Rusted Sedge (Caricetum ferrugineae s. lat.). Still more, the growth site is not approachable without easier rock-climbing. The numerical classification of the phytosociological relevés, taken on the afore mentioned growth sites, showed that the relevé from the locality 4 suits, together with the relevé from the locality 1 and other similar stands from the Julian Alps, to the association Caricetum ferrugineae Lüdi 1921 s. lat. and the stand from the locality 3 to Avenastro parlatorei-Festucetum calvae (Aichinger 1933 corr. Franz 1980) Poldini & Feoli Chiapella in Feoli Chiapella & Poldini 1993 var. Genista radiata prov. Nevertheless, serious scruples about the indigenous origin of the Spur Pansy in the Julian Alps still exist. Namely, during the 1st world war the area represented the front line between the Austro-Hungarian and Italian army and was therefore well attended. According to the historical data, in the period from May 1915 till March 1916, only in the vicinity of Mt. V. Lemež the whole Austro-Hungarian battalion (around 800 soldiers) was situated. There were also several cableways conecting the surrounding of the lake Krnsko jezero and neighbouring summits supplying the army posts. Therefore the omnipresence of soldiers at the same place increase the possibility of incidental transportation or even plantation. Inspite new findings the phenomenon of its indigenous origin in the Julian Alps is still not solved.

 

Spur Pansy (Viola cornuta L.).



Austro-Hungarian soldiers from the 1st World War by the aerial ropeway on the Jastrovec in the Upper Soča Valley (photo archives David Erik Pipan).

Known (1) and new localities (2-4) of the Spur Pansy (Viola cornuta L.) in the Krn area on the  photography taken from the Batogniška škrbina.

  • Contributors: Branko Vreš, Igor Dakskobler (both Institute of Biology, Scientific and Research Center of the Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia), Tone Wraber (Polhov Gradec, Slovenia)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Razprave IV. Razreda SAZU, 44 (2): 87-102, 2003


POLLINATION ECOLOGY


Foraging behaviour of the bee Osmia apicata Smith, 1853 (Hymenoptera: Megachilidae)

With observations and photography, the collecting of Onosma echioides (Boraginaceae) pollen and nectar by Osmia apicata Smith, 1853 bees was studied. Pollen is collected by shaking the anther cone using the legs, an alternative method to the buzzing in this oligolectic bee species.

  • Contributors: Andrej Gogala (Slovenian Museum of Natural History, Ljubljana, Slovenia)
  • Reference: Acta Entomologica Slovenica, 19 (2): 139-149, 2011
Osmia apicata on Onosma echioides. Photo: Andrej Gogala


STUDIES OF CRYPTOGAMS

Tortella densa (Lorentz & Moledno) Crundwell & Nyholm: the first record for Croatia


Tortella densa, a species with a disjunctive Holarctic distribution, and considered to be a temperate element in Europe, is recorded for the first time in Croatia. Several specimens of T. densa were found in crevices of vertical boulders and cliffs of limestone rock, among four stands of a chasmophytic community including the narrow endemic Campanula tommasiniana K.Koch. Other frequent and rather interesting phanerogams in stands, covering 20–40% of the releve´ area, included Sesleria tenuifolia Schrad., Athamanta turbith Brot., Leontopodium alpinum Cass., Globularia cordifolia L., Arabis scopoliana Boiss., Silene saxifraga L., etc. In these stands, bryophytes were only poorly represented and covered around 1% (and up to 5%) of the releve´ area. Besides Tortella densa, other mosses recorded included Encalypta vulgaris Hedw., Homalothecium philippeanum (Spruce) Schimp., H. sericeum (Hedw.) Schimp., Hypnum cupressiforme Hedw. var. cupressiforme, Tortella tortuosa (Hedw.) Limpr., T. nitida (Lindb.) Broth., and Weissia sp. Here, Tortella densa preferred open and sunny sites between 1180 and 1265 m a.s.l., and was exposed to the Bora wind regardless of its position. In the Balkan Peninsula and elsewhere in SE Europe, Tortella densa is known only from Slovenia, Serbia, Romania and Greece. In neighbouring Slovenia, it occurs in south-eastern Calcareous Alps, pre-Alps and Dinaric Alps between 500–2100 m a.s.l.

  • Contributors: Andrej Martinčič (Ljubljana, Slovenia)
  • Funding: Učka Nature Park
  • Reference: Journal of Bryology 34 (2): 131, 2012


Cyrtomnium hymenophylloides (Mniaceae): the first record for Croatia

Cyrtomnium hymenophylloides, an easily recognized, rare, northern hemisphere moss of the Mniaceae was recorded for the first time in Croatia (Dinaric Alps, Velebit Mts, Rožanski kukovi peaks area). The species has a scattered circumboreal distribution with an interesting arctic-alpine disjunction. It is a relict species already recognized from fossil material in the latest Pliocene some four million years ago. It was found in a snowbed scree with long-lasting snow cover at the bottom of a f reezing ravine. The species was recorded sterile, which is expected owing to its dioecous condition and the distinctly different distribution of male and female plants throughout its range. Female plants are widely distributed, while male plants are restricted to the far northern fringe of its range where they survived glaciations. Even there, sporophytes and male plants are rare. Cyrtomnium hymenophylloides was recorded in stands of the association Drepanoclado uncinati-Heliospermetum pusillae (alliance Salicion retusae) with Saxifraga sedoides L. subsp. prenja (Beck) Beck dominant in a herb layer; other frequent and rather frigoriphilous taxa in the stand are Heliosperma pusillum, Poly gonum viviparum L., Poa alpina and Festuca nitida among the phanerogams, and Sanionia uncinata, Campylium stellatum and Orthothecium rufescens among the cryptogams.


Cyrtomnium hymenophylloides (photo: Dmitar Lakušić).

  • Contributors: Marko Sabovljević and Dmitar Lakušić (both University of Belgrade, Serbia)
  • Funding: Ministry of Culture of Republic of Croatia
  • Reference: Journal of Bryology 31: 203, 2009



Contribution s to the lichen flora of Slovenia X.: Lichens from the Slovenian Julian Alps

The epiphytic lichen flora of the Slovenian Julian Alps, with special regard to the Triglav National Park, is discussed. The following taxa are reported for the first time for Slovenia: Bacidia incompta, Bacidina delicata, Candelariella kuusamoensis, Cladonia norvegica, C. ramulosa, Diplotomma alboatrum, Gyalecta ulmi, Hyperphyscia adglutinata, Lecanora boligera, Lecidea betulicola, L. plebjea, Micarea cinerea, M. misella, Thelomma ocellatum and Usnea barbata. The geography, geology, climate and vegetation of the area are described briefly.
  • Contributors: Franc Batič, Tadeja Trošt (both University of Ljubljana, Slovenia), Katja Primožič (Koper, Slovenia), Helmut Mayrhofer (University of Graz, Austria)
  • Reference: Herzogia, 16: 143-154, 2003



The lichens of the Ždrocle forest reserve

88 lichenized and 2 lichenicolous fungi are reported from the Forest Reserve Ždrocle (Mt Snežnik, Liburnian karst, SW Slovenia). The association Ranunculo platanifolii-Fagetum var. geogr. Calamintha grandiflora hosts the greatest biodiversity of lichens. One species (Biatora flavopunctata) is new for the flora of Slovenia, two species (Collema furfuraceum and Lecanora subintricata) and one variety (Cladonia macilenta ssp. floerkeana) are new for the Dinaric phytogeographical region.


One of the sinkholes (doline) in the Ždroclje area (Snežnik plateau).
























  • Contributors: Johannes Prügger, Helmut Mayrhofer (both University of Graz, Austria)
  • Funding: Ministry of Higher Education, Science and Technology of the Republic of Slovenia
  • Reference: Zbornik gozdarstva in lesarstva 63: 7-25, 2000
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