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Karst as a geological phenomenon and karst poljes as the largest depressions in the Dinaric Alps represent one of the most important and most diverse relief units globally. The diversity of karst poljes in the Dinaric Alps is one of the most significant worldwide and one of the most heterogeneous, too. In this study, by using the most relevant variables and delimitation criteria, all recognized Dinaric karst poljes were divided and classified into 30 well differentiated groups and organized in nine main biogeographical regions. During the process of classifying the Dinaric karst poljes, for some of the most important parameters recognizable patterns were defined or established for each group. The mycobiota of 10 Dinaric karst poljes classified into six groups have been intensively studied in Bosnia and Herzegovina and Croatia during various research sessions. All results were evaluated by using standardized variables and a corresponding binary grading scale in order to classify poljes according to their importance for mycobiota. Based on the “cumulative ecological score” (CES), the ratio of the CES/number of all recorded species and the CES/number of research days ratio, the mycologically most important karst poljes were identified. A brief description of specific habitat types of top evaluated poljes is given together with recommendations for the implementation of proper conservation measures in these areas. Several rare and very vulnerable species of ascomycetes, like Mollisia uda, Patinella hyalophaea, Podostroma leucopus, Scutellinia peloponnesiaca and Lamprospora leptodictya, were recorded in the investigated poljes. Some of the species which were found during the present study, represent the only known records in the Dinaric Alps and their conservation status has never been assessed before. Recommendations for the protection of these species, based on IUCN criteria, are presented.
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DINARIC
KARST POLJES
NATURE CONSERVATION
AND RURAL DEVELOPMENT
Edited by:
Peter Sackl,
Stefan Ferger,
Nermina Sarajlić,
Dražen Kotrošan
& Goran Topić
Sarajevo, 2019
Contents
I. Preface
Dinaric karst poljes – Jewels of the Western Balkans
Tobias Salathé
Dinaric karst poljes – a look into the future
Dražen Kotrošan & Nermina Sarajlić
II. Karst poljes as wetlands of national and international importance - Worskhop
proceedings and projects results
Spatial protection of Livanjsko polje in the framework of the UNEP/GEF project: Achieving
biodiversity conservation through creation, effective management and spatial designation of
protected areas and capacity building
Maja Jaćimovska
The role of local breeds for the preservation of the ecosystems of karst pasture areas
Gordan Šubara, Ida Štoka & Ante Ivanković
Dinaric karst poljes and their importance for mycobiota
Neven Matočec, Nedim Jukić, Nihad Omerović & Ivana Kušan
Birds of Pešter karst polje
Slobodan Puzović, Vladan Vučković, Nikola Stojnić, Goran Sekulić, Miloš Radaković, Nenad Dučić,
Brano Rudić, Milan Ružić, Dimitrije Radišić, Bratislav Grubač, Marko Šćiban & Marko Raković
Results of two years of research of the bird fauna of Popovo polje
Aleksandar Vukanović
Analysis of the occurrence of Lesser Kestrel (Falco naumanni) and Red-footed Falcon
(Falco vespertinus) in the karst poljes of Bosnia and Herzegovina in the 2012-2017 period
Goran Topić, Biljana Topić, Dražen Kotrošan, Mirko Šarac & Josip Vekić
Northern Lapwing (Vanellus vanellus) in the karst poljes of Bosnia and Herzegovina
Dražen Kotrošan, Goran Topić, Mirko Šarac, Josip Vekić & Nermina Sarajlić
Karst poljes in Bosnia and Herzegovina – newly identified Important Bird Areas
Borut Rubinić
III. Dossier of karst poljes of Bosnia and Herzegovina meeting the criteria
to be identified as Important Bird Areas (IBAs)
List of karst poljes in Bosnia and Herzegovina proposed as new Important Bird Areas
Borut Rubinić, Jovica Sjeničić, Peter Sackl, Dražen Kotrošan & Nermina Sarajlić
5
6
9
17
27
51
87
93
113
123
143
26 DINARIC KARST POLJES
Helvella corium (Photo: Neven Matočec)
27
NATURE CONSERVATION AND RURAL DEVELOPMENT
Dinaric karst poljes and their importance for mycobiota
Neven Matočec1, Nedim Jukić2, Nihad Omerović2 & Ivana Kušan1
1 Ruđer Bošković Institute,
Bijenička cesta 54, 10000 Zagreb, Croatia;
E-mail: nmatocec@irb.hr
2 Mycological Association MycoBH,
Trg Zlatnih ljiljana 34, 71000 Sarajevo, Bosnia and Herzegovina
Summary
Karst as a geological phenomenon and karst
poljes as the largest depressions in the Dinaric
Alps represent one of the most important and
most diverse relief units globally. The diversity
of karst poljes in the Dinaric Alps is one of the
most significant worldwide and one of the most
heterogeneous, too.
In this study, by using the most relevant variables
and delimitation criteria, all recognized Dinaric
karst poljes were divided and classified into 30
well differentiated groups and organized in nine
main biogeographical regions. During the process
of classifying the Dinaric karst poljes, for some
of the most important parameters recognizable
patterns were defined or established for each
group. The mycobiota of 10 Dinaric karst poljes
classified into six groups have been intensively
studied in Bosnia and Herzegovina and Croatia
during various research sessions. All results were
evaluated by using standardized variables and a
corresponding binary grading scale in order to
classify poljes according to their importance for
mycobiota. Based on the “cumulative ecological
score” (CES), the ratio of the CES/number of all
recorded species and the CES/number of research
days ratio, the mycologically most important
karst poljes were identified. A brief description
of specific habitat types of top evaluated poljes
is given together with recommendations for the
implementation of proper conservation measures
in these areas.
Several rare and very vulnerable species
of ascomycetes, like Mollisia uda, Patinella
hyalophaea, Podostroma leucopus, Scutellinia
peloponnesiaca and Lamprospora leptodictya,
were recorded in the investigated poljes. Some of
the species which were found during the present
study, represent the only known records in the
Dinaric Alps and their conservation status has
never been assessed before. Recommendations
for the protection of these species, based on
IUCN criteria, are presented.
Keywords: Ascomycota, classification, conserva-
tion, ecological score, fungi, Bosnia and Herze-
govina, Croatia
Sažetak
Krš kao geološki fenomen, zajedno s krškim polji-
ma koja predstavljaju najveće depresije na povr-
šini Dinarida, globalno predstavlja jednu od naj-
važnijih i najraznolikijih tipova reljefnih cjelina.
Dinarska krška polja spadaju među najraznolikije
i najkompleksnije krške formacije na svijetu.
28 DINARIC KARST POLJES
(Sauro 2012), whereby recent ice-free karst
landscapes occupy between 13% (Stevanović et
al. 2016) and 20-25% (Breg Valjavec et al. 2017,
Bonacci 1987b) of the total global land area
(depending on estimations of different authors).
Closed karst poljes (or „poljes inside karst“)
are larger flat-bottomed depressions in karstic
landscapes that are completely surrounded
with elevated terrain (with at least one steep-
sloped side). Closed poljes have significant
historic and/or present hydrological functions
including sedimentation processes and karstic
drainage through underground outflows. In the
close vicinity of local underground water bodies,
they were most frequently formed by tectonic
activities followed by marginal fluvial, corrosive
and sedimentation processes (Roglić 1954,
Ford & Williams 2007). Therefore, the bottoms
of today’s karst poljes are covered by younger
(Pleistocene) sediments. Due to their size and
specific hydrological-sedimentation functions,
karst poljes play an important role in large
karstic complexes. This is especially evident when
bearing in mind that roughly 50% (or more) of
the total land area of Bosnia and Herzegovina,
Croatia, Montenegro and Slovenia is covered by
karstic terrain. Altogether, karst poljes of all types
cover 25% of the area of the Dinaric Alps.
In the Dinaric karst, following to hydrology and
fertile soils, karst poljes constitute areas that
are most suitable for many groups of organisms,
including human civilization. Surrounded by vast
areas of porous carbonate massifs and plateaus
without surface watercourses, the bottoms of
the poljes represent fluvial islands on whose
edges karstic springs that feed various surface
waterbodies (rivers, streams, lakes, swamps,
bogs and fens) are situated. The main goal of
this study was to investigate the importance
and association of fungi with local habitat types
that are formed by specific abiotic factors which
U ovom su istraživanju Dinarska krška polja podije-
ljena i klasificirana u 30 dobro diferenciranih sku-
pina koje su raspoređene u devet glavnih biogeo-
grafskih regija, temeljem većeg broja odabranih
relevantnih varijabli i kriterija. U postupku klasifi-
kacije dinarskih krških polja utvrđeni su prepo-
znatljivi obrasci za svaku grupu polja u okviru ne-
kih od najvažnijih parametara. Na odabranih 10 di-
narskih krških polja Bosne i Hercegovine te Hrvat-
ske (iz šest skupina polja) provedena su intenzivna
istraživanja tijekom različitih istraživačkih projeka-
ta. Kako bi se ocijenila važnost istraživanih polja
za mikobiotu, dobiveni rezultati su vrednovani po-
moću standardiziranih kriterija uz provedbu binar-
nog ocjenjivanja. Mikološki najvažnija polja iden-
tificirana su na temelju postignute “kumulativne
ekološke vrijednosti” (CES), omjera CES vrijedno-
sti i broja svih zabilježenih vrsta te omjera CES vri-
jednosti i broja dana istraživanja (mjere uloženog
istraživačkog napora). Uz analizu, priložen je i kra-
tak opis specifičnih tipova staništa najvažnijih za
gljive zajedno s preporukama za provedbu odgo-
varajućih mjera zaštite u tim područjima.
Na istraživanim krškim poljima zabilježen je odre-
đeni broj rijetkih i vrlo ranjivih vrsta gljiva iz od-
jeljka Ascomycota, npr. Mollisia uda, Patinella
hyalophaea, Podostroma leucopus, Scutellinia
peloponnesiaca i Lamprospora leptodictya. Nalazi
nekih vrsta iz posljednjeg bloka istraživanja, pred-
stavljaju jedine poznate nalaze na Dinaridima pa
tako ni njihov status ugroženosti još nije ocije-
njen. Zbog toga se za te vrste ovdje daju prepo-
ruke za njihovu zaštitu na temelju IUCN kriterija.
Ključne riječi: Ascomycota, klasifikacija, zaštita,
ekološka vrijednost, gljive, Bosna i Hercegovina,
Hrvatska
Introduction
Generally, karst poljes represent the largest of
five main types of depressions of karstic reliefs
29
NATURE CONSERVATION AND RURAL DEVELOPMENT
dominate the bottoms of the karst poljes. We
further tried to classify the Dinaric karst poljes
according to the occurrence of different habitat
types and their influence on the composition of
mycobiota.
Methods
To understand the variability of the ecological
conditions that are important for mycobiota
and which dominate in the karst poljes of the
Dinarides, regardless of previous delimitations
of the Dinaric karst since Roglić (1969) onwards,
we firstly delimited as precisely as possible the
extent of the Dinaric karst (mainly using the
micro-relief features with karst-determinative
elements). The next step was to identify and
characterize all larger karstic depressions
attributable to closed karst poljes according to
the criteria given by Ford & Williams (2007) and
Bonacci (2013).
This was necessary because a full list and
database of the Dinaric karst poljes was not
available for the authors. It was expected that
the extremely high bioclimatic variability and the
differences in a number of features among karst
poljes are reflected by the occurrence of different
habitat types important for fungi and by the
regional importance that certain poljes play for
their wider surroundings.
Standards and terminology
For the purpose of standardisation and saving
space in the text the following terms are here
introduced and recommended for international
use:
Low karst poljes (± ”poljes of first horizon” in Jelavić 1982), with bottoms between ~0-150 m a.s.l.
Elevated
karst poljes (± “poljes of second horizon” in Jelavić 1982) with bottoms in the range from 150-700 m
a.s.l.
Medium high
karst poljes (± “poljes of third horizon” in Jelavić (1982), except for Kupreško and Vukovsko/Ravno
polje near Kupres) with bottoms in the range of 700-1000 m a.s.l.
High karst poljes with bottoms situated at elevations between 1000-1700 m a.s.l.
Karst
micro-polje
flat-bottomed closed karstic depressions with at least one steep slope, smaller than
“small” poljes (e.g. Srijansko polje, Mt. Mosor or Njeguško polje, Mt. Lovćen) with an
underground outflow and with historical and/or currently significant sedimentation
processes. In the Dinaric karst they are mostly situated on the Adriatic islands
(Bonacci 1991) or in the highest limestone massifs.
Blatina
flooded karst polje near sea-level (cf. Ljubenkov et al. 2010) occupied by a continuous
surface of Mediterranean stagnant freshwater (swamps), different from karstic lakes
that represent the native (ancient) morphogenetic stage of the development of karst
poljes, best represented by Vrana Lake on the Island of Cres (cf. Ožanić & Rubinić 1992,
Šegota & Filipčić 2001).
Slatina karst polje approximately situated at sea-level and covered by a continuous surface of
brackish water.
Ponornica a watercourse that sinks into karstic underground at that point where it reaches
contact with porous bedrock (also swallet, stream sink, swallow hole or ponor; see
Monroe 1970, Bonacci 1987a, Ford & Williams 2007).
30 DINARIC KARST POLJES
Evaluation of existing maps and remote
sensing data
Topographic maps (scale 1:25000) which cover
the whole Dinaric Alps, including the Adriatic
lowlands and islands, were used for the detection
of karst poljes. Together with Gams (1976) and
Ranković et al. (1981) the Atlas of the climate
of former Yugoslavia for the 1931-1960 period
(Hidrometeorološka služba SFRJ 1931-1960) was
used for climatic characterisation. All recognised
karst poljes were mapped on physical maps with
superimposed Dinaric karst range boundaries
(own database) and bioclimatic belts compiled
from Ozenda (1975), Rivas-Martinez (1980),
Stefanović et al. (1983), Quezel & Barbero (1985),
Jovanović et al. (1986), Bertović & Lovrić (1992),
Antonić et al. (2000) and Kutnar & Kobler (2011). A
map of the biomes of former Yugoslavia (Matvejev
& Puncer 1989) was used for the same purpose.
As a myco-ecological criterion (modelled for
Scutellinia spp., Matočec et al. 1995) the optimal
conditions (temperature and precipitation)
for fructification in late spring, summer and
early autumn on a monthly scale was used to
visualize the variability of ecological conditions
in different karst poljes for fungi. Although a
minimal size of karstic depressions (length,
width, surface) was claimed by some authors
to determine karst poljes, this criterion varies
greatly between authors (cf. Ford & Williams
2007, Gams 1978, Krklec et al. 2015). Therefore, we
used geomorphological, morphogenetical and
hydrological criteria. According to these criteria
even very small flat-bottomed karstic depressions
can be determined as karstic micro-poljes (cf.
Frelih 2003, Bonacci 2013). A preliminary database
of karst poljes (in Excel software application)
was constructed for the analysis and progressive
classification of Dinaric karst polje, partly relying
on Jelavić’s (1967, 1982) classification concept,
the only earlier attempt of the classification of
the Dinaric karst poljes available to us. Since
closed karst poljes are mutually highly diverse
but individually entirely confined, we were able to
define discrete and mutually completely isolated
groups with homogenous abiotic conditions
(apart of hydrological objects) which can be
classified readily on the descriptive level.
Fieldwork
Ten karst poljes belonging to six groups (Fig. 1;
Tab 1) that greatly differ from each other viz.
Northwestern Dinaric montane poljes (E1), Mala
Kapela poljes (D1), Eastern-Bosnian and Sandžak
poljes (I2), Čvrsnica-Vran micro-poljes (G4), West-
ern Bosnian-Herzegovinian mid-altitude poljes
(G1) and Dalmatian-Herzegovinian low poljes (A3)
have been the subject of recent (2017-2018) and
earlier (1981-2010) intensive mycological research.
The aim of these studies was to explore the di-
versity of mycobiota in habitat types that are im-
portant for fungi and to ascertain the importance
of the given karst polje for mycobiota on a basis
of the presence of stenovalent, often sensitive,
rare and/or protected species of the largest fun-
gal phylum, Ascomycota.
The following karst poljes were investigated: (1)
Ponikve near Tršće (2) Sungerski lug high hyper-
karst small polje (3) large Ogulinsko-Plaščansko
polje in continental area with Mediterranean-like
climatic regime, all situated in the northwestern
Dinarides (Croatia), (4) altimontane Bijambare
karst randpolje, (5) Dugo polje and (6) Gornje/
Donje bare of Čvrsnica-Vran high hyperkarst
area; (7) Šujičko polje, (8) Duvanjsko polje and (9)
Livanjsko polje, all of them medium high poljes of
the Central Dinarides (Bosnia and Herzegovina),
and (10) Konavosko polje in the Mediterranean
lowlands (Croatia).
For calibration of the present evaluation the in-
tensively researched Ponikve on Mt. Medvednica
(situated outside the Dinaric karst in Croatia) was
31
NATURE CONSERVATION AND RURAL DEVELOPMENT
selected as an outgroup polje for which previous
studies ascertained that it is of low significance
for regional mycobiota.
Evaluation of karst poljes by myco-
bioindication
The importance of karst poljes which were studied
by relatively intensive fieldwork, was preliminary
evaluated by summing up the binary (except (d)
and (e) rarity categories) grading scale assigned to
the recorded ascomycete species, i.e. the “cumula-
tive ecological score” (CES), as follows: (a) ecologi-
cal stenovalence; (b) local species’ habitat exclu-
sivity; (c) species’ habitat vulnerability/stability/
threat; (d) regional rarity of the species: species
known from a single locality in the Dinarides = 1,
for species known from 2-5 localities = 0.5, and
known from 6 or more localities = 0; and (e) con-
servation status IUCN status or any other evalua-
tion in any of the countries of the Dinaric Alps.
Results
Stepwise classification of Dinaric karst poljes
according to habitat availabilty for fungi
We recognized 30 karst polje groups that can be
assembled in nine geographic units (regions) on
the basis of geographical, geological, morphologi-
cal, hydrological, climatic, bioclimatic and myco-
ecological criteria (Tab 1).
Those groups are differentiated from each other
by highly diverse abiotic conditions.
Generally, it is known that annual precipita-
tion and cloudiness decreases from the NW,
with the NW Dinaric karst poljes receiving
around 3,000 mm annual precipitation, to the
SE, where some Sandžak high poljes receive be-
low 1,000 mm as well as from the SW to the NE
(cf. Hidrometeorološka služba SFRJ (1931-1960),
Radičević et al. 1980, Poje et al. 1984). But in the
Central-Adriatic arid zone some micro-poljes on
the Adriatic Islands receive <600 mm precipita-
tion p.a. On the other hand, global radiation on
horizontal surfaces (which approximates the situ-
ation on the bottom of karst poljes) increases sig-
nificantly from the NW Dinarides towards the SE
as well as from the N to the S (Matić 2007, www.
meteonorm.com). Additionally, along the N-S
axis the continental climatic regime (precipita-
tion minimum in the cooler half of the year) turns
into the maritime regime (precipitation minimum
in the warmer half of the year, cf. Ranković et al.
1981) which has a strong influence on the ap-
pearance and fructification period of fungi (see
column “Myco-phenology” in Tab. 1). Temperature
greatly depends on altitude with higher annual
air temperatures in lower altitudes (karst polje
bottoms vary from below sea level up to 1,700
m a.s.l.). Besides altitude, the amplitude of an-
nual air temperature increases from the SW, i.e.
the Adriatic Sea, towards the NE. Thus, all karst
poljes of Mediterranean groups have all-year i.e.
365 days with mean daily temperature above 0
°C (karst poljes from group A with mean annual
air temperature ~14–17 °C and poljes from group
B with annual air temperature ~10-14 °C). On the
other hand, the highest Southeast and Central Di-
naric karst poljes (groups G4 and I 1-4) have only
220-260 days with mean daily temperature above
0 °C per year, with mean annual air temperature
of only (2) 4–8 °C (see Tab. 1).
The relationship between vegetation and the lo-
cal climate has been ascertained by Bertović
(1975). The Dinaric karst poljes are situated in all
bioclimatic belts from the thermo-Mediterranean
up to the alti-Mediterranean/Southeastern Di-
naric alpine “islands”, i.e. from the Mediterranean
side to the highest ridges of the Dinaric Alps, and
from the alpine to the colline-continental belt on
the Pannonian side of the mountains.
32 DINARIC KARST POLJES
Fig. 1: Dinaric karst poljes overview – Classification and distribution of geographical groups with highlighted position of karst poljes
that have been investigated. (Author: Neven Matočec, derived from personal database “Dinaric karst poljes and canyons 1984-2018”)
List of studied karst poljes
1 - Ponikve, Mt. Medvednica
2 - Ponikve (Tršće), Gorski kotar
3 - Sungerski lug (Mrkopalj), Gorski kotar
4 - Bijambare (Nišići)
5 - Ogulinsko-plaščansko polje
6 - Livanjsko polje
7 - Duvanjsko polje
8 - Šujićko polje
9 - Dugo polje (Mt. Čvrsnica - Mt. Vran)
10 - Gornja and Donja bara, Mt. Čvrsnica
11 - Konavosko polje (Dubrovnik)
Bioclimatic border-lines
after Ozenda (1975), Rivas-Martinez (1980), Stefanović
et al. (1983), Quezel & Barbero (1985), Jovanović et al.
(1986), Bertović & Lovrić (1992), Antonić et al. (2000),
Kutnar & Kobler (2011), and personal field data.
33
NATURE CONSERVATION AND RURAL DEVELOPMENT
Regarding area size, micro-poljes are prevalent either in
the lowest altitudes up to 150 m a.s.l., on islands (groups
A1a-b, B1) or in the highest altitudes, mainly between 900
and 1,700 m a.s.l. (groups E2, G2, I1, I3, I4). The largest karst
poljes (nearly all being structural, cf. Ford & Williams 2007)
except Konavosko and Peštersko polje, are situated on
external Dinaric thrust geological unit (cf. Pamić & Jurković
2002, Placer et al. 2010).
The occurrence of different habitat types that are important
for fungi, depends largely on hydrological conditions of
karst polje bottoms. Together with Mediterranean freshwater
swamps (blatinas) or brackish waterbodies (slatinas), karst
lakes occur either in the lowest Mediterranean region (groups
A1b, B1 and B2) or in the highest elevations of the alti-
Mediterranean up to the Dinaric-alpine zone (groups I3 and
I4). Karst poljes with perennial watercourses and high water
levels dominate in the NW Dinaric region. These areas are
characterized by very high annual precipitation and a strong
maritime climatic regime (groups C3, D1, E1-E3). Central and
South-eastern Dinaric poljes that are in hydrological contact
with high and large, highly porous carbonate massifs and
plateaus with huge underground aquifers (groups G1, G3 and
H1) are also rich in perennial watercourses.
Evaluation of karst poljes based on preliminary analyses
of the ascomycete fungi
The significance of the investigated karst poljes for
mycobiota was evaluated by five criteria, already explained
above, and the ascomycetes who were recorded in frame
of various projects in 10 karst poljes during the last four
decades. The numbers of species recorded per polje, the
cumulative “ecological score” (CES) and estimated research
level are given in Tab. 2.
As an auxiliary tool for estimating the importance of Dinaric
karst poljes for ascomycete fungi and the exploration
level we used an extra-Dinaric micro-polje (Ponikve on Mt.
Medvednica) situated in a small patch of isolated karst
previously assessed as a polje of low significance to regional
mycobiota.
Delimitation lines
among groups of
karst poljes
Karst polje
Studied
karst polje
External boundary
of Dinaric karst
Extent of Adriatic-
Dinaric carbonate
platform (external
Dinarides), after Pamić
& Jurković (2002)
G3
4
34 DINARIC KARST POLJES
Tab. 1: Karst poljes polyphasic classification based on seven types of different delimitation criteria
Geogr
group Group name Bioclimat.
zone Drainage Geological
subdivision Altitude Mean annual
temp. 10-d seq.
T>°C Mean annual
precip. Myco-fenology (model-Scutellinia) Prevalent
size Polje
type
I II III IV V VI VII VIII IX X XI XII
A1a Dalmatian Adriatic island poljes (HR) TM-MM Adriatic Ext-Din-imbricate ~0-150 15-17 36 600-1250 × × × × × × micro DP
A1b South Dalmatian island blatinas/Baćina lakes (HR) TM-MM Adriatic Ext-Din-imbricate (-)50-15 16-17 36 1000-1250 × × × × × × × × × × micro SMP
A2 Korčula-Pelješac elevated poljes (HR) MM-SM Adriatic Ext-Din-imbricate 150-350 14-15 36 1250-1500 × × × × × × × × × small-large BP
A3 Dalmatian-Herzegovinian low poljes (HR/BA) MM-SM Adriatic External-Dinaric ~0-250 14-16 36 1500-2500 × × × × × × × × × × small-large DP / BP
B1 Kvarnerian-Kras poljes (HR/IT) MM-SM Adriatic Adr/Ext-Din-imbr (-)30-100 (12)14-16 36 1000-1500 × × × × × × × × × × micro-small SMP
B2 Ravni kotari-Šibenik low poljes (HR) MM-SM Adriatic Ext-Din-imbricate ~0-150 14-16 36 800-1000 × × × × × × small-large SMP
B3 Dalmatian Zagora poljes (HR) SM Adriatic External-Dinaric 250-650 10-14 36 1250-1750 × × × × × × × × × × micro-small DP / BP
B4 Northern Dalmatian poljes (HR) SM Adriatic Ext-Din-imbricate 200-300 12-14 36 1000-1250 × × × × × × × × micro BP
B5 Dalmatian-Herzegovinian elevated poljes (HR/BA) SM Adriatic Ext-Din-thrust 250-600 10-14 36 1250-1750 × × × × × × × × × small-large SP / DP
C1 South-eastern Lika poljes (HR) SM-OM Adriatic Ext-Din-thrust 600-800 6-10 28-30 1500-1750 × × × × × × × × × × micro-med. RP
C2 Ćićarija-Primorje poljes (HR/SI) SM-OM Adriatic External-Dinaric 200-750 6-12 30-35 1000-2500 × × × × × × × × × × × × small-med. SP
C3 Postojna region (SI) SM(OM) Adr/Bl_Sea Ext-Din-thrust 300-600 8-10 30-35 1500-2500 × × × × × × × × × × × × small-large SP
G1 W-Bosnian-Herzegovinian mid-altitude poljes (BA) SM-OM Adriatic Ext-Din-thrust 700-1000 8-12 27-30 1250-1750 × × × × × × × × × × small-large SP
D1 Mala Kapela poljes (HR) MD(MT) Black Sea Ext-Din-thrust 300-500 8-10 28-30 1500-2500 × × × × × × × × × × × × small-large SP
D2 Central & Western Lika poljes (HR) MD Adr/Bl_Sea Ext-Din-thrust 400-700 6-10 28-30 1000-1750 × × × × × × × × × × small-large BP
D3 NW Dinaric colline-continental poljes (SI/HR) COL Black Sea Ext-Din-thrust 200-450 8-10 30-32 1000-1500 × × × × × × × × × × × micro-med. BP
F1 Una drainage poljes (BA/HR) MD/OM Black Sea Ext-Din-thrust 500-700 6-10 29-30 1250-1750 × × × × × × × × × × small-large RP
F2 Sana-Vrbas poljes (BA) MT-AMT Black Sea Internal-Dinaric 400-1000 7-10 28-31 1000-1500 × × × × × × × × × × × micro-med. RP
E1 NW Dinaric montane poljes (SI/HR) MT-AMT(SA) Black Sea* Ext-Din-thrust 500-1000 6-10 26-30 1750-3000 × × × × × × × × × × × × small-large SP
E2 Velebit high micropoljes (HR) AMT-SA Adriatic Ext-Din-thrust 700-1250 4-8 26-29 2000-3000 × × × × × × × × × × × × micro SP
E3 Northern Lika montane poljes (HR) MT-AMT Adriatic** Ext-Din-thrust 600-800 6-8 26-28 1000-1500 × × × × × × × × × × small-med. BP
G2 Čemernica-Dinara high micropoljes (HR) OM-AM Adriatic Ext-Din-thrust 900-1250 4-8 25-28 1500-2000 × × × × × × × × × × × micro DP
G3 W-Bosnian-Herzegovinian high poljes (BA) AMT Adr/Bl_Sea Ext-Din-thrust 900-1250 4-8 (10) 24-28 1250-1500 × × × × × × × × × × × small-large SP
G4 Čvrsnica-Vran high micropoljes (BA) AM Adriatic Ext-Din-thrust 1200-1400 4-8 25-26 1750-2000 × × × × × × × × × × × small-med. SP / DP
H1 E-Herzegovinian-Montenegrin elevated poljes (BA/MN) SM Adriatic Ext-Din-thrust 200-700 8-12 36 1500-2500 × × × × × × × × × × small-large SP / DP
H2 E-Herzegovinian-Montenegrin high poljes (BA/MN) OM Adriatic Ext-Din-thrust 700-1200 6-10 24-28 1500-3000 × × × × × × × × × × small-large SP / DP
I1 Morača-Zeta high micro-poljes (MN) OM-AM Adriatic Ext-Din-thrust 900-1680 (2) 4-8 22-26 2000-2500 × × × × × × × × × × × micro SP / DP
I2 E-Bosnian-Sandžak high poljes (BA/SRB/MN) AMT-DA Black Sea Internal-Dinaric 700-1700 4-8 22-26 750-1500 × × × × × × × × × × × × small-large RP
I3 Bosnian-Montenegrin high micro-poljes (BA/MN) AM-DA Adr/Bl_Sea Internal-Dinaric 1000-1700 (2) 4-8 21-26 1500-2000 × × × × × × × × × × × × micro AP
I4 Prokletian high micropoljes (MN/AL) AM-DA Adr/Bl_Sea Internal-Dinaric 1000-1500 (2) 4-8 21-25 1500-2000 × × × × × × × × × × micro AP
Karst polje geographical units (regions)
AMediterranean region of lower Adriatic BMediterranean region of upper Adriatic CNW peri-Mediterranean Dinarides
DNW Lower Dinarides ENW higher Dinarides FCentral peri-Pannonian Dinarides
GCentral Dinarides HExternal SE Dinarides IAlpine SE Dinarides
× monthly mean precipitation >70 mm monthly mean air temperature 10-22 °C (after Hidrometeorološka služba SFRJ (1931-1960)
Karst polje types (after Ford & Williams 2007, supplemented)
RP surface spring → sinkhole (Randpoljes) SP / BP karstic spring → sinkhole (Structural poljes or Baselevel poljes)
DP no significant hydrology - Dry poljes AP alpine poljes
SMP maritime karstic lakes (freshwater or brakish) - Submerged poljes
*
** Gomance - Adriatic drainage;
Čujića krčevina - Black Sea drainage.
SA
AMT
MT
COL
TM
MM
SM
OM
AM
DA
MD
Subalpine (continental);
Altimontane (continental);
Montane (continental);
Colline (continental);
Thermo-Mediterranean;
Meso-Mediterranean;
Sub-Mediterranean;
Oro-Mediterranean;
Alti-Mediterranean;
Dinaric-alpine;
Mediterranoid (continental,
colline to montane).
A1a
A1b
A2
A3
B1
B2
B3
B4
B5
C1
C2
C3
D1
D2
D3
E1
E2
E3
F1
F2
G1
G2
G3
G4
H1
H2
I1
I2
I3
I4
Blatsko polje, Korčula;
Blatsko polje, Mljet;
Orlovo polje, Pelješac;
Jezero polje (Vrgorac);
Ponikve, Krk;
Bokanjačko blato (Zadar);
Mućko polje;
Bare (Đevrske);
Imotsko polje;
Velikopopinsko polje (Gračac);
Grobničko polje (Rijeka);
Postojnska kotlina;
Drežnički lug;
Krbavsko polje;
Grosupeljsko polje;
Cerkniško polje;
Bunovac, Mt. Velebit;
Vrhovinsko polje;
Petrovačko polje;
Podrašničko polje (Mrkonjić Grad);
Livanjsko polje;
Brezovac, Mt. Dinara;
Kupreško polje;
Dugo polje (Mt. Čvrsnica-Vran);
Dabarsko polje;
Nevesinjsko polje;
Cigovića bare, Mt. Lola;
Peštersko polje;
Masna bara, Mt. Lelija;
Rikavačko jezero, Mt. Žijovo.
Karst poljes representatives:
Key:
35
NATURE CONSERVATION AND RURAL DEVELOPMENT
Tab. 1: Karst poljes polyphasic classification based on seven types of different delimitation criteria
Geogr
group Group name Bioclimat.
zone Drainage Geological
subdivision Altitude Mean annual
temp. 10-d seq.
T>°C Mean annual
precip. Myco-fenology (model-Scutellinia) Prevalent
size Polje
type
I II III IV V VI VII VIII IX X XI XII
A1a Dalmatian Adriatic island poljes (HR) TM-MM Adriatic Ext-Din-imbricate ~0-150 15-17 36 600-1250 × × × × × × micro DP
A1b South Dalmatian island blatinas/Baćina lakes (HR) TM-MM Adriatic Ext-Din-imbricate (-)50-15 16-17 36 1000-1250 × × × × × × × × × × micro SMP
A2 Korčula-Pelješac elevated poljes (HR) MM-SM Adriatic Ext-Din-imbricate 150-350 14-15 36 1250-1500 × × × × × × × × × small-large BP
A3 Dalmatian-Herzegovinian low poljes (HR/BA) MM-SM Adriatic External-Dinaric ~0-250 14-16 36 1500-2500 × × × × × × × × × × small-large DP / BP
B1 Kvarnerian-Kras poljes (HR/IT) MM-SM Adriatic Adr/Ext-Din-imbr (-)30-100 (12)14-16 36 1000-1500 × × × × × × × × × × micro-small SMP
B2 Ravni kotari-Šibenik low poljes (HR) MM-SM Adriatic Ext-Din-imbricate ~0-150 14-16 36 800-1000 × × × × × × small-large SMP
B3 Dalmatian Zagora poljes (HR) SM Adriatic External-Dinaric 250-650 10-14 36 1250-1750 × × × × × × × × × × micro-small DP / BP
B4 Northern Dalmatian poljes (HR) SM Adriatic Ext-Din-imbricate 200-300 12-14 36 1000-1250 × × × × × × × × micro BP
B5 Dalmatian-Herzegovinian elevated poljes (HR/BA) SM Adriatic Ext-Din-thrust 250-600 10-14 36 1250-1750 × × × × × × × × × small-large SP / DP
C1 South-eastern Lika poljes (HR) SM-OM Adriatic Ext-Din-thrust 600-800 6-10 28-30 1500-1750 × × × × × × × × × × micro-med. RP
C2 Ćićarija-Primorje poljes (HR/SI) SM-OM Adriatic External-Dinaric 200-750 6-12 30-35 1000-2500 × × × × × × × × × × × × small-med. SP
C3 Postojna region (SI) SM(OM) Adr/Bl_Sea Ext-Din-thrust 300-600 8-10 30-35 1500-2500 × × × × × × × × × × × × small-large SP
G1 W-Bosnian-Herzegovinian mid-altitude poljes (BA) SM-OM Adriatic Ext-Din-thrust 700-1000 8-12 27-30 1250-1750 × × × × × × × × × × small-large SP
D1 Mala Kapela poljes (HR) MD(MT) Black Sea Ext-Din-thrust 300-500 8-10 28-30 1500-2500 × × × × × × × × × × × × small-large SP
D2 Central & Western Lika poljes (HR) MD Adr/Bl_Sea Ext-Din-thrust 400-700 6-10 28-30 1000-1750 × × × × × × × × × × small-large BP
D3 NW Dinaric colline-continental poljes (SI/HR) COL Black Sea Ext-Din-thrust 200-450 8-10 30-32 1000-1500 × × × × × × × × × × × micro-med. BP
F1 Una drainage poljes (BA/HR) MD/OM Black Sea Ext-Din-thrust 500-700 6-10 29-30 1250-1750 × × × × × × × × × × small-large RP
F2 Sana-Vrbas poljes (BA) MT-AMT Black Sea Internal-Dinaric 400-1000 7-10 28-31 1000-1500 × × × × × × × × × × × micro-med. RP
E1 NW Dinaric montane poljes (SI/HR) MT-AMT(SA) Black Sea* Ext-Din-thrust 500-1000 6-10 26-30 1750-3000 × × × × × × × × × × × × small-large SP
E2 Velebit high micropoljes (HR) AMT-SA Adriatic Ext-Din-thrust 700-1250 4-8 26-29 2000-3000 × × × × × × × × × × × × micro SP
E3 Northern Lika montane poljes (HR) MT-AMT Adriatic** Ext-Din-thrust 600-800 6-8 26-28 1000-1500 × × × × × × × × × × small-med. BP
G2 Čemernica-Dinara high micropoljes (HR) OM-AM Adriatic Ext-Din-thrust 900-1250 4-8 25-28 1500-2000 × × × × × × × × × × × micro DP
G3 W-Bosnian-Herzegovinian high poljes (BA) AMT Adr/Bl_Sea Ext-Din-thrust 900-1250 4-8 (10) 24-28 1250-1500 × × × × × × × × × × × small-large SP
G4 Čvrsnica-Vran high micropoljes (BA) AM Adriatic Ext-Din-thrust 1200-1400 4-8 25-26 1750-2000 × × × × × × × × × × × small-med. SP / DP
H1 E-Herzegovinian-Montenegrin elevated poljes (BA/MN) SM Adriatic Ext-Din-thrust 200-700 8-12 36 1500-2500 × × × × × × × × × × small-large SP / DP
H2 E-Herzegovinian-Montenegrin high poljes (BA/MN) OM Adriatic Ext-Din-thrust 700-1200 6-10 24-28 1500-3000 × × × × × × × × × × small-large SP / DP
I1 Morača-Zeta high micro-poljes (MN) OM-AM Adriatic Ext-Din-thrust 900-1680 (2) 4-8 22-26 2000-2500 × × × × × × × × × × × micro SP / DP
I2 E-Bosnian-Sandžak high poljes (BA/SRB/MN) AMT-DA Black Sea Internal-Dinaric 700-1700 4-8 22-26 750-1500 × × × × × × × × × × × × small-large RP
I3 Bosnian-Montenegrin high micro-poljes (BA/MN) AM-DA Adr/Bl_Sea Internal-Dinaric 1000-1700 (2) 4-8 21-26 1500-2000 × × × × × × × × × × × × micro AP
I4 Prokletian high micropoljes (MN/AL) AM-DA Adr/Bl_Sea Internal-Dinaric 1000-1500 (2) 4-8 21-25 1500-2000 × × × × × × × × × × micro AP
Karst polje geographical units (regions)
AMediterranean region of lower Adriatic BMediterranean region of upper Adriatic CNW peri-Mediterranean Dinarides
DNW Lower Dinarides ENW higher Dinarides FCentral peri-Pannonian Dinarides
GCentral Dinarides HExternal SE Dinarides IAlpine SE Dinarides
× monthly mean precipitation >70 mm monthly mean air temperature 10-22 °C (after Hidrometeorološka služba SFRJ (1931-1960)
Karst polje types (after Ford & Williams 2007, supplemented)
RP surface spring → sinkhole (Randpoljes) SP / BP karstic spring → sinkhole (Structural poljes or Baselevel poljes)
DP no significant hydrology - Dry poljes AP alpine poljes
SMP maritime karstic lakes (freshwater or brakish) - Submerged poljes
36 DINARIC KARST POLJES
a) Photo: Nedim Jukić b) Photo: Ivana Kušan
e) Photo: Neven Matočec f) Photo: Neven Matočec
i) Photo: Jasmin Jukić j) Photo: Neven Matočec
o) Photo: Nedim Jukićn) Photo: Neven Matočecm) Photo: Nedim Jukić
37
NATURE CONSERVATION AND RURAL DEVELOPMENT
Fig. 2: Different types and groups of karst poljes and karstic depressions in Croatia and
Bosnia and Herzegovina: a) View towards Dugo polje, Bare - Mt. Čvrsnica; b) Šujičko
polje and its appearance in the peak of the dry season; c) Dugo polje landscape with Mt.
Čvrsnica massif in the background; d) Plaščansko polje with ponornica Dretulja, near
the city of Ogulin; e) Duvanjsko polje panorama; f) Čvrsničke bare micro-poljes with
altimontane marsh vegetation; g) View on Blidinje Lake and the NW borders of Dugo
polje from just below Pločno, the highest peak of Mt. Čvrsnica; h) Karstic river Sturba,
one of the most important hydrological elements and watercourses of Livanjsko polje; i)
Livanjsko polje, natural water retention and karstic watercourse near the village Bastasi;
j) Konavosko polje near the city of Dubrovnik (Konavle municipality); k) Acidophilic bog
in the area of Protected landscape Bijambare; l) Ogulinsko-Plaščansko polje and karstic
spring Sabljaki; m) Brčanj stream, main watercourse of Brčanjsko polje; n) Sungerski
lug, acidofilic bog surrounded by coniferous forest; o) Livanjsko polje, scenery near the
Vrbica village; p) Masna Luka, fragmented parts of Dugo polje.
c) Photo: Neven Matočec d) Photo: Neven Matočec
g) Photo: Nedim Jukić h) Photo: Nedim Jukić
k) Photo: Nedim Jukić l) Photo: Neven Matočec
p) Photo: Nedim Jukić
38 DINARIC KARST POLJES
Photo: Neven Matočec Photo: Nedim Jukić c)
Photo: Nihad Omerović
Photo: Neven Matočec i)
Photo: Nedim Jukić
o) Photo: Neven Matočec p) Photo: Neven Matočec
s)
a)
g)
q) Photo: Nedim Jukić Photo: Nedim Jukić
b)
h) Photo: Neven Matočec
Photo: Nedim Jukić
n)
Scale bars:
b), h), j), k), m)
a), g), e), l), q), r), s)
c), f), i)
d), n), o), p)
1 mm
2 mm
0,5 cm
1 cm
39
NATURE CONSERVATION AND RURAL DEVELOPMENT
d)
f)e)
Photo: Nihad Omerović
k)
l) m)
Photo: Ivana Kušan
Fig. 3: Ascomycete fungi recorded in the karst poljes
in Croatia and Bosnia and Herzegovina:
a) Scutellinia peloponnesiaca, Ogulinsko-
plaščansko polje; b) Lamprospora leptodictya,
Bare - Mt. Čvrsnica; c) Scutellinia subhirtella, Gornje
bare - Mt. Čvrsnica; d) Helvella albella, Masna
Luka - Dugo polje; e) Vibrissea calcaria, Sturba river
- Livanjsko polje; f) Myriosclerotinia sp., Vrbica -
Livanjsko polje; g) Parascutellinia carneosanguinea,
Ostrožac stream, Duvanjsko polje; h) Heyderia
abietis, Ponikve (Tršće); i) Helvella corium, Ostrožac
stream - Duvanjsko polje; j) Patinella hyalophaea,
Bijambare; k) Ascobolus xylophilus, Bijambare;
l) Scutellinia superba, Šujičko polje; m) Mollisia
uda, Ogulinsko-plaščansko polje; n) Podostroma
leucopus, Bijambare; o) Sarcoscypha macaronesica,
Konavosko polje; p) Peziza obtusapiculata,
Sungerski lug; q) Boudiera tracheia, Bare - Mt.
Čvrsnica; r) Vibrissea truncorum, Ponikve (Tršće); s)
Scutellinia hyperborea, Masna Luka - Dugo polje.
Photo: Nedim Jukić
Photo: Nihad Omerović
r)
Photo: Nihad Omerović
Photo: Neven Matočecj)
Photo: Nedim Jukić
Photo: Nedim Jukić
40 DINARIC KARST POLJES
Nevertheless, several of the localities which were
studied, and which harbour habitat types that
are important for some ecological or taxonomic
groups, we designated regardless of their index
value as areas of special conservation interest.
These are:
(1) Vrbica is a specific area in Livanjsko polje, the
largest karst polje in Bosnia and Herzegovina
(group G1; cf. Fig. 1) characterised by Salix
spp. stands with dense hygrophilous swamp
vegetation located in a basin filled with a
permanent stagnant waterbody with coarse and
rotten woody debris of the willows and with
pronounced nitrification processes. This habitat
type is extremely rare and difficult to study.
It should be, therefore, in the focus of future
research. In the area we found a still unidentified
species of Myriosclerotinia. Nearly all species
of this genus are rare and endangered in most
countries (i.e., M. dennisii is listed as Critically
Dinaric karst poljes
harbour a high species
diversity of Ascomycota,
as well as a number of
species which can be used
as bioindicators, and also
a number of rare and
threatened species.
In the frame of recent
mycological research, four
species were recorded for the
first time for the mycobiota
of Bosnia and Herzegovina,
and two further species for
the mycobiota of Croatia.
In the frame of recent mycological research, since
1982, four species were recorded for the first time
for the mycobiota of Bosnia and Herzegovina:
Coprotus luteus, Helvella albella, Lamprospora
leptodictya and Scutellinia superba, and two
further species for the mycobiota of Croatia:
Mollisia uda and Scutellinia peloponnesiaca.
Very rare species which are largely confined to
special and often threatened habitat types are:
L. leptodictya, M. uda, Patinella hyalophaea,
Podostroma leucopus and S. peloponnesiaca (Fig.
3). The latter species were used to recommend
regional protection measures.
Discussion
All ascomycete species which were recorded in
the karst poljes that have been studied more
or less sporadically and with different levels of
intensity during the 1982–2018 period, were used
for the evaluation process (Tab. 2). The aim of our
assessment was to find objective correlations
between different groups of karst poljes and the
diversity of mycobiota. We further evaluated the
importance for conservation of all investigated
poljes based on the fungal species which have
been found in the area. According to our analyses,
Konavosko polje and Sungerski lug are the most
valuable karst poljes with highest “Cumulative
ecological scores” (CES). However, at the same
time, the degree of exploration for both areas
is also highest, as well as for Ponikve on Mt.
Medvednica which reached one of the lowest CES
values. As a measure for the potential value we
calculated the ratio between CES and a number of
days representing the research effort for all poljes.
Ogulinsko-plaščansko polje, Ponikve near Tršće,
Dugo polje, below Mt. Čvrsnica, and Duvanjsko
polje reached the highest values, followed by
Gornja and Donja bara on Mt. Čvrsnica, Konavosko
polje and Livanjsko polje. Similar results were
obtained when dividing the CES value through the
number of all recorded species.
41
NATURE CONSERVATION AND RURAL DEVELOPMENT
Endangered in Croatia; cf. Tkalčec et al. 2008).
Hence, this is a very important discovery.
(2) Dugo polje and its surrounding karstic
landscape (group G4; Fig. 1) is very important
for nature protection and its mycobiota due
to its large luxuriantly developed relict forest
dominated by the SE Dinaric endemic tree Pinus
heldreichii accompanied with a number of other
endemic arborescent species (e.g., Lonicera spp.).
The polje is situated in the subalpine bioclimatic
belt. Together with the upper xeronival belt, the
subalpine bioclimatic belt can be found in some
of the highest massifs of SE Dinarides. Both
bioclimatic belts show a strong Mediterranean
influence (cf. Ranković et al. 1981, Jovanović et al.
1986) and represent biogeographical areas that
are sharply separated from their Euro-Siberian
bioclimatic counterparts (cf. Puşcaş & Choler 2012).
Other areas with particularly high indices concern
several freshwater habitats: karstic springs, small
ponornicas with developed fens and the bank
of Blidinje Lake with tall Salix and Phragmites
stands. Among a number of ascomycete fungi
recorded in this locality, Boudiera tracheia and
Helvella albella are most important for nature
protection. This very important area should be
intensively investigated in the future.
(3) For the purpose of this study Gornja and Donja
Čvrsnička bara (group G4; Fig. 1) are considered
as a micro-polje complex. These two small
karstic depressions with steep sides harbour
small watercourses and karstic springs, some
temporary but most of it perennial water bodies.
The special importance of these localities lies
in its location in high altitudes between 1,300
and 1,400 m a.s.l.). They represent the only near-
alpine karst poljes in group G4 (Tab.1, Fig. 1) with
a strong Mediterranean influence. Karst poljes
in comparable localities are known only from
a few sites in the high SE Dinarides (groups I1
and I4; cf. also Ranković et al. 1981, Jovanović
et al. 1986). In some localities, like in Gornje
bare, water is retained on the soil surface for
a whole year. These conditions are favourable
for fungal species associated with cyperaceous
and juncaceous plants and their remnants.
Furthermore, the area soaked with water around
springs and rivulets is the habitat for some very
rare and sensitive arctic-alpine fungal species.
Besides Boudiera tracheia (a species listed as
Endangered in Bosnia and Herzegovina, Jukić
& Omerović 2017) the very rare bryo-parasitic
fungus Lamprospora leptodictya was recorded
in Donje bare polje. This find represents the
first known record in the Dinaric Alps for the
species which is hitherto known from the Arctic
and northern European countries (Schumacher
1993). Therefore, the species should be strictly
protected in Bosnia and Herzegovina.
(4) Bijambare represents a very valuable and
rather large complex of habitat types of old
growth altimontane coniferous forests rich in
coarse woody debris. These forests are adapted
to frequent and heavy frosts during winter (group
I2; Fig.1). They are tremendously rich in fungal
species (cf. Ódor et al. 2006) and therefore of high
importance for mycobiota. The permanently wet
conditions further favoured the development
of a bog, a special freshwater habitat with very
specific assemblages of fungi. In Bijambare we
recorded the highest number of fungi (Tab. 2). The
area is therefore of particular interest. Patinella
hyalophaea and Podostroma leucopus inhabit
very sensitive and endangered habitat types.
Both species were recorded in the Protected
Landscape Bijambare and were identified as
candidates for a special protection status in
Bosnia and Herzegovina.
Very similar habitat types exist in (5) Sungerski
lug and (6) in the Ponikve area near Tršće,
42 DINARIC KARST POLJES
situated in the perhumide climate of the NW
Dinarides (group E1; Fig. 1). In these areas bogs
are much smaller and situated in significantly
lower altitudes then in Bijambare. Nevertheless,
the importance of both localities under extreme
climatic conditions is evident in Tab. 2.
(7) In contrast to the former areas Konavos-
ko polje is situated in the Mediterranean cli-
matic zone (group A3; Fig. 1) with all-year air
mean temperatures above 0 °C and yearly mean
temperature above 14 °C. However, it is situated
in an area of relative high precipitation. The im-
portance of this polje derives from several strong
karstic and flysch springs and rather short ponor-
nicas with some of them embedded in tall ev-
ergreen laurel-oak forest. These few small but
permanently moist and shady habitats are of the
highest importance for mycobiota. The bottom of
the polje is still occupied by extensive Mediterra-
nean traditionally managed grasslands important
for specific grassland fungi (e.g. Geoglossum spp.).
Previous research conducted during the past
decades in Croatian karst poljes, and current
research in the Central Dinarides of Bosnia and
Tab. 2: Evaluation of Dinaric karst poljes based on ascomycetous fungi. In addition, protected species and/or species which were used for the present evaluation are shown
Polje name Geographical orientation Country Size class Spec. no. Res. days CES CES/sp CES/day Exclusive polje’s habitat of special value to mycobiota Protected and/or evaluated species
Ponikve Mt. Medvednica, NW from Zagreb HR micro 28 16 9,5 0,34 0,59 none Helvella phlebophora (HR: VU)
Ponikve Tršće, NW from Delnice HR micro 9 3 24 2,67 8 bog Vibrissea truncorum (HR: VU),
Heyderia abietis (HR: NT)
Sungerski lug Sunger, SSE from Delnice HR small 46 24 70,5 1,53 2,94 small bogs surrounded by tall old growth coniferous forest
Discina fastigiata (HR: CR),
Caloscypha fulgens (HR: VU),
Geopyxis majalis (HR: VU),
Trichoglossum hirsutum (HR: VU),
Peziza obtusapiculata (HR: DD),
Heyderia abietis (HR: NT)
Bijambare Nišići, NNE from Sarajevo BA small 23 13 49,5 2,15 3,81 bog, old growith coniferous altimontane forest
Cudonia circinans (HR: VU),
Heyderia abietis (HR: NT),
Ascobolus xylophilus (BA: CR),
Plicaria trachycarpa (BA:VU),
Patinella hyalophaea (BA candidate: EN),
Podostroma leucopus (BA: candidate EN)
Ogulinsko-plaščansko polje Ogulin HR large 6 2 21 3,5 10,5 fens and riparian forest in mediterranoid continental belt Mollisia uda (HR candidate: EN),
Scutellinia peloponnesiaca HR candidate: VU)
Livanjsko polje Livno BA large 5 2 8 1,6 4 fens, two types of riparian forests, swamp, traditionally managed grassland Myriosclerotinia sp. (M. dennisii HR: CR)
Duvanjsko polje Tomislavgrad BA large 4 1 6,5 1,63 6,5 various oro-Mediterranean brooks, traditionally managed grassland Helvella corium (BA: EN),
Parascutellinia carneosanguinea (BA: VU)
Šujičko polje Šujica BA medium 3 1 1 0,33 1 tall riparian forest none
Dugo polje Blidinje jezero BA medium 14 4 26 1,86 6,5 fens and Pinus heldreichii endemic (alti-Mediterranean) forest, traditionally
managed grassland Helvella albella (HR: CR),
Boudiera tracheia (BA: EN)
Gornja i donja bara pl. Čvrsnica BA micro 3 2 8,5 2,83 4,25 fens in alti-Mediterranean belt Boudiera tracheia (BA: EN)
Lamprospora leptodictya (BA candidate: EN)
Konavosko polje Čilipi HR large 33 17 73,5 2,23 4,32 karstic springs and riparian forest in Mediterranean belt Geoglossum cookeianum (HR: VU),
Geoglossum umbratile (HR: VU),
Sarcoscypha macaronesica (HR: VU)
43
NATURE CONSERVATION AND RURAL DEVELOPMENT
(Lakušić 1970, Lovrić & Rac 1989, Bertović & Lovrić
1992, Ozimec et al. 2013b, 2013c, Stumberger 2015)
and with analyses of the presence of particular
habitat types in some large karst poljes (Schwarz
2013, Sackl et al. 2014). Like other researchers, we
conclude that many karst poljes, in comparison
to surrounding vast areas of waterless dry lime-
stone massifs (see Fig. 1) are some kind of oases
of high habitat diversity with otherwise rare and
exclusive habitats which function as key refugia
and strongholds for many rare, endangered and
endemic plants, fishes, birds, invertebrates and
also for a vast number of fungi.
Herzegovina, show that a number of the Dinaric
karst poljes harbour a high species diversity of
Ascomycota, as well as a number of species which
can be used as bioindicators, and also a number
of rare and threatened species. Their significance
is evident (Tab. 2) by comparing these localities to
isolated, extra-Dinaric micro-polje in the colline
continental humid zone (Ponikve, Mt. Medvednica;
Fig. 1). Presumably, much more species of fungi,
characteristic for the biogeographic region, will
be discovered by future research which would
be in line with research that up to now has been
conducted on other groups of living organisms
Tab. 2: Evaluation of Dinaric karst poljes based on ascomycetous fungi. In addition, protected species and/or species which were used for the present evaluation are shown
Polje name Geographical orientation Country Size class Spec. no. Res. days CES CES/sp CES/day Exclusive polje’s habitat of special value to mycobiota Protected and/or evaluated species
Ponikve Mt. Medvednica, NW from Zagreb HR micro 28 16 9,5 0,34 0,59 none Helvella phlebophora (HR: VU)
Ponikve Tršće, NW from Delnice HR micro 9 3 24 2,67 8 bog Vibrissea truncorum (HR: VU),
Heyderia abietis (HR: NT)
Sungerski lug Sunger, SSE from Delnice HR small 46 24 70,5 1,53 2,94 small bogs surrounded by tall old growth coniferous forest
Discina fastigiata (HR: CR),
Caloscypha fulgens (HR: VU),
Geopyxis majalis (HR: VU),
Trichoglossum hirsutum (HR: VU),
Peziza obtusapiculata (HR: DD),
Heyderia abietis (HR: NT)
Bijambare Nišići, NNE from Sarajevo BA small 23 13 49,5 2,15 3,81 bog, old growith coniferous altimontane forest
Cudonia circinans (HR: VU),
Heyderia abietis (HR: NT),
Ascobolus xylophilus (BA: CR),
Plicaria trachycarpa (BA:VU),
Patinella hyalophaea (BA candidate: EN),
Podostroma leucopus (BA: candidate EN)
Ogulinsko-plaščansko polje Ogulin HR large 6 2 21 3,5 10,5 fens and riparian forest in mediterranoid continental belt Mollisia uda (HR candidate: EN),
Scutellinia peloponnesiaca HR candidate: VU)
Livanjsko polje Livno BA large 5 2 8 1,6 4 fens, two types of riparian forests, swamp, traditionally managed grassland Myriosclerotinia sp. (M. dennisii HR: CR)
Duvanjsko polje Tomislavgrad BA large 4 1 6,5 1,63 6,5 various oro-Mediterranean brooks, traditionally managed grassland Helvella corium (BA: EN),
Parascutellinia carneosanguinea (BA: VU)
Šujičko polje Šujica BA medium 3 1 1 0,33 1 tall riparian forest none
Dugo polje Blidinje jezero BA medium 14 4 26 1,86 6,5 fens and Pinus heldreichii endemic (alti-Mediterranean) forest, traditionally
managed grassland Helvella albella (HR: CR),
Boudiera tracheia (BA: EN)
Gornja i donja bara pl. Čvrsnica BA micro 3 2 8,5 2,83 4,25 fens in alti-Mediterranean belt Boudiera tracheia (BA: EN)
Lamprospora leptodictya (BA candidate: EN)
Konavosko polje Čilipi HR large 33 17 73,5 2,23 4,32 karstic springs and riparian forest in Mediterranean belt Geoglossum cookeianum (HR: VU),
Geoglossum umbratile (HR: VU),
Sarcoscypha macaronesica (HR: VU)
44 DINARIC KARST POLJES
In 2017, in particular during the research in the
karst poljes of the Central Dinarides (groups G1
and G4), we encountered climatic extremities
never before recorded in the area since the
beginning of meteorological observations. For
several months there was a severe drought
with high temperatures and frequent strong
winds. However, even under such harsh
conditions our research was surprisingly fruitful.
In particular, in sheltered and shady stands
with high substrate humidity we found many
fungi. The occurrence of drought intolerant
ascomycetes in karst poljes is mainly owed to
their specific hydrological features. Large water
bodies in karstic underground that feed surface
watercourses in structural polje’s bottoms are not
so susceptible to drying out as dense networks of
watercourses over watertight bedrock (cf. Bonacci
2015). Simultaneously, a number of Pannonian
permanent rivulets and large fishponds outside
of the Dinaric karst dried out completely. Even
the smallest karst polje often generates its own
microclimate with temperature inversions and the
retention of fog (cf. Gams 1978) as an additional
ecological influence that is often reflected in
an “inverse” zonation of vegetation (Horvat
1953). Therefore, wet structural karst poljes have
tremendously high capabilities for the retention
of substrate moisture which is important for
fungi in a number of habitat types, e.g. a variety
of freshwater habitats (especially swamps and
fens) and their inundation zones and in riparian
old growth forests (in particular, stands of Salix
spp., Fraxinus angustifolia, Alnus glutunosa and
in Adriatic Quercus robur, Q. frainetto and Dinaric
Q. cerris tall forests; cf. Jovanović et al. 1986).
Beside numerous epigean habitat types present
in structural poljes typically there are also
permanent cave springs, artesian (vauclusian)
karstic springs and ponors situated at the polje’s
edges that play an important role not only for
water circulation (Bonacci 1987a), but also (like
during certain geological periods) for many
fungal groups. In karst areas owing to long and
intensive corrosive processes large subterranean
spaces developed which provided a shelter
for many fungi during several ice ages. Many of
those species evolved afterwards under specific
ecological conditions (constant and complete
darkness, high humidity, low but above zero
temperatures, poor nutrition levels) into true
cave organisms incapable of resettling surface
habitats after the withdrawal of the ice and
climate warming (Matočec et al. 2014).
Following to current global climate warming,
modern observers will probably witness consid-
erable changes of karst polje’s ecosystems. It
can be expected that the magnitude of change
will differ between karst poljes (some poljes may
show more environmental shifts while others
may be more resilient to environment change). In
that case differences between certain poljes will
probably remain comparably high over longer pe-
riods of time. Measurements of the hydrological
regime of Vrgorsko polje (Jezero) between 1926
and 2006 (Bonacci 2013) may illustrate certain
climatological trends. A number of fungi from
different taxonomic and ecological groups have
been ascertained as good bioindicators of cli-
mate shifts (e.g., Ing 2005, Kauserud et al. 2009,
Buentgen et al. 2013, Ohenoja et al. 2013). This is
confirmed also by our own monitoring data some
of it covering a period of more than 30 years.
The general ecological importance of karst poljes,
the international significance of the large Dinaric
karst poljes, their natural and cultural values as
well as their need for protection is unquestion-
able (Bonacci 2015, Sackl et al. 2014). Following to
considerable morphological, climatological and
hydrological differences between the karst poljes
of the Dinaric Alps, the species diversity and im-
45
NATURE CONSERVATION AND RURAL DEVELOPMENT
portance of different karst poljes for mycobiota
differs, according to the present assessment,
heavily between the different groups of poljes in
Tab. 2. In this respect we can draw the following
general but preliminary conclusions:
1) The overall importance for mycobiota (here
modelled for ascomycete fungi) is greater
for karst poljes (especially of those with
continual fluvial inflow) situated in regions
which are exposed to longer period of
drought and/or higher temperatures, and a
precipitation deficiency during the warmer
half of the year (resulting in the intensive
and fast drying of the substrate).
2) The importance of the Dinaric karst poljes
for general diversity of ascomycete fungi,
the number of rare as well as aquatic/
subaquatic species is greater if the climate
at the polje’s bottom is more humid and
when the polje harbours some freshwater
waterbody (ponornica, spring, lake, swamp,
bog or fen).
3) The importance of Dinaric karst poljes for
regional/local ascomycete mycobiota is
greater if the polje is situated farther from
the karst area edge or is more isolated from
the continuous area of the Dinaric karst on
the islands in the Adriatic Sea. Structural
poljes of the central Dinaric karst or in
the Adriatic lowlands and micro-poljes on
the Adriatic Islands are more pronounced
“oases” than randpoljes situated along
the continental edge of the Dinaric karst.
They make more contrasts in ecological
conditions and habitat types favourable to
most fungi in relation to surrounding dry
karstic massifs and plateaus.
4) The overall importance of Dinaric karst
poljes for ascomycetes is not correlated
with polje size unless the polje houses a
high number of different habitat types.
5) Small micro-poljes situated in higher
altitudes in semi-alpine or alpine areas,
especially those poljes which are partly
influenced by Mediterranean or sub-
Mediterranean climates, are particularly
important for ascomycetes because these
karst poljes harbour arctic-alpine species.
Our preliminary analyses show that many habitat
types of the highest importance for fungi in the
vast area of the Dinaric karst are largely restricted
to the bottoms of karst poljes. Regardless of
their size and the bioclimatic zone these habitat
types are stable in many karst poljes and provide
the basis for high habitat diversity and species
diversity of mycobiota in the poljes. The most
significant habitats for fungi are periodical or
perennial freshwater habitats, like ponornicas,
riparian old growth forests, bogs, fens, swamps,
lakes, karst springs, humid cave entrances etc. In
addition, traditionally used grasslands, artificial
ponds and old, shady and wet foot trails are of
great importance for mycobiota as well. These
habitats should be protected and carefully
managed by, e. g., traditional and sustainable
forms of land use or by eco-tourism.
In particular, the mycobiota of karst poljes with
low exploration level but high “ecological scores”
(Tab. 2) as well as some poljes that were not yet
studied, should be intensively investigated in the
future. This is especially important in wet poljes
with unique, rare and widely endangered habitats
(e.g., poljes of the groups D1, E1, E2, G1, G3 and G4).
Because human interests often have negative
effects for the biodiversity of karst poljes (cf.
Bonacci 1985) this research will be a significant
contribution to the regional and international
Important Fungus Area system (cf. Anderson
2002, Tkalčec et al. 2005, Jukić & Omerović 2017).
Thanks to the usefulness of different myco-
46 DINARIC KARST POLJES
bioindication methods, calibrated for regional
ecological conditions (e.g., Matočec 2000, Matočec
et al. 2000, Alebić-Juretić & Arko-Pijevac 2005,
Ódor et al. 2006, Ozimec et al. 2013a, Kušan &
Matočec 2017, 2018) the results of such research
may also help in planning and implementing
regional sustainable economies.
Acknowledgements
The authors wish to thank the NGO “Naša
Baština” from Tomislavgrad, Bosnia and Her-
zegovina, and NGO “ADIPA”, especially Roman
Ozimec for the organisation of the expeditions
in the vicinity of Tomislavgrad. The expeditions
provided us with the valuable opportunity for in-
tensive mycological research in the remote areas
around Tomislavgrad, Livno and in Blidinje Nature
Park. We further thank Mićo Šarac for his guid-
ance and for transportation to several important
localities on Mt. Čvrsnica. We also appreciate
the help by Smiljan Tomić with fieldwork and the
collecting of fungal material. The second author
wishes to thank the Rufford Small Grant Founda-
tion for financial support during some of the field
investigations.
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... These authors observed similar climate and vegetation conditions of their collection site (coastal eastern Canada) and the type locality in montane northeastern Italy, including Fagus as the dominant forest tree species. Baral and Carter [84] also suggested an association with semi-aquatic habitats subjected to occasional flooding, which is in line given the recent report of P. hyalophaea from a torrent watercourse in a bog complex of an altimontane karst polje in the Dinaric Alps [92]. Patinella hyalophaea was isolated from historic wood in Deception Island, Antarctica [93], and most recently from buried wooden artifacts at five sites in Western Greenland and lacustrine sediment cores from a lake in King George Island, Antarctica [94,95]. ...
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