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ORIGINAL ARTICLE
Distribution and ecological preferences of the species of the family
Athericidae in three hydrobiological ecoregions of Central Europe
Eva Bulánková
1
&Jan Špaček
2
&Pavel Beracko
1
&Igor Kokavec
3
Received: 11 December 2018 /Accepted: 19 March 2019
#Institute of Zoology, Slovak Academy of Sciences 2019
Abstract
From the family Athericidae, only three species (Atherix ibis, Ibisia marginata, Atrichops crassipes) are widespread in Central
and Western Europe. Although predatory larvae of these species are an importantcomponent of many benthic communities, little
is known about their distribution and ecological preferences. Aiming to fill these gaps, the distribution and ecological preferences
of these species were studied in three ecoregions of central Europe (Central highlands, The Carpathians, Hungarian lowlands).
We found that A. ibis was present in the most streams in all of the studied ecoregions. I marginata clearly preferred the streams of
the Carpathians ecoregion, whilst A. crassipes was more frequent in the Central highlands and Hungarian lowlands and it
occasionally inhabited streams in the Carpathian ecoregion. The occurrence of the species was explained by the physico-
chemical parameters of water (explained variability = 18.6%), site characteristics (3.8%), as well as catchment characteristics
(3.3%). Four environmental variables (temperature, conductivity, percentage of agricultural land, catchment area) from three
defined groups represented significant gradients, which explained species distribution in running waters of Central Europe. For
the Central European streams, a correction of the saprobic index of the species was made, and the altitudinal, temperature, current
and pH preferences for each species were also calculated. These values can be used for completion of the Bfreshwaterecology.
info^database, which includes several biological and ecological traits of most European benthic species.
Keywords Atherix ibis .Ibisia marginata .Atrichops crassipes .Ecological parameters .Saprobity
Introduction
Athericidae is a small family of aquatic true flies (Diptera) with
only ten species in four genera that occur in Europe (Rozkošný
2007). Only three species are known in the Czech Republic and
Slovakia, as well as from several other European countries:
Atherix ibis (Fabricius, 1798), Ibisia marginata (Fabricius,
1781) and Atrichops crassipes (Meigen, 1820). Occurrence of
these species covers a wide range of habitats across Europe
except Iceland, Tajga and the Caspic depression. A. ibis and
I. marginata occur in most European hydrobiological ecoregions
definedbyIllies(1978), whereas A. crassipes is mainly distrib-
uted in the ecoregions of Central and Western Europe (Appendix
1(Table 4)). Some data about the occurrences and ecology of
these species of the family Athericidae from former
Czechoslovakia were published by Rozkošný and Spitzer
(1965), Tuša(1993,1994), as well as from Slovakia by
Bulánková and Ďuričková (2009).
Thomas (1974,1975) described the nomenclature, morphol-
ogy and feedings habits of adults and larvae of all three species
for the first time. He also determined the principal factors lim-
iting the distribution of these species based on 237 samplings
sites in the south of France. Most of scientific papers relating to
A. ibis have so far been focused on the taxonomic position,
zoogeographical distribution, some habitat preferences and
adult feeding habits (Nagatomi 1962,1984a,b;Nagatomiand
Rozkošný 1997;Thomas1974,1975,1976,1997;Tuša1994;
Malmqvist 1996). Many papers have been published about the
occurrence of A. ibis females and their clusters in Europe
(Itämies et al. 1990,1993; Buttstedt et al. 2001;Feldmann
2010;Möller2010;M
adsen2012).
*Pavel Beracko
beracko@fns.uniba.sk
Jan Špaček
spacekj@pla.cz
1
Department of Ecology, Faculty of Natural Sciences, Comenius
University, B-2 Mlynská dolina, SK-842 15 Bratislava, Slovakia
2
River Elbe Board s. e., Víta Nejedlého 951, Hradec Králové 500 03,
Czech Republic
3
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta
9, SK-845 06 Bratislava, Slovakia
Biologia
https://doi.org/10.2478/s11756-019-00244-9
Predatory larvae of Athericidae have an important impact
on the higher and lower trophic levels in aquatic ecosystem,
and they are also considered as good bio-indicators of the
saprobity level (Sládeček 1973), acidity (Braukmann 2001)
and stream habitat degradation (Lewin et al. 2014). Heino
and Mikrä (2006) also showed high affinity of these species
to stream size. To date, the ecology of I. marginata and
A. crassipes has been scarcely studied. According to current
knowledge, I. marginata is less tolerant to organic pollution
and prefers small colder streams (Thomas 1976;Tuša1994;
Bulánková and Ďuričková 2009; Jentzsch and Kleinsteuber
2012). Larval development of I. marginata under natural con-
ditions was described forthe first time by Vaňhara (1975). The
most comprehensive data about larval development and
bionomy of A. crassipes was published by Gerke and
Böttger (2001). In the last decades, some papers have been
published about the first record of A. crassipes in Hungary
(Murányi et al. 2009), Switzerland (Sartori et al. 2011)and
Portugal (Andrade 2014). No paper relating to the ecological
preferences of A. crassipes has been published till now.
The species traits database for freshwater organisms named
Bfreshwaterecology.info^(Schmidt-Kloiber and Hering 2015)
, joins together data from several scientific sources into a
comprehensive overview about the distribution and
ecological preferences of the species occurring in Europe.
For the family Athericidae, there are only a few records
about the ecological preferences of the species. Based on the
current knowledge about the three mentioned species of the
family Athericidae, the aims of our study were: i) to describe
the distribution of three species of the family Athericidae in
the three studied ecoregions, ii) to identify responsible
environmental variables that determine the occurrence of the
species, as well as the composition of the Athericidae
assemblage in streams, iii) to calculate some ecological
indices and preferences of the Athericidae species, which
may be useful for the freshwaterecology.info database
(Schmidt-Kloiber and Hering 2015).
Material and methods
Study area and sampling sites
The study area and sampling sites of the species of the
family Athericidae took place in three ecoregions (Fig. 1)
sensu Illies (1967,1978): ecoregion 9 - Central highlands,
ecoregion 10 - The Carpathians, ecoregion 11 - Hungarian
lowlands. According to AQEM:STAR stream typology
(Hering et al. 2004), the following typed of streams were
investigated: A04 Mid-sized streams in the Bohemian
Massif and COl Mid-sized streams in the central sub-
alpine mountains (ecoregion 9), COl Mid-sized streams
in the central sub-alpine mountains and C03 Mid-sized
streams in the Carpathians (ecoregion 10), AOI Mid-
sized streams in the Hungarian Plains (ecoregion 11).
Large sub-mountain and lowland rivers were also includ-
ed in the investigation of the species distribution and pref-
erences in the three studied ecoregions.
Sampling procedure
Altogether, we evaluated distribution of the species of the family
Athericidae at 770 localities in the „Central highlands
Becoregion, and at 99 localities in the „Carpathians Becoregion,
as well as at 26 localities in the „Hungarian lowlands
Becoregion. In the Czech Republic, larvae of the family
Athericidae as part of benthic samples were taken within a na-
tional hydrobiological survey from 2007 to 2014. According to
the Czech national sampling scheme PERLA (ČSN 757701
2008), the 3-min long multi-habitat sampling is used to obtain
a benthic sample. In Slovakia, larvae of the family Athericidae
were sampled according to the STAR-AQEM method (Hering
et al. 2003), which involves the sampling of macroinvertebrates
from microhabitats with a coverage higher than 5%. These sam-
plings were performed as a part of several hydrobiological re-
search studies provided from 2001 to 2017.
Environmental variables
According to the Czech national scheme, basic physico-
chemical measurements and hydromorphological evaluations
are performed at each sampling site (Table 1). The physico-
chemical parameters of water were measured using a WTW
Multi 3240 portable device with a multi-parametric probe YSI
600 LM, or by using a WTW pH/cond 340i and Hach Lange
HQ4D portable multimeter. Total phosphorus and nitrogen were
measured by SP100 BOD analyzer and SP2000 BOD analyzer
(Skalar Analytical B.V. Instruments). A 5-day biochemical ox-
ygen demand (BOD 5) was measured by Continuous Flow an-
alyzer (AMS Alliance Instruments). Site characteristics were
recorded directly at the sampling site or they were found later
using the GISyPoNET application (http://igis.pla.cz). Catchment
land use was derived from the CORINE land cover database in
ARGIS 9.3 (ESRI 2011). Land use was expressed as the per-
centage of seven principal land use types (Forest, Shrubs,
Meadows and pastures, Orchards, Agricultural land, Tilled land,
Urban). In the Central highlands, 211 localities had a complete
dataset of the variables given in Table 1, therefore, only these
sampling sites were included in the further statistical analyses in
the given ecoregion.
Data analyses
Multivariate analyses were performed with CANOCO 5 soft-
ware (Ter Braak and Šmilauer 2012). First, the Variance
Inflation Factors (VIFs) were inspected. High values for the
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VIF (>20) indicated multicollinearity between some variables
and this should be avoided (Ter Braak 1986). Hence, the var-
iable with the highest VIF was removed in the next analysis.
Second, for assessment of the relative importance of
physico-chemical data versus site characteristics versus catch-
ment characteristics for the occurrence of the species, we used
Variation Partitioning in the Bdistance-based^Redundancy
Analysis (db_RDA) with Bray-Curtis distance matrices. This
method allowed the decomposition of the variance of the spe-
cies matrix among sets of explanatory variables in order to
identify their pure and shared contributions to the total vari-
ance (Borcard et al. 1992; Legendre and Legendre 1998).
Third, for explanation of the variation in the species data, we
performed three different db_RDAs with the forward selection
method of significant factors involved at each of the three hier-
archical levels of acting of environmental conditions (Fig. 2):
run 1) db_RDA of species data constrained by physico-chemical
variables, run 2) db_RDA of species data constrained by
physico-chemical variables + site characteristics, run 3)
db_RDA of species data constrained by physico-chemical vari-
ables + site characteristics + catchment characteristics. The re-
sponse curves, which explained the probability of species occur-
rence along the gradient of significant continuous environmental
parameters in db_RDA, were generated in a generalized linear
model (GLM) with a Bquasi-logit^distribution function. The
Blogit^distribution function in the GLM was applied for testing
of the effects of the significant categorical variable from
db_RDA. Non-significant effects of the variable were eliminat-
ed by step-by-step merging procedure. Univariate analyses were
performed in the base package of the R 3.2.2 software environ-
ment (R Core Team 2016). Species distribution maps were cre-
ated in ArcGIS software version 10.4 (ESRI 2016).
Calculation of some ecological metrics (saprobic valence,
saprobic index and current preference) for all three species
was based on the result of Asterics 4.04 software downloaded
from the web site Bhttp://www.fliessgewaesser-bewertung.de^.
The database consisting of the benthic macroinvertebrate
community composition at each site was imported into
Asterics 4.04. As a result of this, we received a proportion of
the species in each saprobic/current preference class. The pro-
portion in each class was multiplied by the species abundance
and then the sum for each saprobic/current preference class was
calculated. The proportions of these sums on the total sum were
recalculated into a 10-point scale (Zelinka and Marvan 1961;
Moog 1995). The saprobic index Si was calculated for each
species according to the equation (Pantle and Buck 1955):
Si¼1*SC1 þ2*SC2 þ3*SC3 þ4*SC4 þ5*SC5
=10;
where SC1 –SC5 were the preference points of the species (on
a 10-point scale) to the saprobic class in the order from
xenosaprobity to polysaprobity. Indication weights ranging
Fig. 1 Map of ecoregions according to Illies (1967,1978) (modified from European Environment Agency 2004). Dark grey polygon represents the
sampling area
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from 1 (for the eurysaprobic species) to 5 (for the stenosaprobic
species) were identified for each species using the approach of
Sládeček et al. (1981).
Other ecological parameters (pH preference, temperature pref-
erence and altitudinal preference) were calculated directly from
field measurements of the environmental parameters. The altitu-
dinal categories were based on Ellenberg classification for low
mountain regions (Ellenberg 1996). Temperature and pH catego-
riesreflectedthecategorizationusedinthefreshwaterecology.
info database. Prior to the calculation, the number of occurrences
in each category was standardized at 100 sampling sites. Affinity
of the species to each category of the parameter was designed
based on the number of occurrences in each category. In the chi–
squared goodness of fit test, we tested the observed number of
occurrences in each category against a null hypothesis: Bthere is
the same number of occurrences in each category^.
Results
Species distribution and environmental factors
In the studied ecoregions, Atherix ibis was present at the most of
sampling sites situated in the Central highlands (576 sites) and
Hungarian lowlands (19 sites) (Fig. 2). In the Carpathians
ecoregion, the number of occurrences in A. ibis (57 sites) was
similar to Ibisia marginata, which occurred there at 62 sites
(Fig. 3). In the Central highlands, I. marginata occurred in one
third of the sampling sites (216 sites), while its presence was only
occasional in the Hungarian lowland ecoregion. Atrichops
crassipes occasionally inhabited streams in the Carpathian
ecoregion, and was found only at five sites there. A. crassipes
was more frequently present in the Central highlands (85 sites)
and Hungarian lowlands (9 sites) (Figs. 4and 5).
In the db_RDA with variation partitioning, the physico-
chemical variables accounted for 18.6% of the total variation
(46.2% of explained variability) of the Athericidae communi-
ty composition (Fig. 6). Together, site characteristics and
catchment land-use explained 17.6% of the explained variabil-
ity, with a similar proportion of each component. The highest
proportion of the shared effects was recorded between
Fig. 2 Scheme of the running Bdistance-based^Redundancy Analyses explaining the presence of three species of the family Athericidae in relation to
factors acting on three hierarchical levels
Table 1 Environmental variables measured and evaluated at the
sampling sites and in the river basins
Variables Min. Max. Mean Median
Maximal summer water temperature 5 19.3 10.13 9.9
pH 5.42 8.67 7.81 7.85
Dissolved oxygen 5.6 14.4 9.69 9.7
Oxygen saturation 49.3 133.4 87.08 88.65
Conductivity 24 1656 455.17 393.4
BOD5 1 8.9 2.73 2.35
Total phosphorus 0.4 35 4.23 3.6
Total nitrates 0.01 4 0.15 0.1
Present Absent
Algae 39 172
Moss 69 142
Floating roots 51 160
Artificial banks 17 194
Range (%) Present
Continuous line trees 10–100 126
Isolated trees 10–50 52
Shrubs 10–100 101
Meadows/pasture 10–60 48
Tilled land 10 1
Suburban 10–100 76
Suburban –Buildings 10–100 118
Agricultural land 20–100 86
Forest 10–100 50
Orchard 10–100 9
Natural/seminatural open land 20–100 21
12 3
Bridge 68 98 45
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physico-chemical variables and site characteristics, which
accounted for 17.1% of the explained variability in the com-
position of the Athericidae community.
Four environmental variables represented significant
gradients that explained the structure of the Athericidae
community. The probability of occurrence of A. ibis clear-
ly increased with higher mean summer water temperature,
unlike I. marginata, where the probability of its occur-
rence decreased with higher mean summer water
temperature (Fig. 7). The probability of occurrence
A. crassipes increased with higher conductivity and
higher proportion of agricultural land in the catchment
area. A significantly higher probability of the occurrence
of A. ibis and A. crassipes was in the medium/large river
basin (no significant difference between large and medi-
um), rather than in smaller river basins (Fig. 8). In
I. marginata, significant differences in the probability of
occurrence were observed among all three scales of the
Fig. 3 Distribution of Atherix ibis in the Central highlands, Carpathians and Hungarian lowlands ecoregions
Fig. 4 Distribution of Ibisia marginata in the Central highlands, Carpathians and Hungarian lowlands ecoregions
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river basin, with decreasing probability of occurrence
from small to large river basins.
A hierarchical model of ordination setting showed that
at the first level (physico-chemical variables), temperature
and conductivity were the factors that significantly affect-
ed the composition of the Athericidae community
(Table 2). No environmental factor acting at the second
level (site characteristics) added the significant informa-
tion to explained variability at the first level. At the third
level (catchment characteristics), the catchment area and
proportion of agricultural land were the significant factors
that contributed to the increase of the explained variability
performed by factors acting at the first two lower levels.
Species preferences and ecological indices
In affinity to the organic pollution, all three species had the
optimum of their occurrence in beta-mesosaprobic habitats
(Table 3). However, I. marginata and A. ibis inhabited more
frequently less-organically polluted streams at the
oligosaprobity level, while A. crassipes generally tolerated
alpha-mesosaprobic conditions. In altitudinal distribution,
I. marginata mainly preferred streams in sub-montane and
montane zones, while A. ibis was widespread in streams
from planar to montane zone. A. crassipes can occasionally
be found in streams of higher altitudes, although it predom-
inantly inhabited streams and rivers of the planar zone.
Considering pH preference, all three species have affinity
to neutral and alkaline water. Temperature preference of the
species linked with maximal water temperature during
summer showed that A. ibis and A. crassipes clearly pre-
ferred warmer habitats compared to I. marginata.
Nevertheless, A. ibis was usually found in streams with
low as well as with high summer water temperature. The
occurrence of I. marginata was clearly associated with cold
and very cold streams, and it was never found in warmer
habitats. In current preference of the species, A. ibis and
I. marginata could be classified as rheophilous taxa, while
A. crassipes preferred slow flowing lowland streams and
rivers. In fast flowing streams, it was found only in the
marginal dead waters or pools.
Fig. 5 Distribution of Atrichops crassipes in the Central highlands, Carpathians and Hungarian lowlands ecoregions
Fig. 6 Result of the variation partitioning analysis showing the relative
influence of: i) unique effect of physico-chemical, site and catchment
characteristics, ii) shared effects of the three above-defined sets of
environmental variables on the Athericidae assemblages in the Central
highlands, Carpathians and Hungarian lowlands ecoregions.
Abbreviations: Uco –unconstrained axes
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Discussion
Species distribution and environment
All three common European snipeflies species occurred in the
territories of the Slovak and the Czech Republic as well. In
comparison to previous studies performed by Rozkošný and
Spitzer (1965), Tuša(1993), Bulánková and Ďuričková (2009),
Bulánková (2011), Mišíková Elexová et al. (2010), Varecha and
Fehérová (2012), our study brought many new records of the
presence of the species in Bohemia and Slovakia. Many of these
records have been represented by unpublished data obtained dur-
ing monitoring and scientific programmes running over the last
decade. Therefore, maps of the distribution of each species found
in the Czech and Slovak Republics represent the current state of
species distribution in these countries. Moreover, these countries
are situated in three different hydrobiological ecoregions on a
relatively small area with various aquatic habitats, which allowed
us to re-evaluate their ecological preferences as well.
In this study we found that the physico-chemical parameters
of water limited the occurrence of the examined snipe flies spe-
cies much more than the site and catchment characteristics. At
the lowest hierarchical level (physico-chemical parameters of
water), only water temperature and conductivity were the
significant factors explaining the presence of the species. In
A. ibis, the probability of occurrence increased with the increas-
ing maximal summer water temperature, whilst in I. marginata,it
was the opposite. The preference of I. marginata for lower water
temperature compared with other species was observed by sev-
eral authors (Thomas 1976; Jentzsch and Kleinsteuber 2012).
Colas et al. (2014) found that the A. ibis population was favoured
by the conditions below reservoirs, especially when warmer wa-
ter is released (Kokavec et al. 2017). Although A. crassipes
showed preference to warmer lowland habitats, which is also
documented by Gerke and Böttger (2001) and Daufresne et al.
(2003), its presence was clearly influenced by the value of con-
ductivity. In the studied ecoregions, the habitats with conductiv-
ity higher than 800 μScm
−1
had a probability of A. crassipes
presence over the 50%. In Western Plain ecoregions, A. crassipes
was found in watercourses with low conductivity, maximally up
to 700 μScm
−1
(Lock et al. 2014). Although the correlation
between the occurrence of the other two athericid species and
conductivity changes was very weak, some studies, for example
Kazanci and Dügel (2010), found a strong negative relation be-
tween conductivity and abundance of A. ibis. At the second
hierarchical level (site characteristics), no factors added signifi-
cant more variability to the model based on significant factors of
the first level. Sartori et al. (2011) also found that some site
Fig. 7 Plot of Bdistance-based^Redundancy Analysis with forward
selection of the significant environmental variables. Response curve
explaining the probability of species occurrence along the gradient of
significant continuous environmental parameters in db_RDA were
generated in the generalized linear model Abbreviations: Cond
(conductivity), Temp (water temperature), TL_Agric (Agricultural
land), S (small catchment, M (medium catchment), L (large catchment)
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characteristics, e.g. substrate composition or type of bankside
vegetation, strongly varied between sites with the presence of
A. crassipes. A similar weak affinity to the substratum type
was also found for whole family Athericidae (Thirion 2016).
At the highest hierarchical level of environment acting (catch-
ment characteristics), the catchment area and the percentage of
Fig. 8 Results of testing of a significant categorical factor (catchment area) from Bdistance-based^Redundancy analysis with the forward selection of
environmental variables based on the analysis of generalized linear model with step-by-step merging procedure of non-significant effects
Table 2 Results of Bdistance-based^Redundancy analyses with the forward selection of the significant environmental variables reflecting the
hierarchical levels of the environmental factors (see Fig. 2)
Physico-chemical factors: Total variation = 22.02, explanatory variables explained - 17.1%
Explanatory variables after forward selection Explains % pseudo-F pvalue
Conductivity 12.7 15.9 <0.01
Temperature 4.4 5.7 <0.01
Axis 1 Axis 2 Axis 3 Axis 4
Explained variation (cumulative %) 15.23 17.08 68.56 100
Explained fitted variation (cumulative %) 89.17 100
Physico-chemical factors + Site characteristics: Total variation = 22.02, explanatory variables explained - 17.1%
Explanatory variables after forward selection Explains % pseudo-F pvalue
Conductivity 12.7 15.9 <0.01
Temperature 4.4 5.7 <0.01
Axis 1 Axis 2 Axis 3 Axis 4
Explained variation (cumulative %) 15.23 17.08 68.56 100
Explained fitted variation (cumulative %) 89.17 100
Physico-chemical factors + Site characteristics +Catchment characteristics: Total variation =22.02, explanatory variables explained - 25.5%
Explanatory variables after forward selection Explains % pseudo-F pvalue
Conductivity 12.7 15.9 <0.01
Temperature 3.2 4.6 <0.05
Catchment area 6.1 8.1 <0.01
TL_agriculture 3.5 4.8 <0.05
Axis 1 Axis 2 Axis 3 Axis 4
Explained variation (cumulative %) 23.32 25.5 68.56 100
Explained fitted variation (cumulative %) 91.46 100
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Table 3 Ecological preferences of the three species of the family Athericidae
Saprobity Xenosaprobity Oligosaprobity Beta-mesosaprobity Alpha-mesosaprobity Polysaprobity Indication weight Saprobic index
A. ibis 03 5 2 02 1.9
I. marginata 04 5 1 02 1.7
A. crassipes 02 5 3 02 2.1
Altitude preference Planar Colline Submontane Montane Subalpine Chi-squared Degree of freedom pvalue
A. ibis 1 3 3 3 0 67.2 4 <0.01
I. marginata 0 2 4 4 0 72.2 4 <0.01
A. crassipes 7 3 0 0 0 143.9 4 <0.01
pH preference Acid Neutral to alkaline Indifferent Chi-squared Degree of freedom pvalue
A. ibis 01 0 14 1 <0.01
I. marginata 01 0 21 1 <0.01
A. crassipes 01 0 10 1 <0.01
Temperature preference Very cold Cold Moderate Warm Eurytherm Chi-squared Degree of freedom pvalue
A. ibis 2 2 2 4 0 43.7 3 <0.01
I. marginata 3 4 2 0 0 24.3 3 <0.01
A. crassipes 0 2 3 5 0 30.4 3 <0.01
Current preference Limnobiont Limnophil Limno to rheophil Rheo to limnophil Rheophil Rheobiont
A. ibis 00 1 2 61
I. marginata 00 0 2 62
A. crassipes 01 2 4 21
Preference categories were defined according to the Bfreshwaterecology.info^database
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agricultural land were the determinants adding significant more
explained variability to the model created at the first two hierar-
chical levels. Organic, inorganic and toxic pollution of the
streams, as well as the total degradation of stream are clearly
correlated with the percentage of the agricultural land in the river
basin (Mateo-Sagasta et al. 2017).
In this study, catchment area is one of the most significant
determinants of athericid community structure which can be
closely linked with stream size. Several studies found that
A. ibis preferred large streams (Paavola et al. 2000;Heinoand
Mikrä 2006). Bulánková (1999) found that A. ibis and
I. marginata were typical inhabitants of the metarhithral zone
of streams in the Carpathians region. We found evidence that
the probability of occurrence of A. ibis in streams with large
and medium catchment area was relatively high. Regarding small
river basins, the probability of occurrence of A. ibis and
I. marginata was similar. However, the preference of
I. marginata for small streams was recorded by many authors in
the past (e.g. Thomas 1976;Tuša1994; Bulánková and
Ďuričková 2009). Atrichops crassipes had relatively low the prob-
ability of occurrence in all three large types of river basin, but it
preferred streams with medium to large river basin. This statement
was confirmed by Tachet et al. (2000) and Sartori et al. (2011).
In general, our results also confirmed the facts that the
occurrence of the three studied species is mainly determined
by physico-chemical variables of stream as well as the stream
size (Thomas 1976;Malmqvist1996; Daufresne et al. 2003;
Colas et al. 2014).
Revalidation of ecological preferences and indices
Revision of autecological preferences and indices of the
athericids is essential for an accurate assessment of the ecolog-
ical status of aquatic environments. In the trait database
Bfreshwaterecology.info^, which represents a base for the
assessment of aquatic biotopes in Europe (Schmidt-Kloiber
and Hering 2015), there is insufficient information and
incorrect data about some ecological preferences, as well as
for the saprobic index of the three athericid species in the
Czech Republic and Slovakia. For instance, the value of the
saprobic index for A. ibis ranges from 0.9 to 2 (Schmidt-
Kloiber and Hering 2015). In our results, the value of the
saprobic index for A. ibis (1.9) was very close to the value of
the German saprobic index, although it was greatly different
from the original value of this index for the Czech Republic and
Slovakia, which was 0.9. We are convinced that this difference
was caused by insufficient data when the saprobic index was
counted for the species in the past (Šporka 2003;ČSN 757716
2008). In this study, the generalization of results has a much
higher value due to the large number of sites that accompanied
it. In the database, the saprobic index for I. marginata had a
similar range (from 1 to 1.9) as it was for A. ibis (Schmidt-
Kloiber and Hering 2015). The newly calculated saprobic
index was just slightly higher than its original value given for
the Czech Republic and Slovakia. According to our results, the
species did not occur in xenosaprobic habitats and prefers beta-
mesosaprobic to oligosaprobic conditions. For the A. crassipes
have not specified value of the saprobic index in the database
for any European country. For Central European countries, the
value of the saprobic index for this species was determined to
be equal to 2.1.
In the studied ecoregions, we found that A. ibis occurred from
planar to montane zone i.e. up to 1000 m a.s.l. From the same
ecoregions, similar altitudinal distribution of this species was also
given by Tuša(1994). In the Alps and Pyrenees, the occurrence
of A. ibis reached altitudes up to 2350 m a.s.l., but the maximal
water temperature was extremely high there - about 20 °C
(Thomas 1976). Khamis et al. (2014)assignedAtherix spp. as
indicator taxa of temperature increase in alpine streams of the
French Pyrenees. Based on our results, I. marginata reached the
highest altitude compared to the other two species, and its altitu-
dinal distribution was limited by the presence of broadleaf/mixed
woodland in the river basin. This association is probably given
by eggs laying on the underside of tree leaves. The dominant
coniferous plantation can also be the cause of the absence of
I. marginata in the High Tatras Mts. In the studied ecoregions,
the distribution of A. crassipes was limited to planar and colline
zone which confirmed the previous findings about altitudinal
distribution of this species in other European regions (Thomas
1976; Murányi et al. 2009). In A. crassipes, preference of slow
flowing streams with a current speed of approximately 0.2 to
0.4 m/s (Tachet et al. 2000; Sartori et al. 2011) reflected our
findings, where this species mainly preferred slow flowing and
lentic zones of lowland brooks and rivers. Based on our results, I
marginata and A. crassipes preferred neutral to alkaline condi-
tions, which fully corresponds with the statement that these spe-
cies are sensitive to acidification (Braukmann 2001; Nagatomi
and Stuckenberg 2004). In our research, although A. ibis was
foundmainlyinnon-acidifiedhabitats, it sometimes tolerated
low acidification as well (Thomas 1997). Whereas some prefer-
ences in the studied species are still absent in the
freshwaterecology.info database (Schmidt-Kloiber and Hering
2015), outputs of this research can be useful for gradual
completion of this database.
Acknowledgments We would like to thank two anonymous reviewers for
their constructive comments, which helped us to improve the quality of
the manuscript. This work was supported by the Slovak Scientific Grant
Agency (Project No. 1/0119/16).
Compliance with ethical standards
Ethical approval This article does not contain any studies with animals
performed by any of the authors.
Conflict of interest The authors declare that they have no conflict of
interest.
Biologia
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Ecoregion Atherix ibis Ibisia marginata Atrichops crassipes
ER1 Ibero Macaronesian *
1
*
1
*
1
Er2 Pyrenees *
3
*
3
*
3
ER3 Italy, Corsica, Malta *
4
*
5,6
*
5
ER4 Alps *
1
*
1
ER5 Dinaric western Balkan *
7
*
8
ER6 Hellenic Western Balkan *
9
*
2
ER7 Eastern Balkan *
10
ER8 Western Highlands *
1
*
1
*
1
ER9 Central Highlands *
1
*
1
*
1
ER10 The Carpathians *
1
*
1
*
1
ER11 Hungarian lowlands *
1
*
2
*
11
ER12 Pontic province *
2
ER13 Western plains *
1
*
1
*
1
ER14 Central plains *
1
*
1
*
1
ER15 Baltic province *
12
ER16 Eastern plains *
2
ER17 Ireland and Northern Ireland *
1
*
1
ER18 Great Britain *
1
*
1
*
1
ER19 Iceland
ER20 Boreal uplands *
1
ER21 Tundra *
1
ER22 Fenno-Scandian field *
1, 13
*
14
ER23 Taiga
ER24 The Caucasus *
15
*
16
ER25 Caspic depression
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