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\begin{document}

\title{Species composition of shoreline wolf spider communities vary with
salinity but their diets vary with wrack inflow}



\author[1]{Peter Hambäck}%
\author[2]{Alyssa R. Cirtwill}%
\author[1]{Magdalena Grudzinska-Sterno}%
\author[1]{Alexander Hoffmann}%
\author[1]{Marie Langbak}%
\author[1]{David Åhlén}%
\affil[1]{Stockholm University}%
\affil[2]{University of Helsinki}%


\vspace{-1em}



  
  \date{May 03, 2024}


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\selectlanguage{english}
\begin{abstract}
Wolf spiders are typically the most common group of arthropod predators
on both lake and marine shorelines, because of the high prey
availability in these habitats. However, shores are also harsh
environments due to flooding and, in proximity to marine waters, to
toxic salinity levels. Here, we describe the spider community, prey
availabilities and spider diets between shoreline sites with different
salinities, albeit with comparatively small differences (5 vs. 7%0).
Despite the small environmental differences, spider communities between
low and higher saline sites showed an almost complete species turnover.
At the same time, differences in prey availability or spider gut
contents did not match changes in spider species composition but rather
changed with habitat characteristics within region, where spiders
collected at sites with thick wrack beds had a different diet than sites
with little wrack. These data suggest that shifts in spider communities
are due to habitat characteristics rather than prey availabilities, and
the most likely candidate restricting species in high salinity would be
saline sensitivity. At the same time, species absences from low-saline
habitats remain unresolved.%
\end{abstract}%



\sloppy


\textbf{Title:} \textbf{Species composition of shoreline wolf spider
communities vary with salinity but their diets vary with wrack inflow.}

Hamb\selectlanguage{ngerman}äck, P. A.\textsuperscript{*}, A. Cirtwill\textsuperscript{\#}, M.
Grudzinska-Sterno, A. Hoffmann, M. Langbak and D. Åhlén

\textsuperscript{1}Department of Ecology, Environment and Plant
Sciences, Stockholm University, 106 91 Stockholm, Sweden

*Corresponding author,
email:\emph{peter.hamback@su.se}\textsuperscript{\#}Current address:

\textbf{Abstract}

Wolf spiders are typically the most common group of arthropod predators
on both lake and marine shorelines, because of the high prey
availability in these habitats. However, shores are also harsh
environments due to flooding and, in proximity to marine waters, to
toxic salinity levels. Here, we describe the spider community, prey
availabilities and spider diets between shoreline sites with different
salinities, albeit with comparatively small differences (5 vs. 7\selectlanguage{english}%0).
Despite the small environmental differences, spider communities between
low and higher saline sites showed an almost complete species turnover.
At the same time, differences in prey availability or spider gut
contents did not match changes in spider species composition but rather
changed with habitat characteristics within a region, where spiders
collected at sites with thick wrack beds had a different diet than sites
with little wrack. These data suggest that shifts in spider communities
are due to habitat characteristics rather than prey availabilities, and
the most likely candidate restricting species in high salinity would be
saline sensitivity. At the same time, species absences from low-saline
habitats remain unresolved.

\textbf{Key words} : \emph{Pardosa} , Molecular gut content
analysis,\emph{Fucus} , Shoreline fauna, \emph{Alopecosa} ,
\emph{Arctosa}



\textbf{Introduction}

Shorelines and other riparian habitats are often described as hotspots
for arthropod predators, and spiders in particular seem to thrive in
these habitats (Polis \& Hurd 1995; Mellbrand \& Hamback 2010; Batzer \&
Wu 2020). There are several reasons underlying these high spider
densities, but an important factor seems to be the high prey density in
these near-water habitats (Polis \& Hurd 1995; Sanchez-Ruiz \emph{et
al.} 2018). Prey densities are high in these sites both because of a
direct inflow of insects from the aquatic environment, such as midges
with aquatic larvae and terrestrial adults, and because large inflows of
organic material are deposited on shorelines providing food for
detritivores and fertilizing plants (Colombini \& Chelazzi 2003; Baxter,
Fausch \& Saunders 2005; Hyndes \emph{et al.} 2022). At the same time,
shorelines are often harsh environments due to flooding and wave
disturbance, and on marine shorelines due to a high salinity and a high
turnover of organic material (Defeo \& McLachlan 2013; Barboza \& Defeo
2015). Species diversity on shorelines may therefore be poor,
particularly on marine shorelines where communities often consist of a
range of habitat specialists that can endure high salinity levels (Cheng
1976; Irmler \emph{et al.} 2002).

Despite these general patterns, there is a lack of understanding on how
physical processes and prey availability interact in shaping coastal
arthropod communities. In fact, the spatial variability of arthropod
communities in these habitats is poorly documented compared to inland
habitats. For instance, what differences in the species composition
between limnic and marine shorelines are due to direct effects from a
saline environment and what differences are rather due to differences in
prey communities? Prey communities on limnic shorelines are often
dominated by midges and a range of other taxa (Benke 1998; Delettre \&
Morvan 2000; Salvarina, Gravier \& Rothhaupt 2017), whereas prey
communities on marine shorelines are more dominated by species
developing in rotting wrack beds (Schlacher \emph{et al.} 2017;
Hyndes\emph{et al.} 2022). Similarly, what is the relative importance
from the inflow of dead organic matter versus prey that developed in the
water for shoreline predators? Previous studies suggest that the
importance of these different resources for spiders and other shoreline
predators may vary both between sites, between life stages and over time
(Mellbrand\emph{et al.} 2011; Verschut \emph{et al.} 2019). The diet
analysis of spiders across the season by Verschut et al. (2019)
indicated that adult wolf spiders during early summer on marine
shorelines feed largely on terrestrial dipterans such as dung flies,
which have developed in wrack beds, whereas juvenile wolf spiders later
in season had fed more on aquatic dipterans such as chironomids, where
the larvae had fed on algae or detritus in the water.

To approach these questions, we studied prey communities, spider diets
and spider community structure in a salinity gradient along the Swedish
coast. Using this gradient allows us to explore effects from
comparatively small salinity differences, as the salinity along the
gradient changes continuously from freshwater (\textless{}1\selectlanguage{english}%0) in the
inner parts of the Bothnian Bay to oceanic conditions
(\textgreater{}30\selectlanguage{english}%0) on the western coastlines. In this study, we
included two coastal regions with 5 \selectlanguage{english}%0 and 7 \selectlanguage{english}%0 respectively, where
previous studies have indicated shifting dominance of spider species
(Hamback \emph{et al.} 2016; Verschut \emph{et al.} 2019). We focus our
attention on wolf spiders because these typically dominate the shoreline
predator community in the area (Mellbrand \& Hamback 2010). To account
for the role of marine inflow, we aimed to include sites with and
without thick wrack beds in each region. We also needed to control for
climatic effects because the salinity gradient for our study is also a
latitudinal gradient. For this reason, we used a similarly collected
data set of spider communities on shores by inland waters along the same
latitudinal gradient. Finally, to examine the role of a changing prey
community and spider diet, we estimate prey densities using SLAM traps
and collected spiders for gut metabarcoding in the same sites. Prey
densities and spider diets were estimated two times, to cover seasonal
changes in prey availability and diet differences between adult and
juvenile spiders (cf. Verschut \emph{et al.} 2019).

\textbf{Methods}

\emph{Study sites} : The coastal regions included in the study were (a)
Uppland north of Stockholm with the lowest salinity (\selectlanguage{english}[?]5\selectlanguage{english}%0, northern
region) and (b) Kalmar and Oland in southeastern Sweden with somewhat
higher salinity (\selectlanguage{english}[?]7\selectlanguage{english}%0, southern region) (Fig. 1, Supplementary
Information Table 1). The numbers of coastal sites were 13 (Uppland) and
7 (Kalmar). Among these, two sites respectively had thick wrack beds,
and the other sites were similar but without thick wrack beds and often
with short-cut grass due to grazing. The thick wrack beds had a
thickness of more than 20cm with a considerable extension (several 10s
of meters). The non-wrack sites either lacked wrack almost completely
(as in the Uppland region) or that wrack occurred in scattered patches
and never so thick as to provide a suitable habitat for detritivores (as
in the Kalmar region). The inland regions included 15 and 23 shoreline
sites in Uppland and southern Halland (same latitude as Kalmar)
respectively (Fig. 1, SI Table 1), as part of a broader study focusing
on both insect and spider communities in wetlands.

\emph{Field sampling} : Coastal wolf spider communities were sampled
using 10 pitfall traps per site placed in the wrack (wrack sites) or in
open ground (non-wrack sites) for three nights in early May 2022
(Kalmar) or early-mid June 2021 and late May 2022 (Uppland). Inland wolf
spider communities were sampled during June 2020. These times were
chosen because wolf spiders are then adults or subadults, which
simplifies species identification. Spiders were placed in 70\% ethanol
and brought to the laboratory for identification. Spiders from the
inland sites were identified by R. Vicente and those from coastal sites
by M. Langbak and A. Hoffmann, with assistance from R. Vicente for
complicated cases (mainly involving \emph{Pardosa
agrestis/agricola/monticola} ).

Spiders used for diet analyses were only collected from 13 sites, six
from Uppland and seven from Kalmar, including all wrack sites. Spiders
were individually collected by hand (30 per site), to reduce
contamination risk, at two times (June and August 2019) and directly
transferred to 95\% ethanol. In the lab, samples were placed in a
freezer (-20\textsuperscript{o}C) until DNA extraction and further
processing. Finally, prey densities were estimated by placing one SLAM
(Sea Land Air Malaise) trap for two nights at the same time when
collecting spiders for diet analyses. SLAM trap catches were placed in
70\% ethanol, and brought to the laboratory for sorting to a family
level.

\emph{Diet analyses} : To metabarcode prey content of the hand-collected
spiders, DNA was extracted from either a dissected abdomen (larger
spiders) or the whole specimen (small spiders). To reduce the DNA yield
of the focal spiders, we used a forward primer designed not to amplify
wolf spider DNA (NoSpi2, Lafage \emph{et al.} 2020) in combination with
a general reverse primer (fwhR2n, Vamos, Elbrecht \& Leese 2017) to
amplify a section within the Folmer region of COI (Folmer \emph{et
al.}1994). Procedures for PCR amplification and library building follow
Hamback et al. (2021), and sequencing of the spider samples was
performed in one batch on the Illumina MiSeq3 platform at SciLifeLab in
Stockholm. To detect individual samples after sequencing, a dual tagging
approach was used where the 5'-end of both primers included an 8
base-pair tag (Binladen \emph{et al.} 2007). Illumina-adaptors bearing
unique indices were then ligated to the phosphorylated amplicons without
a PCR step to preclude tag jumping errors (Bohmann \emph{et al.} 2021).
Due to problems with low DNA content, we had to change strategy and add
a second PCR step with a low cycle number (6). Because this additional
step increases the risk of tag jumping errors, we built libraries
separately for each site, using SMARTer ThruPLEX DNA-seq library
preparation kit excluding fragmentation of DNA (Takara Bio), as tag
jumps between spiders within site do not affect the results due to
pooling at this level before analysis. In each library, we also included
at least 25\% empty combinations to estimate tag jumping errors (which
was about 6\%). After sequencing, we used ObiTools (Boyer \emph{et
al.}2016) within the Galaxy Platform (Jalili \emph{et al.} 2020) to
assemble paired end sequences of high quality (score \textgreater{} 40),
trim primers, clean sequences using `obiclean', and demultiplex
resulting sequences to individual samples using `NGSFILTER' after
filtering for size. These procedures resulted in a data set of 367
spider individuals and about 384,600 prey sequences that were grouped
based on 97\% similarity and where representative sequences were
taxonomically assigned using BoLD (Ratnasingham \& Hebert 2007) before
further analyses.

\emph{Statistical analyses} : Spider communities were modelled as the
abundance of each spider species per site in a multivariate analysis
with region, inland/coast, wrack and the region-by-inland/coast
interactions as independent variables using the command manyglm
(package: mvabund, Wang \emph{et al.} 2012) with a negative binomial
error distribution. Prey communities were similarly modelled as the
abundance of major groups in a multivariate analysis with manyglm
between regions with wrack as an independent variable and a negative
binomial error distribution, but these tests additionally included
season (June and August) as independent variable. Finally, the
proportional number of prey sequences (logit-transformed) of major
groups were pooled for each species within site and season, and was
modelled using adonis2 (package: vegan, Oksanen \emph{et al.} 2019). To
compare diet composition between spider species, we also compared gut
contents while controlling for effects of region. To examine model
assumptions, we used plot.manyglm and all tests showed no pattern in
errors which confirm the model appropriateness. Significant
relationships were further explored using anova with adjusted p-values,
to identify which groups that explained the variation. In all these
tests, prey communities and spider diets were included at the level of
family or higher taxonomic unit and not at a species level.

To study prey diversity and diet consistency within and among species,
we first calculated individual diets using the dynamic threshold model
in Cirtwill and Hamback (2021). We then compared species accumulation
curves in spider guts using specaccum with spider individual as sampling
unit (package: vegan, Oksanen \emph{et al.} 2019), and then estimated
diet consistency by calculating the Jaccard similarity index between
diets of individual spiders' prey species and prey families, first
between pairs of all spider individuals and then between individual
pairs of the same species. Diet similarity was compared between region,
wrack, and their interaction, firstly, depending on if pairs included
all spider individuals or where restricted to within species comparison,
and secondly, depending on if diets were based on prey species or prey
family. If the interaction terms did not contribute, models were re-fit
without the interaction. We then tested for pairwise differences between
region-wrack combinations using a Tukey's HSD test applied to the
analysis of variance of the above linear models, including the
interaction term between region and wrack. All tests were performed
using R 3.6.3 (R Core Team 2020).

\textbf{Results}

The analysis on spider communities included 3931 spider individuals
separated into 16 taxa (Fig. 2). The variation in community composition
was explained by a region-by-inland/coast interaction (Wald
statistics=8.3, p\textless{}0.001) and not by the presence or absence of
a thick wrack bed (Wald statistics=4.7, p\textgreater{}0.1). The
region-by-inland/coast interaction arose because of a larger difference
between southern and northern coastal sites compared with southern and
northern inland sites (Fig. 2). When comparing abundances at the species
level (Table 1), four species (\emph{Pardosa agrestis} , \emph{P.
agricola, Arctosa leopardus} and \emph{Alopecosa cuneata} ) were found
almost exclusively at southern coastal sites and three taxa
(\emph{Pardosa prativaga, P. amentata} and \emph{Pirata} spp.
{[}mainly\emph{P. piraticus} {]}) almost never occurred in these sites
but were abundant elsewhere (Fig. 2). In addition, one species (\emph{P.
monticola} ) was mainly coastal whereas another species (\emph{Pardosa
palustris} ) occurred mainly inland, irrespective of region.

The variation in the prey community was explained by a region-by-wrack
interaction (Wald statistics=11.4, p\textless{}0.001) and by season
(Wald statistics=11.6, p\textless{}0.002) (Fig. 3). The region-by-wrack
interaction occurred because Coleoptera (Deviance=14.1,
p\textless{}0.02) and Sciaridae (Deviance=20.3, p\textless{}0.003) were
positively affected by wrack availability only in northern sites,
whereas Dolichopodidae (Deviance=11.5, p\textless{}0.05) was negatively
affected by wrack availability only in southern sites (Fig. 3). The
seasonal effect occurred because Empididae (Deviance=12.4,
p\textless{}0.04) and Homoptera (Deviance=12.8, p\textless{}0.03) were
more abundant during early season in June, whereas Chironomidae
(Deviance=14.7, p\textless{}0.03) and Trichoptera (Deviance=12.5,
p\textless{}0.04) were more abundant during August.

The number of prey species encountered in the gut of spider individuals
varied between one and 15, with an average of 3.9 prey species. The
dominant order in the guts was Diptera, both Brachycera (60\%) and
Nematocera (18\%), with minor amounts of other groups; Homoptera (10\%,
mainly Cicadellidae and Delphacidae), Collembola (4\%), other flying
prey (3\%, Hymenoptera and Lepidoptera), Formicidae (2\%), Acari (2\%)
and Heteroptera (1\%) (Fig. 4, SI Table 2). The diet contents varied
considerably among sites and were mainly explained by wrack
(Lawley-Hotelling trace statistics=6.1, p\textless{}0.001) and season
(Lawley-Hotelling trace statistics=2.6, p\textless{}0.003), with an
almost significant effect from a region-by-wrack interaction
(Lawley-Hotelling trace statistics=1.7, p\textless{}0.07) (Fig. 4).
However, there was no effect of spider species either when including
this variable alone or in combination with other variables, or when run
separately for region. Because of the almost significant region-by-wrack
interaction on gut contents, we repeated the analysis for sites with or
without wrack separately. In this analysis, region was significant for
sites without wrack (p\textless{}0.03) but not for sites with wrack
(p\textgreater{}0.2). The prey groups explaining the region difference
for sites without wrack were Sphaeroceridae (F\textsubscript{1,6}=141,
p\textless{}0.003) and Enchytraeidae (F\textsubscript{1,6}=111,
p\textless{}0.05), that both had a higher frequency in spider guts from
southern sites (Fig. 4). Finally, the comparison between sites with or
without wrack suggested that mainly Sphaeroceridae
(F\textsubscript{1,11}=17.2, p\textless{}0.06) and Heteroptera
(F\textsubscript{1,11}=19.9, p\textless{}0.05) were more abundant in
spider guts from site with wrack.

The species accumulation curves indicated that prey diversity was higher
in southern sites and in sites with no wrack compared with northern
sites and wrack sites (Fig. 5). When comparing diet consistency, we
found that individual spiders had, on average, a Jaccard similarity of
diets = 0.056 (sharing approximately 5.6\% of the prey species consumed
by two individuals). Diet consistency between any pair of individual
spiders varied with region (\selectlanguage{greek}β \selectlanguage{english}=0.025, p\textless{}0.001), wrack
(\selectlanguage{greek}β\selectlanguage{english}=0.080, p\textless{}0.001), and their interaction (\selectlanguage{greek}β\selectlanguage{english}=-0.042,
p\textless{}0.001). The interaction arose because spider from northern
wrack sites had higher diet consistency than spiders from southern wrack
sites, whereas spiders from northern non-wrack sites had lower
consistency than those from southern non-wrack sites (Fig. 6). These
diet similarities were larger when performed for pairs of the same
spider species (Fig. 6) but patterns were otherwise similar (region:
\selectlanguage{greek}β\selectlanguage{english}=0.062, p\textless{}0.001; wrack: \selectlanguage{greek}β\selectlanguage{english}=0.108, p\textless{}0.001;
interaction: \selectlanguage{greek}β\selectlanguage{english}=-0.099, p\textless{}0.001) and when diets were estimated
at the prey family level both for all spider individuals (region:
\selectlanguage{greek}β\selectlanguage{english}=0.038, p\textless{}0.001; wrack: \selectlanguage{greek}β\selectlanguage{english}=0.094, p\textless{}0.001;
interaction: \selectlanguage{greek}β\selectlanguage{english}=-0.059, p\textless{}0.001) and for pairs of the same
spider species (region: \selectlanguage{greek}β\selectlanguage{english}=0.070, p\textless{}0.001; wrack: \selectlanguage{greek}β\selectlanguage{english}=0.136,
p\textless{}0.001; interaction: \selectlanguage{greek}β\selectlanguage{english}=-0.124, p\textless{}0.001).

\textbf{Discussion}

The spider community showed large regional changes along the Baltic Sea
seashore despite comparatively small salinity differences (5 vs. 7%0).
Several spider species (\emph{Pardosa agrestis} , \emph{P. agricola,
Arctosa leopardus} and \emph{Alopecosa cuneata} ) were almost
exclusively located on the higher salinity sites compared to lower
salinity sites by the Baltic Sea shore and by inland lakeshores. At the
same time, other taxa (\emph{Pardosa prativaga, P. amentata} and
\emph{Pirata} spp.) had the opposite distribution pattern, and this
pattern was seemingly not explained by either prey availability or
actual spider diets. In fact, there were no detectable diet differences
between spider species or between spiders captured on shores with
different salinity levels. Instead, spider diets varied between shores
with or without thick beds of stranded wrack, a gradient that did not
affect spider community structure. Consequently, and because the species
shift only occurred on coastal sites and not on corresponding inland
sites, it seems that coastal spider communities are directly affected by
the saline conditions.

High salinity has several negative impacts on spiders and other
arthropods, by reducing both survival and reproduction (P\selectlanguage{ngerman}étillon\emph{et
al.} 2011; Puzin \emph{et al.} 2011; Foucreau \emph{et al.}2012). Even
though none of the species found on the Baltic shorelines can be
considered true halophilic and are usually not found on more marine
seashores (Pétillon \emph{et al.} 2008), it seems reasonable to assume
that species vary in their sensitivity to saline conditions. However,
please note that previous studies on wolf spiders tested the responses
of individuals at a much higher salinity (\textgreater{}30\selectlanguage{english}%0) than in our
gradient, and it is unclear to what extent that their conclusions could
be extrapolated to our study. Irrespective of the mechanisms, our data
in combination with previous studies suggest a gradient in salinity
thresholds of the dominant wolf spider species on marine shorelines in
northwestern Europe where \emph{P. prativaga}typically dominates low
salinity sites, \emph{P. agricola} dominates intermediate salinity sites
and \emph{P. purbeckensis} dominates high salinity sites. The species
abundance distributions of wolf spider communities are often highly
skewed with one dominant species having more than 60\% of all
individuals and a tail of rare species. Even though low salinity sites
are not always dominated by \emph{P. prativaga} , two-thirds are
dominated by this species and then more rarely by \emph{P. amentata} ,
\emph{P. palustris} and some other species (see also Meriste, Helm \&
Ivask 2016).

Whereas the restriction to low salinity sites can likely be explained by
salt sensitivity, the corresponding absence of other species at the same
low salinity sites seems more puzzling. First, it is evident that the
absence from low salinity sites is not absolute as both \emph{Pardosa
agrestis} and \emph{P. agricola} are frequently reported also from
inland habitats in central Europe and more rarely inland also from
northern Europe (GBIF.org). Moreover, studies on \emph{P. purbeckensis}
, perhaps the most halophilic species, suggest that fitness is not
reduced on low salinity sites (Petillon \emph{et al.} 2011). It is
possible that some other habitat characteristics restrict the occurrence
in low salinity sites or that distributions are restricted by species
interactions. Several wolf spider species are known for intraguild
predation of other wolf spider species, at least in the laboratory, and
dominance is mainly governed by size differences (Buddle, Walker \&
Rypstra 2003; Rypstra \& Samu 2005; Rickers, Langel \& Scheu 2006;
Rypstra \emph{et al.} 2007; Turney \& Buddle 2019), but no study this
far has evaluated the role of intraguild predation on the spatial
distribution of wolf spiders.

Whatever the reason is for the difference in wolf spider community
composition, the patterns are not likely explained by different dietary
niches among spider species or by differences in prey availability. Both
this, and previous studies using either molecular gut content analysis
or other methods, indicate large overlaps in the diet of wolf spider
species (Mellbrand \& Hamback 2010; Verschut \emph{et al.} 2019). Diet
differences observed in this study instead seem to depend on whether
spiders were collected on sites with or without accumulated wrack, but
these diet shifts did not coincide with shifts in the wolf spider
community. By far the most abundant prey group in the wolf spider guts
on sites with either wrack or no wrack were dipterans (typically taxa
with smaller individuals) and to some extent homopterans. This general
prey composition of wolf spiders is of course well-known from
non-molecular studies (e.g., Nyffeler 1999), but the relative importance
of small dipterans is perhaps larger in our study habitats. Some
differences between molecular and non-molecular studies may occur
because the former provide an improved representation of small prey
items, which are easily overlooked in non-molecular studies due to more
rapid consumption. In either case, wolf spiders are likely quite
opportunistic predators where prey choice perhaps depend more on
encounter probabilities and catchability of prey in their selected
habitat than on prey qualities. This opportunistic behavior is perhaps
also reflected in the different number of prey species, where the number
is higher in southern sites, as expected, and in sites with no wrack.
Similarly, diet consistency was also higher on wrack sites, and both
patterns observed for wrack sites may reflect that wrack beds are
dominated by a small set of detritivorous species. More surprising was
the higher diet consistency of spiders on southern non-wrack sites
compared with northern non-wrack sites, despite the lower total prey
diversity observed for the spiders in the southern region.

Even though opportunism seems to be a dominant pattern, particularly
dark-winged fungus gnats (Sciaridae) are underrepresented in wolf spider
guts despite their comparatively high occurrence at these sites, similar
to what was found previously (Verschut \emph{et al.} 2019). The reason
for spiders to avoid fungus gnats may be that they represent low quality
food (as suggested by Toft \& Wise 1999b; Toft \& Wise 1999a). Diet
differences between sites with or without accumulated wrack otherwise
reflect availability, even though we refrained from testing the
availability-use relationship due to the bias in SLAM traps. Many small
flies often occurring on wrack beds, such as Drosophilidae, Ephydridae,
Sepsidae and Sphaeroceridae are underrepresented in Malaise type traps
on shore lines because these flies tend not to stick to the ground. In
either case, these small detritivorous flies that likely developed in or
close to the decomposing wrack made up more than 75\% of all prey in
spider guts when collected from sites with heavy wrack beds and the diet
composition was surprisingly similar for spiders collected on northern
and southern wrack beds. More unexpected was perhaps the low frequency
of chironomids in the spider gut contents, particularly in the non-wrack
sites. In a previous study (Verschut \emph{et al.} 2019), not far from
the sites included in this paper, chironomids dominated the spider gut
contents and particularly late in the season. In this study, there were
no seasonal differences and spiders on non-wrack sites instead consumed
a range of terrestrial prey groups, such as Homoptera and various
terrestrial Diptera (Chloropidae, Empididae, Dolichopodidae etc.), and
it seems that spiders were less strongly connected to the nearby marine
environment than previously assumed. In either case, this variability
among studies indicate how dynamic food choice of spiders may be.

To summarize, our study indicates that quite a small difference in
salinity caused the species composition of wolf spider communities to
change almost completely. The mechanism underlying this community shift
is less obvious, both why species disappear in the high salinity and in
the low salinity ends, but we can conclude that prey availability or
differences in the trophic niche between species is likely not involved.

\textbf{Acknowledgements}

This work was supported by the National Genomics Infrastructure (NGI)
and Uppmax through the Science for Life Laboratory, which is funded by
the Knut and Alice Wallenberg Foundation, and the Swedish Research
Council Vetenskapsradet.

\textbf{Data accessibility}

The data that support the findings of this study are openly available in
Dryad at http://doi.org/{[}to be added{]} (Reference to be added)

\textbf{Competing interest statement}

The authors have no competing financial or personal interests that would
conflict with the content of this paper

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Table 1. Marginal deviance and adjusted significances for the abundance
of wolf spider species relative to coast/inland (CI), region (R, north
{[}Uppland{]}/south {[}Kalmar/Halland{]}) and the CI-by-R interaction
(ns = non-significant {[}p\textgreater{}0.1{]}).\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
Species & CI & R & CI*R\tabularnewline
\midrule
\endhead
\emph{Alopecosa cuneata} & 1.4 ns & 0.3 ns & 10.4
(p\textless{}0.02)\tabularnewline
\emph{A. pulverulenta} & 0.1 ns & 2.9 ns & 0.8 ns\tabularnewline
\emph{Arctosa leopardus} & 24.5 (p\textless{}0.001) & 2.9 ns & 8.1
(p\textless{}0.05)\tabularnewline
\emph{Pardosa agrestis} & 11.8 (p\textless{}0.02) & 25.8
(p\textless{}0.001) & 0.2 ns\tabularnewline
\emph{P. agricola} & 25.9 (p\textless{}0.001) & 16.1 (p\textless{}0.005)
& 0.0 ns\tabularnewline
\emph{P. amentata} & 1.5 ns & 1.0 ns & 10.6
(p\textless{}0.05)\tabularnewline
\emph{P. fulvata} & 6.8 ns & 3.5 ns & 5.0 ns\tabularnewline
\emph{P. monticola} & 42.7 (p\textless{}0.001) & 0.7 ns & 0.0
ns\tabularnewline
\emph{P. palustris} & 13.5 (p\textless{}0.005) & 0.8 ns & 5.6
ns\tabularnewline
\emph{P. prativaga} & 0.4 ns & 14.6 (p\textless{}0.006) & 36.6
(p\textless{}0.001)\tabularnewline
\emph{P. pullata} & 2.1 ns & 0.7 ns & 3.2 ns\tabularnewline
\emph{Pirata} spp. & 1.1 ns & 4.6 ns & 13.1
(p\textless{}0.007)\tabularnewline
\emph{Trochosa ruricola} & 3.0 ns & 1.7 ns & 1.7 ns\tabularnewline
\emph{T. terricola} & 2.1 ns & 19.7 (p\textless{}0.001) & 0.0
ns\tabularnewline
\bottomrule
\end{longtable}



\textbf{Figure legends}

Fig. 1. Map showing the location of regions (Northern sites = Uppland,
Southern coastal sites = Kalmar, Southern inland sites = Halland). For
site information see SI Table 1.

Fig. 2. Relative abundance of wolf spider species in inland and coastal
sites in Halland/Kalmar (S) and in Uppland (N).

Fig. 3. Relative abundances of prey catches in SLAM-traps, separated by
wrack occurrence, region (south = Kalmar, north = Uppland) and season (A
= August, S = July). Detritivore flies include Sepsidae, Sphaeroceridae
and Coelopidae. Other flying prey include Hymenoptera and Lepidoptera
but also a range of terrestrial Diptera.

Fig. 4. Relative contents of spider guts from sites with or without
wrack, and in the northern (Uppland) or southern (Kalmar) region. Other
detritivore flies include those connected to wrack, such as
Anthomyiidae, Coelopidae, Ephydridae and Sepsidae. Other large flies
include Dolichopodidae, Dryomyzidae, Empididae, Fannidae, Heleomyzidae,
Hybotidae, Muscidae, Rhinophoridae, Scatophagidae, Sciomyzidae,
Syrphidae, Tabanidae and Tachinidae. Other small flies include
Acroceridae, Agromyzidae, Asteiidae, Canacidae, Carnidae, Chamaemyiidae,
Chloropidae, Lonchopteridae, Opomyzidae, Phoridae, and Pipunculidae.
Other Nematocera include Cecidomyiidae, Ceratopogonidae, Keroplatidae,
Limoniidae, Mycetophilidae, Psychodidae, Scatopsidae and Sciaridae.

Fig. 5. Species accumulation curves (\selectlanguage{ngerman}±S.D.) relative to the number of
sampled spiders for northern sites (Uppland) and southern sites
(Kalmar), with or without wrack accumulation.

Fig. 6. Individual diet similarity estimated as Jaccards similarity
index (±S.E.) separated for region (N=Uppland, S=Kalmar) and wrack
presence. The diet similarity was estimated between all pairs of
individuals (red) or between pairs of the same species (white) and when
prey where included at the species (circles) or family (squares) level.
N-values refer to the number of pairs of individuals.

Figure 1\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

Figure 2\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

Figure 3\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

Figure 4\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

Figure 5\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

Fig. 6\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

Appendix 1. Site information, where spider communities were quantified
in all sites and where diet data and prey availabilities were collected
from a subset of sites.\selectlanguage{english}
\begin{longtable}[]{@{}lllllll@{}}
\toprule
Site & Coast/Inland & Region & Wrack & Diet data & Latitude &
Longitude\tabularnewline
\midrule
\endhead
Barnens \selectlanguage{ngerman}Ö & Coast & Uppland & Y & Y & 59\selectlanguage{ngerman}°55'38''N &
18\selectlanguage{ngerman}°56'29''E\tabularnewline
Forsmark & Coast & Uppland & N & Y & 60\selectlanguage{ngerman}°23'41''N &
18\selectlanguage{ngerman}°13'12''E\tabularnewline
Gudinge & Coast & Uppland & N & & 60\selectlanguage{ngerman}°30'31''N &
17\selectlanguage{ngerman}°59'46''E\tabularnewline
Klungsten & Coast & Uppland & N & Y & 60\selectlanguage{ngerman}°32'44''N & 18\selectlanguage{ngerman}°
1'10''E\tabularnewline
Raggarön & Coast & Uppland & N & & 60\selectlanguage{ngerman}°12'18''N &
18\selectlanguage{ngerman}°33'55''E\tabularnewline
Rådmansholmen & Coast & Uppland & N & & 59\selectlanguage{ngerman}°37'18''N &
18\selectlanguage{ngerman}°56'52''E\tabularnewline
Senneby Hake & Coast & Uppland & Y & Y & 59\selectlanguage{ngerman}°58'23''N &
18\selectlanguage{ngerman}°54'21''E\tabularnewline
Skedvik & Coast & Uppland & N & & 58\selectlanguage{ngerman}°58'15''N & 17\selectlanguage{ngerman}°45'
1''E\tabularnewline
Storsten & Coast & Uppland & N & Y & 60\selectlanguage{ngerman}°31'16''N & 18\selectlanguage{ngerman}°
0'23''E\tabularnewline
Sveden & Coast & Uppland & N & & 59\selectlanguage{ngerman}°35'12''N &
18\selectlanguage{ngerman}°39'58''E\tabularnewline
Tranviken & Coast & Uppland & N & & 60\selectlanguage{ngerman}° 9'52''N &
18\selectlanguage{ngerman}°46'43''E\tabularnewline
Tullgarn & Coast & Uppland & N & & 58\selectlanguage{ngerman}°57'22''N &
17\selectlanguage{ngerman}°35'10''E\tabularnewline
Tvärnö & Coast & Uppland & N & Y & 60\selectlanguage{ngerman}°12' 9''N &
18\selectlanguage{ngerman}°32'42''E\tabularnewline
Bergianska & Inland & Uppland & N & & 59\selectlanguage{ngerman}°22' 3''N & 18\selectlanguage{ngerman}°
2'46''E\tabularnewline
Frötuna & Inland & Uppland & N & & 59\selectlanguage{ngerman}°54'45''N &
17\selectlanguage{ngerman}°51'52''E\tabularnewline
Jönsbolsjön & Inland & Uppland & N & & 59\selectlanguage{ngerman}°51'21''N & 18\selectlanguage{ngerman}°
1'52''E\tabularnewline
Kromsta & Inland & Uppland & N & & 59\selectlanguage{ngerman}°40'38''N &
17\selectlanguage{ngerman}°15'13''E\tabularnewline
Kärven & Inland & Uppland & N & & 59\selectlanguage{ngerman}°54'54''N & 18\selectlanguage{ngerman}°
9'10''E\tabularnewline
Ludden & Inland & Uppland & N & & 59\selectlanguage{ngerman}°46'18''N &
18\selectlanguage{ngerman}°40'23''E\tabularnewline
Ribbingebäck & Inland & Uppland & N & & 59\selectlanguage{ngerman}°50'17''N &
17\selectlanguage{ngerman}°10'54''E\tabularnewline
Segersta viltvatten & Inland & Uppland & N & & 59\selectlanguage{ngerman}°38'12''N &
17\selectlanguage{ngerman}°24'27''E\tabularnewline
Senneby & Inland & Uppland & N & & 59\selectlanguage{ngerman}°57'47''N &
18\selectlanguage{ngerman}°51'13''E\tabularnewline
Örsundsbro & Inland & Uppland & N & & 59\selectlanguage{ngerman}°44'47''N &
17\selectlanguage{ngerman}°20'47''E\tabularnewline
U27 & Inland & Uppland & N & & 60\selectlanguage{ngerman}°12'59''N & 18\selectlanguage{ngerman}°12'38''E\tabularnewline
U28 & Inland & Uppland & N & & 60\selectlanguage{ngerman}°12'39''N & 18\selectlanguage{ngerman}°15'41''E\tabularnewline
U46 & Inland & Uppland & N & & 59\selectlanguage{ngerman}°54'46''N & 17\selectlanguage{ngerman}°23'45''E\tabularnewline
U113 & Inland & Uppland & N & & 59\selectlanguage{ngerman}°41'11''N & 17\selectlanguage{ngerman}°54'19''E\tabularnewline
U132 & Inland & Uppland & N & & 59\selectlanguage{ngerman}°27'41''N & 17\selectlanguage{ngerman}°41'56''E\tabularnewline
Dösjön & Coast & Kalmar & N & Y & 56\selectlanguage{ngerman}°28'35''N & 16\selectlanguage{ngerman}°
8'39''E\tabularnewline
Enetri & Coast & Kalmar & N & Y & 56\selectlanguage{ngerman}°15' 7''N & 16\selectlanguage{ngerman}°29'
8''E\tabularnewline
Enudden & Coast & Kalmar & N & Y & 56\selectlanguage{ngerman}°30'59''N &
16\selectlanguage{ngerman}°11'13''E\tabularnewline
Fågelmara & Coast & Kalmar & N & Y & 56\selectlanguage{ngerman}°13'28''N & 16\selectlanguage{ngerman}°
0'53''E\tabularnewline
Grisbäck & Coast & Kalmar & N & Y & 56\selectlanguage{ngerman}°19'59''N & 16\selectlanguage{ngerman}°
4'10''E\tabularnewline
Sandvik & Coast & Kalmar & Y & Y & 56\selectlanguage{ngerman}°22'22''N &
16\selectlanguage{ngerman}°24'14''E\tabularnewline
Ventlinge & Coast & Kalmar & Y & Y & 56\selectlanguage{ngerman}°17' 2''N &
16\selectlanguage{ngerman}°23'49''E\tabularnewline
BA6 & Inland & Halland & N & & 56\selectlanguage{ngerman}°34'60''N & 13\selectlanguage{ngerman}° 4' 5''E\tabularnewline
BA700 & Inland & Halland & N & & 56\selectlanguage{ngerman}°34'52''N & 13\selectlanguage{ngerman}°
5'27''E\tabularnewline
D2 & Inland & Halland & N & & 56\selectlanguage{ngerman}°38'25''N & 12\selectlanguage{ngerman}°59'49''E\tabularnewline
D3 & Inland & Halland & N & & 56\selectlanguage{ngerman}°37'58''N & 13\selectlanguage{ngerman}° 0'15''E\tabularnewline
D7 & Inland & Halland & N & & 56\selectlanguage{ngerman}°36'34''N & 12\selectlanguage{ngerman}°55'28''E\tabularnewline
D12 & Inland & Halland & N & & 56\selectlanguage{ngerman}°26' 7''N & 12\selectlanguage{ngerman}°56'30''E\tabularnewline
D15 & Inland & Halland & N & & 56\selectlanguage{ngerman}°26'52''N & 13\selectlanguage{ngerman}° 0'37''E\tabularnewline
D16 & Inland & Halland & N & & 56\selectlanguage{ngerman}°27' 0''N & 13\selectlanguage{ngerman}° 3'41''E\tabularnewline
D17 & Inland & Halland & N & & 56\selectlanguage{ngerman}°40' 5''N & 12\selectlanguage{ngerman}°46'46''E\tabularnewline
D18 & Inland & Halland & N & & 56\selectlanguage{ngerman}°38'40''N & 12\selectlanguage{ngerman}°47'19''E\tabularnewline
D19 & Inland & Halland & N & & 56\selectlanguage{ngerman}°45'28''N & 12\selectlanguage{ngerman}°39'43''E\tabularnewline
D20 & Inland & Halland & N & & 56\selectlanguage{ngerman}°44'15''N & 12\selectlanguage{ngerman}°46' 4''E\tabularnewline
D22 & Inland & Halland & N & & 56\selectlanguage{ngerman}°45' 3''N & 12\selectlanguage{ngerman}°52'52''E\tabularnewline
D23 & Inland & Halland & N & & 56\selectlanguage{ngerman}°49'47''N & 12\selectlanguage{ngerman}°53'28''E\tabularnewline
D25 & Inland & Halland & N & & 56\selectlanguage{ngerman}°52'13''N & 12\selectlanguage{ngerman}°45'31''E\tabularnewline
D28 & Inland & Halland & N & & 56\selectlanguage{ngerman}°52'44''N & 12\selectlanguage{ngerman}°44'57''E\tabularnewline
D29 & Inland & Halland & N & & 56\selectlanguage{ngerman}°57'55''N & 12\selectlanguage{ngerman}°24'15''E\tabularnewline
EA16 & Inland & Halland & N & & 56\selectlanguage{ngerman}°27'44''N & 13\selectlanguage{ngerman}° 4'27''E\tabularnewline
EA18 & Inland & Halland & N & & 56\selectlanguage{ngerman}°27'40''N & 13\selectlanguage{ngerman}° 5'13''E\tabularnewline
EA60 & Inland & Halland & N & & 56\selectlanguage{ngerman}°27'44''N & 13\selectlanguage{ngerman}° 8'43''E\tabularnewline
KA7 & Inland & Halland & N & & 56\selectlanguage{ngerman}°49'23''N & 12\selectlanguage{ngerman}°39'51''E\tabularnewline
MA18 & Inland & Halland & N & & 56\selectlanguage{ngerman}°25'38''N & 13\selectlanguage{ngerman}° 5'24''E\tabularnewline
VA4 & Inland & Halland & N & & 56\selectlanguage{ngerman}°33'45''N & 13\selectlanguage{ngerman}° 6'26''E\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

Appendix 2. Total list of prey species with DNA in spider guts, and the
number of spiders where the prey DNA was detected. Species id is
provided as multiple species or genus level when the identity cannot be
resolved (i.e., multiple species with sequence similarity \textgreater{}
97\%).\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
Prey order & Family & Species & No of spiders\tabularnewline
\midrule
\endhead
Annelidae & Enchytraeidae & Enchytraeus albidus & 6\tabularnewline
Annelidae & Enchytraeidae & Enchytraeus buchholzi & 1\tabularnewline
Annelidae & Enchytraeidae & Enchytraeus moebii/albidus &
4\tabularnewline
Annelidae & Enchytraeidae & Lumbricillus pagenstecheri &
3\tabularnewline
Aranae & Clubionidae & Clubiona phragmitis & 3\tabularnewline
Aranae & Clubionidae & Clubiona reclusa/norwegica & 2\tabularnewline
Aranae & Linyphiidae & Erigone arctica & 17\tabularnewline
Aranae & Linyphiidae & Erigone longipalpis & 7\tabularnewline
Aranae & Theridiidae & Anelosimus vittatus & 1\tabularnewline
Aranae & Theridiidae & Theridion varians & 1\tabularnewline
Coleoptera & Anthicidae & Anthicus flavipes & 2\tabularnewline
Coleoptera & Carabidae & Bembidion pallidipenne & 2\tabularnewline
Coleoptera & Carabidae & Elaphrus uliginosus & 1\tabularnewline
Coleoptera & Carabidae & Pterostichus rhaeticus/nigrita &
1\tabularnewline
Coleoptera & Carabidae & Trechus secalis & 2\tabularnewline
Coleoptera & Chrysomelidae & Chrysolina staphylaea & 2\tabularnewline
Coleoptera & Curculionidae & Pelenomus quadrituberculatus &
6\tabularnewline
Coleoptera & Dermestidae & Dermestes szekessyi & 5\tabularnewline
Coleoptera & Hydraenidae & Ochthebius marinus/minimus & 1\tabularnewline
Coleoptera & Hydrophilidae & Cercyon depressus & 3\tabularnewline
Coleoptera & Scarabaeidae & Melolontha melolontha & 1\tabularnewline
Coleoptera & Scarabaeidae & Serica brunnea & 1\tabularnewline
Coleoptera & Silphidae & Oiceoptoma thoracicum & 2\tabularnewline
Coleoptera & Staphylinidae & Acrolocha sulcula & 1\tabularnewline
Coleoptera & Staphylinidae & Aleochara bipustulata/verna &
2\tabularnewline
Coleoptera & Staphylinidae & Atheta fungi & 1\tabularnewline
Coleoptera & Staphylinidae & Atheta vestita & 3\tabularnewline
Coleoptera & Staphylinidae & Cafius xantholoma & 1\tabularnewline
Coleoptera & Staphylinidae & Carpelimus rivularis & 18\tabularnewline
Coleoptera & Staphylinidae & Cordalia obscura & 1\tabularnewline
Coleoptera & Staphylinidae & Gnypeta carbonaria & 4\tabularnewline
Coleoptera & Staphylinidae & Omalium riparium & 9\tabularnewline
Coleoptera & Staphylinidae & Tachyporus nitidulus & 1\tabularnewline
Collembola & Entomobryidae & Desoria grisea & 8\tabularnewline
Collembola & Entomobryidae & Entomobrya lanuginosa & 2\tabularnewline
Collembola & Entomobryidae & Entomobrya multifasciata & 3\tabularnewline
Collembola & Entomobryidae & Lepidocyrtus lignorum & 1\tabularnewline
Collembola & Entomobryidae & Orchesella cincta/villosa &
5\tabularnewline
Collembola & Entomobryidae & Orchesella flavescens & 2\tabularnewline
Collembola & Hypogastruridae & Hypogastrura viatica & 2\tabularnewline
Collembola & Isotomidae & Halisotoma maritima & 9\tabularnewline
Collembola & Isotomidae & Isotoma anglicana & 3\tabularnewline
Collembola & Isotomidae & Isotoma riparia & 31\tabularnewline
Collembola & Isotomidae & Isotoma viridis(coerulea & 8\tabularnewline
Collembola & Isotomidae & Isotomurus fucicolus & 3\tabularnewline
Collembola & Katiannidae & Sminthurinus signatus & 1\tabularnewline
Collembola & Sminthuridae & Sminthurinus aureus & 2\tabularnewline
Collembola & Sminthuridae & unid & 2\tabularnewline
Collembola & Tomoceridae & Pogonognathellus spp & 2\tabularnewline
Diptera & Acroceridae & Acrocera orbicula & 16\tabularnewline
Diptera & Acroceridae & Ogcodes pallipes & 3\tabularnewline
Diptera & Agromyzidae & Agromyza albipennis & 1\tabularnewline
Diptera & Agromyzidae & Agromyza filipendulae & 1\tabularnewline
Diptera & Agromyzidae & Phytomyza horticola & 6\tabularnewline
Diptera & Anthomyiidae & Delia florilega & 1\tabularnewline
Diptera & Anthomyiidae & Delia platura & 10\tabularnewline
Diptera & Anthomyiidae & Fucellia fucorum & 31\tabularnewline
Diptera & Anthomyiidae & Fucellia tergina/maritima & 36\tabularnewline
Diptera & Asteiidae & Asteia amoena & 1\tabularnewline
Diptera & Carnidae & Meoneura sp & 1\tabularnewline
Diptera & Cecidomyiidae & Rhopalomyia sp. & 4\tabularnewline
Diptera & Cecidomyiidae & unid & 26\tabularnewline
Diptera & Ceratopogonidae & Atrichopogon fusculus & 4\tabularnewline
Diptera & Ceratopogonidae & Bezzia annulipes & 4\tabularnewline
Diptera & Ceratopogonidae & Culicoides newsteadi & 1\tabularnewline
Diptera & Ceratopogonidae & Dasyhelea turficola & 12\tabularnewline
Diptera & Ceratopogonidae & Forcipomyia hygrophila & 1\tabularnewline
Diptera & Ceratopogonidae & Palpomyia lineata & 2\tabularnewline
Diptera & Ceratopogonidae & unid & 2\tabularnewline
Diptera & Chamaemyiidae & Chamaemyia geniculata & 2\tabularnewline
Diptera & Chironomidae & Arctopelopia griseipennis & 1\tabularnewline
Diptera & Chironomidae & Chironomus aprilinus/pseudothummi &
6\tabularnewline
Diptera & Chironomidae & Cladopelma virescens & 1\tabularnewline
Diptera & Chironomidae & Cladotanytarsus difficilis & 1\tabularnewline
Diptera & Chironomidae & Cladotanytarsus gedanicus & 4\tabularnewline
Diptera & Chironomidae & Cladotanytarsus mancus & 20\tabularnewline
Diptera & Chironomidae & Cladotanytarsus nigrovittatus &
2\tabularnewline
Diptera & Chironomidae & Cladotanytarsus wexionensis/bicornutus &
1\tabularnewline
Diptera & Chironomidae & Cricotopus caducus/patens/flavocinctus &
11\tabularnewline
Diptera & Chironomidae & Cricotopus ornatus & 1\tabularnewline
Diptera & Chironomidae & Halocladius variabilis & 2\tabularnewline
Diptera & Chironomidae & Limnophyes sp. & 1\tabularnewline
Diptera & Chironomidae & Metriocnemus atriclava & 1\tabularnewline
Diptera & Chironomidae & Paratanytarsus inopertus/Chironomus plumosus &
16\tabularnewline
Diptera & Chironomidae & Paratanytarsus natvigi & 8\tabularnewline
Diptera & Chironomidae & Psectrocladius limbatellus & 4\tabularnewline
Diptera & Chironomidae & Psectrocladius oxyura & 10\tabularnewline
Diptera & Chironomidae & Pseudosmittia trilobata & 3\tabularnewline
Diptera & Chironomidae & Smittia leucopogon & 1\tabularnewline
Diptera & Chironomidae & Smittia sp. & 1\tabularnewline
Diptera & Chironomidae & Tanytarsus gracilentus & 1\tabularnewline
Diptera & Chironomidae & Tanytarsus usmaensis & 5\tabularnewline
Diptera & Chironomidae & unid & 1\tabularnewline
Diptera & Chloropidae & Chlorops pumilionis & 1\tabularnewline
Diptera & Chloropidae & Eutropha fulvifrons & 4\tabularnewline
Diptera & Chloropidae & Incertella sp./Rhopalopterum sp. &
3\tabularnewline
Diptera & Chloropidae & Meromyza nigriventrix/saltatrix &
1\tabularnewline
Diptera & Chloropidae & Oscinella sp. & 56\tabularnewline
Diptera & Chloropidae & Thaumatomyia notata & 2\tabularnewline
Diptera & Coelopidae & Coelopa frigida & 17\tabularnewline
Diptera & Dolichopodidae & Dolichopus nitidus & 1\tabularnewline
Diptera & Dolichopodidae & Dolichopus nubilus & 20\tabularnewline
Diptera & Dolichopodidae & Dolichopus pumilus mfl & 4\tabularnewline
Diptera & Dolichopodidae & Gymnopternus aerosus & 3\tabularnewline
Diptera & Dolichopodidae & Medetera truncorum/petrophiloides &
1\tabularnewline
Diptera & Dolichopodidae & Syntormon pallipes & 1\tabularnewline
Diptera & Dolichopodidae & Xanthochlorus ornatus & 1\tabularnewline
Diptera & Drosophilidae & Cacoxenus argyreator & 4\tabularnewline
Diptera & Drosophilidae & Scaptomyza flava/pallida & 36\tabularnewline
Diptera & Dryomyzidae & Heterocheila buccata & 8\tabularnewline
Diptera & Empididae & Rhamphomyia geniculata & 3\tabularnewline
Diptera & Empididae & unid & 2\tabularnewline
Diptera & Ephydridae & Discocerina obscurella & 5\tabularnewline
Diptera & Ephydridae & Hydrellia griseola & 1\tabularnewline
Diptera & Ephydridae & Lamproscatella sibilans & 3\tabularnewline
Diptera & Ephydridae & Limnellia quadrata & 1\tabularnewline
Diptera & Ephydridae & Paracoenia fumosa & 6\tabularnewline
Diptera & Ephydridae & Philotelma alaskense/defectum & 2\tabularnewline
Diptera & Ephydridae & Psilopa nigritella & 1\tabularnewline
Diptera & Ephydridae & Scatella paludum & 7\tabularnewline
Diptera & Ephydridae & Scatella stagnalis mfl & 23\tabularnewline
Diptera & Ephydridae & Scatella subguttata & 3\tabularnewline
Diptera & Ephydridae & Scatophila despecta & 2\tabularnewline
Diptera & Heleomyzidae & Trixoscelis obscurella & 3\tabularnewline
Diptera & Keroplatidae & Pyratula zonata & 1\tabularnewline
Diptera & Limoniidae & Symplecta stictica & 5\tabularnewline
Diptera & Limoniidae & unid & 1\tabularnewline
Diptera & Lonchopteridae & Lonchoptera bifurcata & 3\tabularnewline
Diptera & Muscidae & Coenosia lacteipennis & 1\tabularnewline
Diptera & Muscidae & Coenosia pedella/testacea & 6\tabularnewline
Diptera & Muscidae & Coenosia pumila & 1\tabularnewline
Diptera & Muscidae & Coenosia testacea & 1\tabularnewline
Diptera & Muscidae & Lispocephala erythrocera & 1\tabularnewline
Diptera & Muscidae & Morellia sinensis/tempestiva & 1\tabularnewline
Diptera & Muscidae & Schoenomyza litorella & 1\tabularnewline
Diptera & Muscidae & Spilogona aerea & 1\tabularnewline
Diptera & Opomyzidae & Geomyza sp. & 1\tabularnewline
Diptera & Opomyzidae & Opomyza germinationis/florum & 2\tabularnewline
Diptera & Phoridae & Megaselia albicans & 2\tabularnewline
Diptera & Phoridae & Megaselia brevicostalis & 14\tabularnewline
Diptera & Phoridae & Megaselia manicata & 4\tabularnewline
Diptera & Phoridae & Megaselia pleuralis & 5\tabularnewline
Diptera & Phoridae & Megaselia pusilla/ignobilis & 1\tabularnewline
Diptera & Phoridae & Megaselia sp. & 1\tabularnewline
Diptera & Phoridae & Metopina sp. & 3\tabularnewline
Diptera & Phoridae & unid & 5\tabularnewline
Diptera & Pipunculidae & Eudorylas fuscipes & 2\tabularnewline
Diptera & Psychodidae & Psychoda lativentris & 2\tabularnewline
Diptera & Rhinophoridae & Tricogena rubricosa & 2\tabularnewline
Diptera & Scathophagidae & Spaziphora hydromyzina & 1\tabularnewline
Diptera & Scathophagidae & Trichopalpus fraternus & 2\tabularnewline
Diptera & Scatopsidae & Coboldia fuscipes & 21\tabularnewline
Diptera & Scatopsidae & Scatopse notata & 10\tabularnewline
Diptera & Sciaridae & Corynoptera inundata & 1\tabularnewline
Diptera & Sciaridae & Lycoriella sativae & 13\tabularnewline
Diptera & Sciomyzidae & Ditaeniella grisescens & 1\tabularnewline
Diptera & Sepsidae & Themira putris & 43\tabularnewline
Diptera & Sphaeroceridae & Coproica hirtula & 2\tabularnewline
Diptera & Sphaeroceridae & Coproica lugubris & 9\tabularnewline
Diptera & Sphaeroceridae & Leptocera curvinervis & 73\tabularnewline
Diptera & Sphaeroceridae & Opacifrons coxata & 5\tabularnewline
Diptera & Sphaeroceridae & Opalimosina mirabilis & 4\tabularnewline
Diptera & Sphaeroceridae & Pullimosina heteroneura & 6\tabularnewline
Diptera & Sphaeroceridae & Pullimosina pullula & 2\tabularnewline
Diptera & Sphaeroceridae & Rachispoda intermedia/fuscipennis &
34\tabularnewline
Diptera & Sphaeroceridae & Rachispoda limosa & 10\tabularnewline
Diptera & Sphaeroceridae & Rachispoda lutosa/breviceps &
10\tabularnewline
Diptera & Sphaeroceridae & Thoracochaeta seticosta & 58\tabularnewline
Diptera & Sphaeroceridae & Thoracochaeta zosterae & 51\tabularnewline
Diptera & Syrphidae & Eristalis sp & 1\tabularnewline
Diptera & Syrphidae & Eupeodes corollae & 1\tabularnewline
Diptera & Syrphidae & Platycheirus sp. & 1\tabularnewline
Diptera & Tabanidae & Chrysops relictus/viduatus/rufipes &
2\tabularnewline
Diptera & Tabanidae & Tabanus cordiger/unifasciatus & 1\tabularnewline
Diptera & Tachinidae & Siphona sp. & 3\tabularnewline
Hemiptera & Anthocoridae & Orius majusculus/minutus & 2\tabularnewline
Hemiptera & Anthocoridae & Orius niger/horvathi & 3\tabularnewline
Hemiptera & Aphididae & Euceraphis betulae & 1\tabularnewline
Hemiptera & Cicadellidae & Arthaldeus striifrons & 3\tabularnewline
Hemiptera & Cicadellidae & Euscelis sordida & 3\tabularnewline
Hemiptera & Cicadellidae & Limotettix striola & 3\tabularnewline
Hemiptera & Cicadellidae & Paralimnus phragmitis & 2\tabularnewline
Hemiptera & Cicadellidae & Planaphrodes bifasciatus & 1\tabularnewline
Hemiptera & Cicadellidae & Psammotettix confinis/alienus &
14\tabularnewline
Hemiptera & Cicadellidae & Psammotettix nodosus/dubius &
1\tabularnewline
Hemiptera & Cicadellidae & Sarhoanus sp. & 2\tabularnewline
Hemiptera & Delphacidae & Delphax crassicornis & 2\tabularnewline
Hemiptera & Delphacidae & Euconomelus lepidus & 3\tabularnewline
Hemiptera & Delphacidae & Evacanthus interruptus & 1\tabularnewline
Hemiptera & Delphacidae & Javesella dubia/pellucida/forcipata &
12\tabularnewline
Hemiptera & Delphacidae & Unkanodes sp. & 13\tabularnewline
Hemiptera & Gerridae & Gerris thoracicus & 2\tabularnewline
Hemiptera & Miridae & Atractotomus mali/Phytocoris pini &
1\tabularnewline
Hemiptera & Miridae & Closterotomus norwegicus & 2\tabularnewline
Hemiptera & Miridae & Orthotylus sp. & 19\tabularnewline
Hemiptera & Nepidae & Nepa cinerea/rubra & 1\tabularnewline
Hemiptera & Piesmatidae & Parapiesma quadratum & 3\tabularnewline
Hemiptera & Saldidae & Saldula sp. & 5\tabularnewline
Hymenoptera & Aphelinidae & Aphelinus asychis & 2\tabularnewline
Hymenoptera & Aphelinidae & Aphelinus sp & 1\tabularnewline
Hymenoptera & Braconidae & Aphaereta minuta & 1\tabularnewline
Hymenoptera & Braconidae & Dolichogenida sp. & 2\tabularnewline
Hymenoptera & Braconidae & Praon flavinode & 1\tabularnewline
Hymenoptera & Braconidae & unid & 1\tabularnewline
Hymenoptera & Encyrtidae & Copidosoma floridanum & 1\tabularnewline
Hymenoptera & Eulophidae & Diglyphus isaea & 1\tabularnewline
Hymenoptera & Eulophidae & Pediobius sp. & 2\tabularnewline
Hymenoptera & Eulophidae & Tamarixia pronomus & 1\tabularnewline
Hymenoptera & Eulophidae & unid & 1\tabularnewline
Hymenoptera & Formicidae & Lasius niger & 14\tabularnewline
Hymenoptera & Formicidae & Lasius sp. & 3\tabularnewline
Hymenoptera & Formicidae & Myrmica rubra & 9\tabularnewline
Hymenoptera & Formicidae & Myrmica ruginodis & 12\tabularnewline
Hymenoptera & Ichneumonidae & Cotesia vestalis & 1\tabularnewline
Hymenoptera & Ichneumonidae & Diadegma armillatum & 1\tabularnewline
Hymenoptera & Ichneumonidae & Diadegma fenestrale/nanus &
2\tabularnewline
Hymenoptera & Ichneumonidae & Homotropus signatus & 1\tabularnewline
Hymenoptera & Mymaridae & Anaphes sp. & 2\tabularnewline
Hymenoptera & Pteromalidae & Halticoptera aenea & 1\tabularnewline
Hymenoptera & Pteromalidae & Psilonotus adamas & 2\tabularnewline
Hymenoptera & Pteromalidae & Pteromalus semotus & 1\tabularnewline
Hymenoptera & Pteromalidae & Trichomalopsis sp. & 3\tabularnewline
Hymenoptera & Pteromalidae & Trichomalus sp. & 6\tabularnewline
Hymenoptera & Pteromalidae & unid & 5\tabularnewline
Lepidoptera & Gelechiidae & Monochroa tetragonella & 1\tabularnewline
Lepidoptera & Gelechiidae & Scrobipalpa obsoletella & 6\tabularnewline
Lepidoptera & Glyphypterigidae & Glyphipterix thrasonella &
1\tabularnewline
Lepidoptera & Noctuidae & Autographa gamma & 1\tabularnewline
Lepidoptera & Plutellidae & Plutella xylostella & 14\tabularnewline
Lepidoptera & Psychidae & Narycia duplicella & 1\tabularnewline
Lepidoptera & Psychidae & Psyche casta & 1\tabularnewline
Mesostigmata & Ascidae & unid & 1\tabularnewline
Mesostigmata & Blattiscociidae & Cheiroseius sp. & 9\tabularnewline
Mesostigmata & Eviphipidae & unid & 14\tabularnewline
Mesostigmata & Macrocheliidae & unid & 11\tabularnewline
Mesostigmata & Parasitidae & Pergamasus crassipes & 2\tabularnewline
Mesostigmata & Parasitidae & unid & 39\tabularnewline
Opiliones & Phalangiidae & Mitopus morio & 1\tabularnewline
Orthoptera & Acrididae & Chorthippus spp. & 5\tabularnewline
Orthoptera & Acrididae & Stethophyma grossum & 2\tabularnewline
Orthoptera & Tetrigidae & Tetrix subulata & 1\tabularnewline
Sarcoptiformes & Ameronothridae & Ameronothrus & 1\tabularnewline
Sarcoptiformes & Ceratozetidae & Trichoribates novus & 1\tabularnewline
Sarcoptiformes & Crotoniidae & Platynothrus thori & 1\tabularnewline
Thysanoptera & Thripidae & unid & 6\tabularnewline
Trichoptera & Leptoceridae & Oecetis ochracea & 1\tabularnewline
Trombidiformes & Erythraeidae & Balaustium & 6\tabularnewline
Trombidiformes & Eupodidae & unid & 3\tabularnewline
Trombidiformes & Hydryphantidae & Hydryphantes crassipalpis &
1\tabularnewline
\bottomrule
\end{longtable}

\selectlanguage{english}
\FloatBarrier
\end{document}

