The article category
Original article (Ecological Research)
Title
Connectivity and short-term flood interactively affect fish community
composition in drainage ditches of paddy fields in an agricultural
landscape
The full names of the
authors
Hideyuki Iwamoto1,4*, Daisuke
Tahara2,4, Takehito Yoshida3,41 Department of General Systems Studies, University of
Tokyo, 3-8-1 Komaba, Meguro, Tokyo 135-8902, Japan2 Research Center of Marine Bioresources, Fukui
Prefectural University, 49-8-2, Katsumi, Obama, Fukui 917-0116, Japan
3 Department of Ecosystem Studies, University of
Tokyo, 1-1-1 Yayoi, Bunkyo, Tokyo 113-8657, Japan4 Research Institute for Humanity and Nature, 457-4
Motoyama, Kamigamo, Kita-Ku, Kyoto 603-8047, Japan
*Corresponding author
Email:
iwamoto.freshwater@gmail.com
ORCiD id: https://orcid.org/0000-0002-9695-4145
Keywords
community assembly; river ecosystem; heavy rain; Bray-Curtis
dissimilarity; temporal turnover;
Abstract
Community assembly can change over time following variations in
connectivity among habitats. Previous studies have demonstrated the
influence of seasonal pulse floods on community structures by
facilitating species dispersal, but the effects of short-term flash
floods have been poorly investigated. We investigated fish community
structures before and after a short-term flood in drainage ditches of
paddy fields in the Kita River basin, Japan. The study sites were
classified into three connectivity types, “Transient” (connected only
during the flood, with a temporally-disappearing vertical gap between
drainage ditches and downstream rivers), “Connected” (always
connected, without the vertical gap), and “Disconnected” (always not
connected, with the vertical gap). The abundance of Misgurnus
anguillicaudatus , the only species that dispersed from paddy fields to
drainage ditches, increased after the flood irrespective of the
connectivity types, although the total abundance of the other species
significantly increased after the flood only in the “Transient” sites.
The dissimilarity of species composition between the (n -1)-th andn -th surveys at each site was consistently low in the
“Connected” and “Disconnected” sites, whereas it rose just after the
flood and gradually declined in the “Transient” sites. Species
composition was significantly different among the connectivity types,
indicating that some fish species need stable connectivity even during
the non-flood period. The results showed that the short-term flood
facilitated fish dispersal from paddy fields by the increased water
discharge and that from downstream rivers by the temporary removal of
vertical gap, emphasizing the importance of hydrological connectivity
for biodiversity conservation in an agricultural landscape.
Introduction
Metacommunity theories have emphasized that local communities are
assembled not only through local processes related to environmental
filtering and species interaction but also through regional processes
based on species dispersal (Leibold et al., 2004). Connectivity is one
of the most important factors affecting community assembly, which is
defined as “the degree to which the landscape facilitates or impedes
movement among resource patches” (Taylor et al., 1993). A decline in
connectivity increases the effect of local niche-driven processes and
decreases the mass effect that leads to the homogenization of community
structure at adjacent localities due to high dispersal (Heino et al.,
2015; Leibold et al., 2004). Moreover, connectivity can temporally
fluctuate on various time scales, such as within a day and around a year
in dynamic ecosystems (Zeigler & Fagan, 2014). As theoretical studies
showed influences of time-varying connectivity on metapopulation and
metacommunity dynamics (Aiken & Navarrete, 2014; Perry & Lee, 2019),
it is one of the main issues to understand how this transient nature of
connectivity drives community assembly and variations in community
structures.
Previous studies in stream ecosystems have provided valuable insight
into the dynamic mechanisms of community assembly. Among them, several
studies demonstrated the effects of seasonal flow fluctuation on the
assembly of the aquatic community, mediated by temporal changes in local
environmental conditions and connectivity, in tropical rivers (Arrington
& Winemiller, 2006; Fernandes et al., 2014; Fitzgerald et al., 2017)
and intermittent rivers (Crabot et al., 2020; Sarremejane et al., 2017).
Seasonal pulse floods, which alternate wet and dry phases over several
months gradually, increase the surface areas and connectivity of habitat
patches, facilitating species dispersal. As a result, community assembly
is more driven by the mass effect at the beginning of the flood, and
then by the niche-driven mechanisms based on the species sorting at the
end of the flood because species can track preferred environmental
conditions (Datry et al., 2016; Fernandes et al., 2014; Sarremejane et
al., 2017). The temporal pattern of assembly processes can be affected
by the details of the flood regime, such as duration, frequency,
intensity, and timing (Datry et al., 2016; Lepori & Malmqvist, 2009),
though empirical studies have been limited to those on the seasonal
pulse floods.
Short-term flash floods are another type of flood regime that
potentially influences community assembly, and characterize temperate
stream ecosystems with the expectation of an increase in magnitude and
frequency with climate change (Ledger & Milner, 2015). Local
physicochemical conditions and hydrological connectivity between
habitats can dramatically change during short-term flash floods and then
return to their original states after the floods. Although empirical
studies are still scarce, Larsen et al. (2019) investigated benthic
Chironomidae and Oligocheta communities in a dynamic river-floodplain
ecosystem for over a year. They found that community structures were
highly influenced by stochastic processes of dispersal shortly after
flash floods and then affected by deterministic processes of species
sorting during the stable hydrological period. In contrast to these
taxonomic groups, different taxonomic groups such as fish can actively
migrate in response to flood not only between habitats within a river
floodplain but also between multiple rivers, such as between a
mainstream and tributaries (Koizumi et al., 2013; Mori et al., 2018). It
is thus expected that the short-term flood can change fish community
structures of tributaries at a short time scale, depending on the degree
of changes in connectivity between a mainstream and tributaries.
However, few studies have examined this expectation by field surveys
with a fine temporal resolution before and after a short-term flood,
partly because of the difficulties in observing sporadic and
unpredictable flood events (McLaughlin et al., 2013).
Drainage ditches of paddy fields are habitats where the short-term
floods potentially influence fish community assembly. The modern
drainage ditches with concrete revetments receive water from paddy
fields and then drain to downstream rivers. Many fish species inhabit
drainage ditches and immigrate from these adjacent water bodies. A few
species resistant to dryness (e.g., mud loach) inhabit paddy fields and
immigrate to drainage ditches, especially during a midsummer drainage
practice of rice farming (Fujimoto et al., 2008), which is usually
conducted in between mid-June and mid-July and makes paddy fields dry up
for several days (Natuhara, 2013). On the other hand, some fish species
move into drainage ditches from downstream rivers, when agricultural
irrigation raises the water level of ditches enough for the species to
spawn and grow up (Onikura et al., 2009; Saitoh et al., 1988). In this
agricultural landscape, although a short-term flash flood is less likely
to provide aftereffects on local environmental conditions of the ditches
with concrete revetment (Nakano, 2017), it is likely to promote these
fish immigrations by improving the hydrological connectivity. The rise
in water level by the short-term flood enables fish species to move to
drainage ditches by removing a vertical gap at a confluence that impedes
the connectivity during non-flood periods (Minagawa, 2021). The
short-term flood also increases water discharge from paddy fields and
thus facilitates the fish immigration to ditches, as the midsummer
drainage practice does. However, few studies have investigated fish
community structures in drainage ditches during pre-flood and post-flood
periods.
We investigated fish community structures in drainage ditches for a
total of three months, once before and six times after a short-term
flash flood, in the Kita River basin, Fukui Prefecture, Japan. To
demonstrate how the short-term flood and connectivity influence fish
community assembly, the following hypotheses were tested. (H1a) The
short-term flood facilitates the drainage of water from paddy fields,
resulting in the increase in the immigration of fish species from paddy
fields to drainage ditches. In contrast, (H1b) the immigration from
downstream rivers to drainage ditches depends on the degree to which the
flood improves the connectivity by removing a vertical gap at a
confluence. (H2) Temporal turnover of local community compositions
sharply rises just after the flood depending on the degree of the
improvement of connectivity, and then gradually declines as time passes
after the flood. Community compositions are highly determined by
stochastic processes at the beginning due to the immigration during the
flood (i.e., mass effect), and subsequently by deterministic
niche-driven processes (i.e., species sorting) as the connectivity
weakens. (H3) The connectivity between drainage ditches and downstream
rivers during both the flood and non-flood periods influence the fish
species distributions. The short-term flood relaxes the dispersal
limitation of some fish species, while other species need stable
connectivity due to their life history.
Materials and methods
Study sites
We set 16 study sites in drainage ditches with concrete revetments in
the Kita River basin, Fukui Prefecture, Japan (Figure 1). These sites
received water from adjacent paddy fields and, in some cases, upstream
rivers via underground pipelines, and then drained water back to the
Kita River (mainstream) directly or via connected tributaries. Each site
was a segment of a drainage ditch with a length of 15m and within 600m
away from the first confluence of downstream rivers (i.e., the Kita
River or tributaries). There were vertical gaps of over 20 cm between
the paddy outlets and the studied drainage ditches. These study sites
did not dry up throughout the year. The width of the study sites varied
from 0.55 to 1.65m, so their area ranged between 8.25 and 24.75
m2.
In the Kita River basin, heavy rain from the seasonal rain front
resulted in a short-term flood event from the 7th to the 8th of July,
2020, which made the water level of the Takatsuka observatory reach the
12th highest level in the last decade, 5.44m (Figure 2a). Also, water
levels of study sites during the flood rose 56.5±13.4cm compared with
pre-flood and returned to the original levels several days after the
flood (Figure 2b). The first and second surveys were conducted several
days before and after the flood, respectively, and then we conducted
surveys approximately once in two weeks until the seventh survey in late
September. We defined the connectivity to downstream rivers based on the
presence/absence of a vertical gap of more than 20cm between the studied
drainage ditches and the downstream rivers. Based on the connectivity to
downstream rivers during the flood and non-flood period, study sites
were classified into three connectivity types, namely “Transient”,
“Connected”, and “Disconnected”. Four “Transient” sites usually
had a vertical gap of more than 20cm between drainage ditches and
downstream rivers but the gap disappeared during the flood, whereas ten
“Connected” sites always had no vertical gap and two “Disconnected”
sites always had the vertical gap even during the flood.
Environmental measurement and fish
sampling
In each survey, we measured nine environmental variables at each site to
examine whether the flood changed local environmental conditions. At the
center of each surveyed segment, we measured water depth and sediment
depth with a measuring rod, as well as water temperature, dissolved
oxygen, pH, and electrical conductivity by Multi-parameter Water Quality
Checker (U-50; HORIBA, Kyoto). We also measured flow velocity as an
average value of the measurements three times by a flow meter with an
impeller (CR-7; Cosmo Riken, Tokyo). The coverage of terrestrial
vegetations over ditches and submerged plant at the bottom of ditches
were examined by visual observations.
We examined fish community compositions of study sites. At each end of a
study site, a trap net (mesh size, 2 mm) was installed to prevent fish
from migrating from and immigrating to the study site. An investigator
collected fish for 15 min using an electrofisher (LR-24; SMITH-ROOT;
USA) in one hand and a D-frame dip net (28 cm wide, 3 mm mesh) in
another hand. After that, the investigator used only the D-frame dip net
for 20 min to mainly collect benthic fish species. Collected fish were
identified and counted in the field, whereas some fish individuals were
too small to accurately identify at the level of genus and therefore
were excluded from the data. All collected fish were immediately
released back to their original ditches after counting.
Additionally, we recorded the presence of unionid species (freshwater
mussels, family Unionidae). This is because some bitterling fish species
(subfamily Acheilognathinae) spawn on the gills of these mussels and
therefore their presence in ditches is closely related to that of these
unionid species (Nagayama et al., 2012; Terui et al., 2011).
Statistical analysis
To statistically examine temporal changes of each environmental
variable, we performed the Friedman test (Friedman, 1937), a
nonparametric equivalent to the repeated measures ANOVA, in each
connectivity type. As for each environmental variable, we grouped
time-series values by surveys, calculated the ranks of sites within each
survey, and analyzed to determine whether statistical differences
existed in the ranks of sites among the seven surveys. We then used post
hoc many-to-one exact multiple comparisons to test the equality of each
post-flood survey (i.e., from the second to seventh one) against the
pre-flood, first survey (Eisinga et al., 2017). The reasons why we did
not conduct multiple comparisons of all pairs were because we were
particularly interested in the difference between the pre-flood and
post-flood surveys. Also, it should be noted that the statistical power
of detection would be low for the ”Disconnected” type as there were
only two sites.
M. anguillicaudatus is the only species that disperse from paddy
fields to drainage ditches in the Kita River basin according to our
previous study (Iwamoto et al., 2022), so we separately considered the
abundance of M. anguillicaudatus and that of all other species.
We also performed a series of Friedman tests and then the post hoc
multiple comparisons for temporal changes in these abundances for each
connectivity type, as mentioned above. Additionally, to illustrate the
temporal patterns, we calculated the relative difference in abundance
between the first and the focal (n -th) surveys at site i ,
as \(\left(x_{\text{in}}-x_{i1}\right)/\bar{x_{i}}\).
We calculated the temporal turnover of fish species composition based on
Bray–Curtis dissimilarity (Bray & Curtis, 1957) between them -th and n -th surveys at each site (referred to as\(D_{m,n}\) hereafter), which can take account of both the
presence/absence and the relative abundance of species. Particularly, we
focused on the two kinds of temporal turnover, \(D_{n-1,n}\) (between
the (n -1)-th and n -th) and \(D_{1,n}\) (between the first
and n -th). \(D_{n-1,n}\) describes whether the turnover in
species composition becomes greater or smaller over time, whereas\(D_{1,n}\) indicates whether the species composition during the
post-flood period becomes different from or similar to that of the
pre-flood as time passes. We conducted the Friedman test and then post
hoc multiple comparisons to analyze whether there were significant
differences between \(D_{1,2}\) and \(D_{1,n}\), and between \(D_{1,2}\)and \(D_{n-1,n}\).
To illustrate the fish species composition and their temporal changes,
we performed a nonmetric multidimensional scaling (NMDS) analysis
(Minchin, 1987) on a community matrix of the fish abundance of all 112
surveys (16 sites × 7 times) using the Bray–Curtis dissimilarity index.
Moreover, we used a permutational multivariate analysis of variance
(PERMANOVA) to clarify whether there are differences in fish community
composition between connectivity types, conducting permutations
constraining samples within each survey period (strata option within the
adonis2 function) to account for the effects of differences in time
(Anderson, 2017).
All analyses were performed in R 4.1.2 (R Core Team, 2021). Friedman
test and post hoc many-to-one exact multiple comparisons were
constructed and computed using the PMCMRplus (Pohlert, 2021).
Bray-Curtis dissimilarity, NMDS, and PERMANOVA were calculated and
conducted with the vegan (Oksanen et al., 2020) package in R.
Result
There were significant temporal changes in vegetation cover, water
temperature, dissolved oxygen, pH, and electrical conductivity in
“Transient” or “Connected” sites according to the Friedman test
(Table S1). These temporal changes seemed to correspond with seasonal
changes rather than the flood event, as post hoc multiple comparisons
showed that there was no significant difference between the first (i.e.,
pre-flood) and the second surveys, and between the first and the third
surveys (Table S2, Figure S1).
We recorded a total of 6895 fish individuals that belonged to 21 species
(15 Cypriniformes, 1 Siluriformes, 4 Perciformes, and 1 Beloniformes)
(Table S3). The mean abundance of fish species varied largely among
study sites, and Misgurnus anguillicaudatus was the most dominant
species, followed by Tanakia limbata , Cobitis sp. BIWAE
type A and Nipponocypris temminckii (Table S3, Figure S2).
The abundance of M. anguillicaudatus changed over time
significantly in the “Transient” and “Connected” sites, but did not
in the “Disconnected” sites, according to the Friedman test (Table 1).
Post hoc multiple comparisons demonstrated that the abundance in the
first survey was significantly different from that in the second and
third in the “Transient” sites and it was also different from that in
the third in the “Connected” sites (Table 1). The abundance generally
increased from the first to the second and third surveys, and then
decreased in the fourth irrespective of the connectivity types (Figure
3a), as it reached the highest in the second or third at 12 sites (four
“Transient”, six “Connected”, and two “Disconnected” sites). In
contrast, although there were also significant temporal changes in the
total abundance of species except for M. anguillicaudatus in the
“Transient” and “Connected” sites, the difference compared with the
first survey only existed in the third in the “Transient” sites (Table
1). The abundance in the “Connected” sites was generally high in the
fourth and seventh surveys, while that in the “Transient” sites
increased from the first to the third survey and then decreased in the
fourth (Figure 3b).
The temporal turnover of fish species composition between the
(n -1)-th and n -th surveys, \(D_{n-1,n}\), was
significantly changed as time passed only in the “Transient” sites
(Table 2). Post hoc multiple comparisons verified that \(D_{1,2}\) was
significantly different from \(D_{4,5}\), \(D_{5,6}\), \(D_{6,7}\) in
the “Transient” sites. The temporal turnover was highest in the second
survey (i.e., the dissimilarity between the first and second) and then
decreased as time passed, while those in the “Connected” and
“Disconnected” sites stayed at relatively low levels (Figure 4a).
There was no significant change over time in the temporal turnover
between the first and n -th survey, \(D_{1,n}\), regardless of the
connectivity types (Table 2). However, the mean of the temporal turnover
was largest in the “Transient” sites (0.807), followed in order by the
“Connected” (0.497) and the “Disconnected” sites (0.305) (Figure
4b).
Fish species composition was significantly different among the
connectivity types (Figure 5), as confirmed by the PERMANOVA results
(R2 = 0.124, p = 0.001).Pseudaspius hakonensis and Gnathopogon elongatus elongatusbelonging to Cyprinidae were representative species after the flood in
the “Transient” sites, as newly observed in the second and third
surveys in four and two “Transient” sites, respectively (Table S2,
Figure 5, Figure S3a-d). T. limbata was one of the dominant
species in the “Connected” sites (Table S3, Figure 5, Figure S3), as
there were more than two individuals found only in seven “Connected”
and one “Transient” sites during the entire period.
We recorded the presence of two unionid mussel species,Pronodularia japanensis only in five “Connected” sites andBuldowskia iwakawai in one “Transient”, two “Connected”, and
one “Disconnected” sites (Table S3). Of the eight sites where T.
limbata was dominant in the fish community, either P. japanensisor B. iwakawai occurred in five “Connected” and one
“Transient” sites, and the other two sites where neither species was
present shared the same confluence of the mainstream. Only in one
“Disconnected” site, there was the unionid mussel species, butT. limbata was not observed.
Discussion
No significant difference in local environmental variables was found
between before and just after the flood (Table S1, Table S2, Figure S1),
although the water level in drainage ditches actually rose during the
flood (Figure 2b). Short-term floods often cause changes in
environmental conditions within intact rivers, such as the modification
of channel shape and depth, but the modern drainage ditches with
concrete revetments have more constant width, shape and water depth
(Nakano, 2017), indicating that changes in environmental conditions that
continue well after the flood would be hard to occur. Therefore, the
observed difference in fish community structure between before and after
the flood was most likely to be derived from temporary changes in
connectivity, rather than from environmental changes associated with the
flood.
The abundance of M. anguillicaudatus increased significantly
after the flood in the “Transient” and “Connected” sites, supporting
the hypothesis (H1a), whereas the similar increase in the
“Disconnected” sites was not significant probably due to their small
number of sites (Figure 3a). The flood facilitated the discharge of
water and the emigration of M. anguillicaudatus from paddy fields
to drainage ditches, as the midsummer drainage practice of rice farming
does usually in mid-June to mid-July in the study area (Fujimoto et al.,
2008). The seasonal timing of the flood seems to be important because
the abundance of M. anguillicaudatus in paddy fields commonly
shows a peak at the end of the spawning season, which is before the
midsummer drainage practice (Fujimoto et al., 2008; Tanaka, 1999). If
the flood had occurred well after the midsummer drainage practice, the
extent of the emigration of M. anguillicaudatus would have been
more modest. It was also possible that M. anguillicaudatus moved
from downstream rivers, but this was less likely to contribute to
increasing the abundance. This is because the emigration of M.
anguillicaudatus from paddy fields to drainage ditches widely observed
around the midsummer drainage practice (Iwamoto et al., 2022), whereas
the fish was observed at few points in downstream rivers during the
summer in the studied Kita River basin (Matsumiya et al., 2001).
Effects of the flood on the immigration from downstream rivers to
drainage ditches depended on the connectivity type with downstream
rivers, supporting the hypothesis (H1b). The total abundance of fish
species except for M. anguillicaudatus significantly increased
after the flood only in the “Transient” sites, but it did not largely
change between before and after the flood in the “Connected” and
“Disconnected” sites (Figure 3b). The flood temporarily removed a
vertical gap between drainage ditches and downstream rivers in the
“Transient” sites, probably resulting in the immigration of fish
species to drainage ditches. Many fish species used the temporary
underwater path to actively immigrate to drainage ditches that serve as
spawning and nursery habitats (Nakano, 2017). Additionally, fish species
could evacuate from the disturbed downstream rivers during flood, as
previous studies showed that fish species inhabiting a mainstream
actively move into tributaries or backwaters for refuge when a flood
increased flow discharge and turbidity in the mainstream (Koizumi et
al., 2013; Mori et al., 2018; Sumida et al., 2019). Also, some fish
species were likely to be passively swept away to drainage ditches by
backflow of water from downstream rivers, which was expected to be
caused by rapid rise of the water level in downstream rivers, although
this backflow was not observed during this study. In contrast, there was
little increase in the abundance of fish species other than M.
anguillicaudatus in the “Connected” sites, although fish could have
moved to drainage ditches at the “Connected” sites as well as at the
“Transient” sites during the flood. This might be because fish species
had already moved freely between downstream rivers and drainage ditches
even before the flood in the “Connected” sites. We would also have
failed to observe fish species that evacuated from a mainstream because
our sampling of fish was done not during but several days after the
flood for safety and practical reasons. Indeed, previous studies showed
that some fish species that once moved to tributaries during a flood
returned to a mainstream as soon as the flood settled (Koizumi et al.,
2013; Mori et al., 2018).
The temporal turnover of fish species composition between the
(n -1)-th and n -th surveys (i.e., \(D_{n-1,n}\))
fluctuated over time depending the connectivity type, coinciding with
the hypothesis (H2). The temporal turnover was consistently low in the
“Connected” and “Disconnected” sites, whereas it rose just after the
flood and gradually declined in the “Transient” sites (Table 2, Figure
4a). Local fish species composition in the “Transient” sites largely
changed between before and right after the flood, resulting from the
immigration of some fish species from downstream rivers (i.e., mass
effect) that was limited before the flood. Although the “Transient”
sites again lost the hydrological connectivity after the flood due to
the vertical gap, the community compositions continued to fluctuate to
some extent for around a month (Figure 4a). Moreover, the abundance of
fish species in “Transient” sites reached their peak around three
weeks after the flood rather than several days after the flood, and then
declined and became stable a month after the flood (Figure 3). These
results may indicate that fish species that once immigrated from
downstream rivers migrated within networks of drainage ditches, and then
returned to downstream rivers or died as time passed in the drainage
ditches. After that, fish species composition was likely to approach the
deterministic one that was influenced by local environmental conditions.
The series of temporal changes occurred within around a month, which is
relatively short compared to seasonal floods influencing fish
communities (Fernandes et al., 2014) or short-term floods determining
invertebrate communities (Larsen et al., 2019). The rapid responses of
fish communities derived from their high dispersal ability and
longitudinally asymmetric connectivity that allowed immigration from
drainage ditches to downstream rivers but prevented that of the opposite
direction after the flood.
The temporal turnover between the first and n -th surveys (i.e.,\(D_{1,n}\)) were generally higher in the “Transient” than in the
“Connected” and “Disconnected” sites, and it showed little change as
time passed irrespective of the connectivity type (Figure 4b). The
reason why the species composition during the post-flood period did not
become similar to that of the pre-flood should be partly because some
fish individuals that immigrated from downstream rivers and paddy fields
survived in drainage ditches. However, even after the final survey in
late September, the species composition would further change due to the
influence of water level decline during the agricultural off-season, so
it would become similar to the pre-flood species composition in the
following spring (Iwamoto et al., 2022).
The connectivity between drainage ditches and downstream rivers during
both the flood and non-flood periods affected the distribution of some
fish species, as hypothesized in (H3). P. hakonensis and G.
elongatus elongatus newly appeared and their abundance rose in the
“Transient” sites after the flood (Figure 5, Figure S3a-d), suggesting
that the flood facilitated their immigration through improvement of the
connectivity. These species seasonally immigrate from downstream rivers
to drainage ditches in order to spawn and grow up during the irrigation
flood (Moriyama et al., 2008; Saitoh et al., 1988; Takemura et al.,
2011). Also, these species move during flood to the backwaters in low
water channels, which is a semi-lentic habitat and functioned as a flow
refugia (Denda et al., 2002). In contrast, T. limbata was one of
the representative species in the “Connected” sites (Figure 5, Table
S2), as well as P. japanensis that T. limbata often use as
spawning beds (Inadome & Yamamoto, 2012; Kitamura, 2007). The continued
connection with downstream rivers was likely to affect the distribution
of T. limbata because the connectivity is also related to the
suitable habitat of unionid mussels (Nagayama et al., 2012; Terui et
al., 2011). The unionid mussels attach to the gills or fins of a host
fish during their first larval stage (glochidia) (Negishi et al., 2008),
so the hydrological connectivity affects their distribution through the
influence on migration of host fish species. Indeed, previous studies
showed that weirs in rivers limited the distribution of unionid mussels
by preventing host fish species from moving upstream (Kuwahara et al.,
2017; Watters, 1996). Negishi et al. (2014) also discussed the
possibility that vertical gaps reduced the connectivity between drainage
ditches and downstream rivers, and thus might have fragmented the mussel
habitat by limiting movement of host fish with attached glochidia.
The present results also emphasize the importance of hydrological
connectivity for biodiversity conservation in an agricultural landscape.
Aquatic communities in rivers have been threatened because of the
shrinking various waterbodies in floodplain by the modern river
improvement (Nagayama et al., 2015) and because of increasing intensity
and frequency of flood disturbance by ongoing climate changes (Ledger &
Milner, 2015). Therefore, drainage ditches in an agricultural landscape
can play more important roles as temporary refuges and seasonal habitats
for spawning and growth as the drainage ditches somewhat resemble
streamlets once existed in natural floodplains (Minagawa, 2021; Nakano,
2017). To secure their roles, it is desirable to recover the
hydrological connectivity between drainage ditches and downstream rivers
by removing vertical gaps (e.g. flap gates) that the modern river
improvement and the drainage system construction for agriculture have
created (Minagawa, 2021; Nakano, 2017). Moreover, the structures
permitting backflow retention of flood from downstream rivers to paddy
fields, such as the floodplain open levee system (kasumi-tei )
(Taki et al., 2021; Teramura & Shimatani, 2021; Yamada et al., 2022),
should allow drainage ditches and paddy fields to play a greater role as
temporary refuges for fish in the event of severer flood than that
observed in this study. In conclusion, our study demonstrated that the
short-term flood and connectivity can influence interactively the
process of fish community assembly in drainage ditches, and this
understanding provides important insights into biodiversity conservation
in an agricultural landscape.
Acknowledgments
The study was supported by the Research Institute for Humanity and
Nature (RIHN: a constituent member of NIHU) Project No. RIHN14200103 and
JSPS KAKENHI Grant Number 20H04377.
Conflict of interest
The authors declare that there is no conflict of interest.
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Tables and figures
Table 1. Results of the Friedman test and post hoc many-to-one exact
multiple comparisons on temporal changes in the abundance ofMisgurnus anguillicaudatus and that of all other species for each
connectivity type. Post hoc multiple comparisons analyzed equality of
each post-flood survey (i.e., from the second to the seventh one)
against the pre-flood, first survey. The significant level of
explanation and difference were shown in bold.