4. Discussion
The temporal variations in the species richness of propagules deposited
by hydrochory show similarities with what has been observed in other
rivers and what we propose in our hypotheses. These temporary changes
are determined by the increase in the level and volume of water in the
rivers during the rainy season (or higher water flow) which in turn
increase the number of species that are dispersed towards the riparian
zones (Boedtelje et al., 2004; Fraaije et al., 2017; Moggridge et al.,
2009). In contrast, the abundance of propagules dispersed by hydrochory
towards the riparian zone did not increase during the rainy season.
Previous work has found that there is a greater number of propagules
during the season of higher water flow in rivers due to the greater
capacity of propagule dispersal through water currents (Boedtelje et
al., 2004; Esper-Reyes et al., 2018; Fraaije et al., 2017; Moggridge &
Gurnell, 2010; Naiman et al., 2010). However, there is no temporal trend
in the deposition of propagules by hydrochory and it has been suggested
that this may be due to the great production of propagules of one
species (e.g. Veronica anagallis-aquatica ) throughout the year
(Gurnell et al., 2008), a similar phenomenon could be occurring in the
study area.
In our case, we observed that the dominant tree species in the riparian
zone (Alnus acuminata ) (Flores-Galicia et al., 2021) had a peak
in propagule release that was detected in the traps for other types of
dispersal at the end of the rainy season (November 2017). Subsequently,
the increase was reflected in the hydrochory traps at the beginning of
the dry season (January 2018) (Annex 1). The effect of A.
acuminata propagule production on temporal variations in the number of
propagules deposited by hydrochory is confirmed if this species is
eliminated from abundance analyses. By doing this, the number of
propagules deposited by hydrochory in the rainy season is greater than
in the dry season (rainy: \(\overset{\overline{}}{x}\) = 21.8, S.D. =
44.3, dry: \(\overset{\overline{}}{\text{x\ }}\)= 3.1, S.D.= 6.3,W = 502, d.f .= 45, p <0.01). The
foregoing indicates that, in this system, the temporal patterns of
abundance of propagules dispersed by hydrochory are not only determined
by the temporal variations in the flow of water from the rivers, but
also that the local production of propagules.
The differences in the composition of the propagules dispersed by
hydrochory between the dry and rainy seasons were consistent with our
hypothesis. These temporary changes in composition are related to
differences throughout the year in the production of propagules of the
species that make up the riparian communities or of the adjacent forests
(Esper-Reyes et al., 2018), the accumulation of these in the riparian
zones and their subsequent remobilization during river floods (Boedtelje
et al., 2004; Fraaije et al., 2017; Gurnell et al., 2008). In our system
we observed that some species with peaks of propagule production during
the rainy season were only observed during the same season in propagules
deposited by hydrochory, such as A. hickelli , Pinus sp.,Solanum sp and some Asteraceae (Asteraceae 2 y Asteraceae 6).
For the spatial dimension, there are contrasts between what our
hypotheses propose and what we observed in this study, specifically for
the richness patterns, the number of propagules and the composition of
the communities of propagules dispersed by hydrochory along the river.
Contrary to what was expected, a greater number of species dispersed by
hydrochory was not observed with increasing distance from the origin
site of the river. The absence of a positive relationship between the
number of species and the distance to the origin of the river may be the
result of the fact that downstream there was no accumulation of species
from upstream, as has been observed in other rivers (Andersson et al.,
2000; Andersson & Nilsson, 2002). The nesting analysis supports the
above since the composition of the communities of the sites close to the
origin of the river (sites one and two) is not a subset of the
communities further away from the origin of the river. The low nesting
indicates that the species near the origin of the river were not
transported downstream (sites beyond 7 km) or if they did, the
propagules were not deposited in the downstream riparian zones and
continued their course in the column. of water beyond the study area.
Regarding the number of propagules deposited by hydrochory along the
rivers, our results contrast with what we propose in our hypotheses and
what has been observed in other studies (Andersson & Nilsson, 2002;
Esper-Reyes et al., 2018). As occurs for the temporal variations in the
number of propagules, the absence of an increase in the propagules
deposited along the river could be related to the local production ofA. acuminata . However, when A. acuminata is eliminated
from the analyses, an increase in the number of propagules is observed
along the rivers in the hydrochory traps in the dry season (estimated=
0.15, z = 2.65, p = <0.01) but not in the rainy
season, where instead a decrease in the deposited propagules is observed
(estimated= -0.14, z = -2.46, p = 0.01). This same
analysis (exclusion of A.acuminata ) in the traps for other types
of dispersal yields similar results (rainy: estimated= -0.20, z =
-3.04, p = <0.01; dry: estimated= 0.17, z =
2.42, p = 0.01). This could indicate that the production of
propagules in the sites near the origin of the river is greater in the
rainy season, which increases the deposition of propagules by hydrochory
in the area, while in the dry season there is a greater production of
propagules in the sites furthest from the origin of the river.
The results that we observed with respect to the spatial dimension
(absence of relationship between the distance to the origin of the river
with the estimated species richness or with the number of propagules,
absence of nesting of the upstream communities in those of downstream),
has important implications for theoretical frameworks such as RHC that
hypothesize regarding longitudinal changes in diversity of riparian
plant communities. The RHC postulate that the number of plant species
increases to a maximum point in the intermediate zone of the rivers from
which it decreases towards its mouth, these changes are associated with:
(i) a linear increase in the disturbance in the riparian zones at along
the river and (ii) a constant increase in the number of species and
propagules dispersed by hydrochory (Nilsson et al., 1994, 2010). As
there is no greater deposition of species and propagules deposited
downstream, the patterns of diversity of the riparian plant communities
that the RHC supposes could not be observed in rivers of mountainous
systems (see for example Flores-Galicia et al., 2021; Pielech, 2021).
Some methodological limitations of this work did not allow us to
conclude whether the contrasts between what we propose in our hypotheses
with respect to the spatial distribution patterns of the propagules
along the river (that is, a greater number of propagules and species
deposited by hydrochory along the river, as well as nesting of the
upstream communities in the downstream communities) and what we observed
(absence of changes in the number of propagules, in the richness and
decrease in the nesting of the communities). This is due to the fact
that in this system: (i) the propagules transported in the water column
travel a shorter distance than has been observed in other studies
(Andersson et al., 2000; Boedeltje et al., 2003) or (ii) the propagules
are effectively transported over long distances but due to the
characteristic erosive capacity of mountain rivers (Wohl, 2010) the
propagules deposited in the riparian zone are quickly remobilized to
downstream and were not detected in our traps. Future studies could
determine which of these hypotheses is correct through controlled
propagule release experiments and the measurement of the distances they
travel (Boland, 2017; da Cunha et al., 2017) or through sampling of
propagules in the water column (Boedeltje et al., 2003).