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Time dynamics of the mollusk communitiesIn this section, the presence-absence of the species recorded in the three periods (T1, T2, T3) are analyzed in relation to time, habitat, and human impact. The list of the 28 species of freshwater mollusks (17 gastropods and 11 bivalves) in T1–T3, their codes, and origins are given in Table 1.The number of mollusk species has increased in time as the river has shifted from lotic to a mixture of flowing and stagnant sectors due to the building of reservoirs. T1 was characterized mainly by rheophilic elements and prosobranchs. Some species became extinct during the hydro-technical works (before or during T2) and are unlikely to recover, such as the rheophilic Theodoxus transversalis and Lithoglyphus naticoides. Other rheophilic species disappeared between T1 and T2 but managed to survive in tributaries and repopulated some sectors during the last years. The most remarkable recovery is that of the thick-shelled river mussel Unio crassus, a species protected by EU legislation. T2 was characterized by some extinctions but also colonization by lentiphilic pulmonates and tolerant, resistant species such as some clams. A few lotic species also survived in the river sectors between the dams. In T3, we encountered a rich and diverse community, including some newly established populations of AIS and the discontinuous presence of both lentic and lotic communities. Overall, the present-day fauna is richer than in former periods, consisting of 15 species of gastropods and 8 bivalves. The AIS included the gastropods Physa acuta and Ferrissia californica, which arrived in the area most likely during the XXth century, Viviparus acerosus, which is native to the Danube, but unknown until after 2000 in the upper-middle Olt River basin, the bivalves Dreissena polymorpha, also native in the Danube but an invader in the middle Olt since 2008–2010, Sinanodonta woodiana, first found in 2015, and Corbicula fluminea, which was first discovered in the Olt (and also in Transylvania) during our survey in February 2020. The mean number of native species per river’s sector increases almost linearly (2.8 species per sector in T1, 3.3 in T2, and 4.6 in T3), while the corresponding values for AIS increase non-linearly (no AIS in T1, 0.6 species per sector in T2 and 3.2 in T3).In the CCA of freshwater mollusk community changes through time (Period as predictor), the adjusted explained variation was 23.6% (test on all axes, pseudo-F = 5.9, p = 0.001). The polygons delimiting the positions of the sites during the three periods of time show no overlap, and they were distinct and separated in the ordination space (Fig. 1a). T2 (the period with maximum human impact) is distinctly placed and separated from the period without impact (T1) along both ordination axes. Meantime, T3 is closer to T1, having an intermediate position between the other two periods, showing a trend of recovery, such as the return of some species. In the CCA of T1–T3 species presence-absence predicted by the selected environmental descriptors (Period, Habitat, and Impact) (Fig. 1b), the adjusted explained variation was 28.36% (test on all axes, pseudo-F = 4.2, p = 0.001). FD(Rao) computed on all FT was plotted as isolines by GAM on the ordination space (model AIC = -17.19, model test F = 5.1, p = 0.003; tests of non-linearity in predictor effects: F = 3.9, p = 0.03). The functional diversity decreased from T1 to T2, then increased sharply to T3; it also decreased from rivers (R) to lakes (L) and along the human impact gradient (Impact).Figure 1Canonical correspondence analysis (CCA) of mollusk communities: (a) classification diagram of sampling sites based on period (as predictors): T1—XIXth century, T2—1995–2000, T3—2020 (adjusted explained variation 23.6%; first axis accounts for 17.6% the second for 6.0%, both axes are significant, p = 0.001); (b) CCA diagram of species occurrence constrained by environmental predictors (period, habitat: L—lakes, lentic sector in reservoirs, R—river, lotic sectors, and Impact—human impact) with functional diversity expressed as Rao quadratic entropy index (FD (Rao)) isolines plotted by generalized additive models (GAM) on the ordination space (adjusted explained variation 28.36%; first axis accounts for 16.3%, the second for 6.0%, both axes are significant, p = 0.001) .Full size imageIn the dc-CA with the selected predictors on T1–T3 presence-absence data, the first two axes separate the communities by period, each positioned in a distinct quadrant (Fig. 2). After a decrease in diversity from T1 to T2, in T3, there were more species and higher functional diversity. In time, there was a reduction in body size, a switch from species with separate sexes to hermaphrodites, a transition of oviposition towards ovo-viviparity (in snails), and external fecundation (in bivalves), and a switch of the feeding type. The dc-CA adjusted explained variation was 16.47%; tests based on sectors and species showed significant relationships (combined test for all axes, pseudo-F = 2.6, p = 0.006), the dimensionality test based on case scores was significant for the first axis (pseudo-F = 4.2, p = 0.001) and marginally significant for the second one (pseudo-F = 1.1, p = 0.053). In contrast, the dimensionality test based on species scores was significant only for the first axis (pseudo-F = 1.6, p = 0.004). The adjusted variation explained by environmental predictors (Hab, Impact, and Period) was 28.36%, and by the selected functional traits (Sexes, FeedT, SizeM, and Ovipos) was 14.64%.Figure 2Double-constrained correspondence analysis (dc-CA) with selected predictors on presence-absence data in T1–T3. The selected functional traits (in blue) are Sexes (circles: H—hermaphrodite, S—separate sexes), Feeding type (squares: SCR—scraper, SS—scraper and sediment, SF—scraper and filter, F—filter, SEDF—suspension and deposit feeder), Oviposition (diamonds: OV—ovo-viviparity, CAP—capsule/eggmass, BE—parental care, juveniles in brood pouches of demibranchs, No—no oviposition, external fecundation), and mean body size (SizeM); the selected environmental predictors (in red) are time (Period, with levels T1—XIXth century, T2—1995–2000, T3—2020), habitat (R—river, lotic sector; L—lake, a lentic sector in reservoirs) and human impact (Impact). Species are coded by the first three letters of the genus and species names. The adjusted explained variation was 16.47%, the first axis accounts for 12.7% and the second for 2.2%. Native species have black labels, while aliens (AIS) are written in green.Full size imageWe have split the binary data describing communities into two parts: natives and AIS, using the latter as predictors. We partitioned the variation in native species composition explained by the three predictor groups (Period, Environment, and AIS) (Fig. 3), subjecting the explanatory variables to an interactive forward selection procedure. We used RDA with centered response variables (CCA can not be used because the empty rows in some tables hinder the use of a proper hierarchical permutation scheme). The adjusted explained variation was 39.6% (the simple effects: time accounted for 22.33%, habitat and impact 24.73%, and the selected AIS 20.82%). All simple and unique effects were significant (p  More

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