Lake responses to long-term disturbances and management practices

1. Long-term human-induced disturbances such as acidification and algal invasions, and management practices such as liming, are known to alter community structure and biodiversity of north temperate lakes. We assessed if they impacted on the trophic ecology and production of apex consumers (i.e. fish) and the overall food-chain length (FCL) of boreal lake ecosystems, and if these functional responses were consistent with the biodiversity changes. We hypothesise that fish production and FCL decrease with decreasing species biodiversity of lake communities, and that long-term environmental perturbations will alter the relative reliance of fish on littoral versus pelagic trophic pathways and their ontogenetic changes in trophic position (TP). 2. We analysed long-term data and stable isotopes of multiple organismal groups - phytoplankton, zooplankton, littoral and sub-littoral/profundal macroinvertebrates, and fish - collected from small boreal lakes that have been subjected to acidification, lime application and/or algal invasion by Gonyostomum semen. Species biodiversity, FCL and fish production (i.e. growth and catch-per-uniteffort) were compared among three lake categories, i.e. acidic, limed and circumneutral (reference) lakes, within each three lakes were selected. Fish TP and their relative littoral versus pelagic reliance were estimated based on stable nitrogen and carbon isotopes respectively. 3. Gonyostomum contributed to 77-98% phytoplankton biovolume in acidic lakes, <1-79% in limed lakes and 0-30% in circumneutral lakes. Its prevalence was correlated with total organic carbon concentration but not with lake pH, alkalinity or any other environmental variable. Diversity and evenness of phytoplankton, macroinvertebrates and fish generally decreased with increasing Gonyostomum biovolume, such that biodiversity was higher in circumneutral and limed lakes than in acidic lakes. 4. Isotopic data revealed that FCL was shortest in limed lakes (3.94 +/- 0.08; least- squares mean +/- SE), intermediate in acidic lakes (4.19 +/- 0.07) and longest in circumneutral lakes (4.38 +/- 0.08). Limed lakes also had the lowest fish growth and CPUE. Overall littoral reliance of fish was higher in acidic lakes (0.53 +/- 0.03) than in limed lakes (0.42 +/- 0.02) and circumneutral lakes (0.30 +/- 0.02), suggesting that fish production and FCL there could have been sustained by the increased littoral reliance when pelagic trophic pathways were hindered by Gonyostomum invasion. European perch (Perca fluviatilis), the most common fish in the lakes, showed faster TP increases in acidic and limed lakes, likely due to their earlier ontogenetic shift from zooplanktivory to zoobenthivory and/or piscivory. 5. Overall, our findings indicate that long-term disturbances (i.e. acidification and algal invasions) and management practices (i.e. liming) can (i) induce contrasting responses in biodiversity, FCL and fish production of boreal lakes; (ii) be the primary driver of FCL variation among small and similar-size ecosystems; and (iii) alter the trophic ecology (i.e. TP change during ontogeny and littoral reliance) of key fish species. The trophic ecology and production of apex consumers and FCL together can provide useful integrated proxies for ecosystem functioning, which can supplement traditional biodiversity measurements for more robust environmental assessments.