Fire impacts on the biology of stream ecosystems: A synthesis of current knowledge to guide future research and integrated fire management.

Freshwater ecosystems host disproportionately high biodiversity and provide unique ecosystem services, yet they are being degraded at an alarming rate. Fires, which are becoming increasingly frequent and intense due to global change, can affect these ecosystems in many ways, but this relationship is not fully understood. We conducted a systematic review to characterize the literature on the effects of fires on stream ecosystems and found that (1) abiotic indicators were more commonly investigated than biotic ones, (2) most previous research was conducted in North America and in the temperate evergreen forest biome, (3) following a control-impact (CI) or before-after (BA) design, (4) predominantly assessing wildfires as opposed to prescribed fires, (5) in small headwater streams, and (6) with a focus on structural and not functional biological indicators. After quantitatively analyzing previous research, we detected great variability in responses, with increases, decreases, and no changes being reported for most indicators (e.g., macroinvertebrate richness, fish density, algal biomass, and leaf decomposition). We shed light on these seemingly contradicting results by showing that the presence of extreme hydrological post-fire events, the time lag between fire and sampling, and whether the riparian forest burned or not influenced the outcome of previous research. Results suggest that although wildfires and the following hydrological events can have dramatic impacts in the short term, most biological endpoints recover within 5-10 years, and that detrimental effects are minimal in the case of prescribed fires. We also detected that no effects were more often reported by BACI studies than by CI or BA studies, raising the question of whether this research field may be biased by the inherent limitations of CI and BA designs. Finally, we make recommendations to help advance this field of research and guide future integrated fire management that includes the protection of freshwater ecosystems.

Forest management influences the effects of streamside wet areas on stream ecosystems.

Riparian zones contain areas of strong hydrological connectivity between land and stream, referred to as variable source areas (VSAs), and are considered biogeochemical control points. However, little is known about whether VSAs influence stream communities and whether this connectivity is affected by forest management. To address this, we used multiple biotic and abiotic indicators to (1) examine the influence of VSAs on riparian vegetation and stream ecosystems by comparing VSA and non-VSA reaches and (2) explore how forest management may affect the influence of VSAs on stream ecosystems. We detected some significant differences between VSA and non-VSA reaches in the riparian vegetation (greater understory and lower tree density) and stream ecosystem indicators (greater dissolved organic matter aromaticity, microbial biomass, peroxidase activity and collector-gatherer density, and lower dissolved organic carbon concentrations, algal biomass, and predatory macroinvertebrate density), which suggests that VSAs may create a more heterotrophic ecosystem locally. However, we show some evidence that forest management activities (specifically, road density) can alter the influence of VSAs and eliminate the differences observed at lower forest management intensities, and that the most hydrologically connected areas seem more sensitive to disturbance. Therefore, we suggest that the heterogeneity in hydrological connectivity along riparian zones should be considered when planning forest harvesting operations and road building (e.g., wider riparian buffers around VSAs).

Increased reliance of stream macroinvertebrates on terrestrial food sources linked to forest management intensity.

Our understanding of how forest management practices affect the relative importance of autochthonous vs. allochthonous resource use by headwater stream food webs is relatively poor. To address this, we used stable isotope (C, N, and H) analyses of food sources and macroinvertebrates from 15 streams in New Brunswick (Canada) and assessed how different catchment conditions arising from the gradient in forest management intensity affect the contribution of autochthonous resources to these food webs. Aquatic primary production contributed substantially to the biomass of invertebrates in these headwater streams, especially for scrapers and collector-gatherers (25-75%). However, the contribution of algae to food webs decreased as forest management intensity (road density and associated sediments, water cations/carbon, and dissolved organic matter humification) increased, and as canopy openness decreased. This trend was probably due to an increase in the delivery of organic and inorganic terrestrial materials (dissolved and in suspension) in areas of greater harvesting intensity and road density, which resulted in more heterotrophic biofilms. Overall, results suggest that, despite the presence of riparian buffers, forest management can affect stream food web structure via changes in energy flows, and that increased protection should be directed at minimizing ground disturbance in areas with direct hydrological connection to streams and at reducing dissolved and particulate matter inputs from roads and stream crossings in catchments with high degrees of management activity.

Effects of forest management on mercury bioaccumulation and biomagnification along the river continuum.

Forest management can alter the mobilization of mercury (Hg) into headwater streams and its conversion to methylmercury (MeHg), the form that bioaccumulates in aquatic biota and biomagnifies through food webs. As headwater streams are important sources of organic materials and nutrients to larger systems, this connectivity may also increase MeHg in downstream biota through direct or indirect effects of forestry on water quality or food web structure. In this study, we collected water, seston, food sources (biofilm, leaves, organic matter), five macroinvertebrate taxa and fish (slimy sculpin; Cottus cognata) at 6 sites representing different stream orders (1-5) within three river basins with different total disturbances from forestry (both harvesting and silviculture). Methylmercury levels were highest in water and some food sources from the basin with moderate disturbance (greater clearcutting but less silviculture). Water, leaves, stoneflies and fish increased in MeHg or total Hg along the river continuum in the least disturbed basin, and there were some dissipative effects of forest management on these spatial patterns. Trophic level (δ15N) was a significant predictor of MeHg (and total Hg in fish) within food webs across all 18 sites, and biomagnification slopes were significantly lower in the basin with moderate total disturbance but not different in the other two basins. The elevated MeHg in lower trophic levels but its reduced trophic transfer in the basin with moderate disturbance was likely due to greater inputs of sediments and of dissolved organic carbon that is more humic, as these factors are known to both increase transport of Hg to streams and its uptake in primary producers but to also decrease MeHg bioaccumulation in consumers. Overall, these results suggest that the type of disturbance from forestry affects MeHg bioaccumulation and trophic transfer in stream food webs and some longitudinal patterns along a river continuum.

Elevated Allochthony in Stream Food Webs as a Result of Longitudinal Cumulative Effects of Forest Management.

The river continuum concept (RCC) predicts a downstream shift in the reliance of aquatic consumers from terrestrial to aquatic carbon sources, but this concept has rarely been assessed with longitudinal studies. Similarly, there are no studies addressing how forestry related disturbances to the structure of headwater food webs manifest (accumulate/dissipate) downstream and/or whether forest management alters natural longitudinal trends predicted by the RCC. Using stable isotopes of carbon, nitrogen and hydrogen, we investigated how: 1) autochthony in macroinvertebrates and fish change from small streams to larger downstream sites within a basin with minimal forest management (New Brunswick, Canada); 2) longitudinal trends in autochthony and food web length compare among three basins with different forest management intensity [intensive (harvest and replanting), extensive (harvest only), minimal] to detect potential cumulative/dissipative effects; and 3) forest management intensity and other catchment variables are influencing food web dynamics. We showed that, as predicted, the reliance of some macroinvertebrate taxa (especially collector feeders) on algae increased from small streams to downstream waters in the minimally managed basin, but that autochthony in the smallest shaded stream was higher than expected based on the RCC (as high as 90% for some taxa). However, this longitudinal increase in autochthony was not observed within the extensively managed basin and was weaker within the intensively managed one, suggesting that forest management can alter food web dynamics along the river continuum. The dampening of downstream autochthony indicates that the increased allochthony observed in small streams in response to forest harvesting cumulates downstream through the river continuum. Supplementary Information: The online version contains supplementary material available at 10.1007/s10021-021-00717-6.

Forest management impacts on stream integrity at varying intensities and spatial scales: Do abiotic effects accumulate spatially?

Though effects of forest harvesting on small streams are well documented, little is known about the cumulative effects in downstream systems. The hierarchical nature and longitudinal connectivity of river networks make them fundamentally cumulative, but lateral and vertical connectivity and instream processes can dissipate the downstream transport of water and materials. To elucidate such effects, we investigated how a suite of abiotic indicators changed from small streams to larger downstream sites (n = 6) within three basins ranging in forest management intensity (intensive, extensive, minimal) in New Brunswick (Canada) in the summer and fall of 2017 and 2018. Inorganic sediments, the inorganic/organic ratios and water temperatures significantly increased longitudinally, whereas nutrients and the fluorescence index of dissolved organic carbon (DOC; indication of terrestrial source) decreased. However, some longitudinal trends differed across basins and indicated downstream cumulative (inorganic sediments, the inorganic/organic ratios and to a lesser extent DOC concentration and humification) as well as dissipative (temperatures, nutrients, organic sediments) effects of forest management. Overall, we found that the effects previously reported for small streams with managed forests also occur at downstream sites and suggest investigating whether different management practices can be used within the extensive basin to reduce these cumulative effects.

Forest management impacts on stream integrity at varying intensities and spatial scales: Do biological effects accumulate spatially?

The effects of forest harvesting on headwaters are quite well understood, yet our understanding of whether impacts accumulate or dissipate downstream is limited. To address this, we investigated whether several biotic indicators changed from smaller to larger downstream sites (n = 6) within three basins that had intensive, extensive or minimal forest management in New Brunswick (Canada). Biofilm biomass and grazer abundance significantly increased from upstream to downstream, whereas organic matter decomposition and the autotrophic index of biofilms decreased. However, some spatial trends differed among basins and indicated either cumulative (macroinvertebrate abundance, predator density, sculpin GSI) or dissipative (autotrophic index, cotton decomposition) effects downstream, potentially explained by sediment and nutrient dynamics related to harvesting. No such among-basin differences were observed for leaf decomposition, biofilm biomass, macroinvertebrate richness or sculpin condition. Additionally, results suggest that some of the same biological impacts of forestry observed in small headwaters also occurred in larger systems. Although the intensive and extensive basins had lower macroinvertebrate diversity, there were no other signs of biological impairment, suggesting that, overall, current best management practices protect biological integrity downstream despite abiotic effects.