Amanda R. McCormick

Ecosystem engineers impact ecosystems by changing the physical environment in which species interact. In Lake Mývatn, Iceland, larvae of the midge Tanytarsus gracilentus function as ecosystem engineers by building tubes in the sediment that solidify the benthic surface and create a 3-dimensional structure on which microalgae can grow. To assess the ecosystem-scale effect of midge engineering, we took advantage of the natural high-amplitude fluctuations in T. gracilentus over a 13-year time series. We combined larval abundance data with benthic primary production under saturating light conditions (P _max ) and pelagic in situ gross primary production (GPP) measured on average 8.4 times per summer. T. gracilentus abundance had a positive effect on benthic P _max , as expected from their ecosystem engineering, while pelagic GPP had a negative effect on benthic P _max , reflecting competition from phytoplankton. A complementary analysis of five replicate sediment cores taken on each of 26 sampling days of the time series shows a quantitatively similar positive effect of local-scale variation in larval midge abundance on benthic net ecosystem production (NEP) in space. Finally, we reanalyzed data from an experiment manipulating light levels and the presence/absence of midge larvae to show that larval engineering increased benthic NEP more at low than at high light levels. This indicates that at least part of the positive engineering effect of midges on benthic P _max arises from amelioration of light limitation through self-shading, likely due to the structure created by midge tubes. These results show that Mývatn's high benthic primary productivity is due not only to high inputs of nutrients but also to the ecosystem engineering of tube-forming midges.

Measuring microalgae density in soft‐sediment benthos has challenges for even the most sophisticated methods. If the goal is to assess the photosynthetic potential of epipelon, then microalgae should be sampled only at the surface of the benthos to the depth of light penetration. Furthermore, microalgae density may show spatial and temporal variability that can only be captured by using many point samples and nondestructive sampling. Here, we use simple near‐infrared (NIR) imagery to assess surface density of microalgae in soft underwater sediments and to infer their photosynthetic capacity. In lab studies, NIR imagery gives estimates of epipelon density that are strongly correlated with standard chlorophyll a (Chl a) assays using pigment extraction and fluorometry ( = 0.70), but NIR imagery is better able to separate experimental treatments. In analyses of sediment samples from a lake, NIR imagery gives estimates of epipelon Chl a density that are strongly correlated to net ecosystem production (NEP). Near‐infrared imagery also gives a fine‐grained assessment of the spatial distribution of epipelon that helps to explain the relationship between epipelon density and NEP. Finally, images from an underwater NIR camera over the course of a wind disturbance event give estimates of the relative density of microalgae that is buried and is likely to be, at least temporarily, photosynthetically inactive. These results show that NIR imagery provides an easy and nondestructive method for sampling surface densities of microalgae which is particularly suitable for remote field locations and for educational settings in which students can generate results with cheap and robust equipment.

Competing species may show positive correlations in abundance through time and space if they rely on a shared resource. Such positive correlations might obscure resource partitioning that facilitates competitor coexistence. Here, we examine the potential for resource partitioning between two ecologically similar midge species (Diptera: Chironomidae) in Lake Mývatn, Iceland. Tanytarsus gracilentus and Chironomus islandicus show large, roughly synchronized population fluctuations, implying potential reliance on a shared fluctuating resource and thereby posing the question of how these species coexist at high larval abundances. We first considered spatial partitioning of larvae. Abundances of both species were positively correlated in space; thus, spatial partitioning across different sites in the lake did not appear to be strong. We then inferred differences in dietary resources with stable carbon isotopes. T. gracilentus larvae had significantly higher δ¹³C values than C. islandicus, suggesting interspecific differences in resource use. Differences in resource selectivity, tube-building behavior, and feeding styles may facilitate resource partitioning between these species. Relative to surface sediments, T. gracilentus had higher δ¹³C values, suggesting that they selectively graze on ¹³C-enriched resources such as productive algae from the surface of their tubes. In contrast, C. islandicus had lower δ¹³C values than surface sediments, suggesting reliance on ¹³C-depleted resources that may include detrital organic matter and associated microbes that larvae selectively consume from the sediment surface or within their burrow walls. Overall, our study illustrates that coexisting and ecologically similar species may show positive correlations in space and time while using different resources at fine spatial scales.

Population cycles can be caused by consumer–resource interactions. Confirming the role of consumer–resource interactions, however, can be challenging due to an absence of data for the resource candidate. For example, interactions between midge larvae and benthic algae likely govern the high‐amplitude population fluctuations of Tanytarsus gracilentus in Lake Mývatn, Iceland, but there are no records of benthic resources concurrent with adult midge population counts. Here, we investigate consumer population dynamics using the carbon stable isotope signatures of archived T. gracilentus specimens collected from 1977 to 2015, under the assumption that midge δ¹³C values reflect those of resources they consumed as larvae. We used the time series for population abundance and δ¹³C to estimate interactions between midges and resources while accounting for measurement error and possible preservation effects on isotope values. Results were consistent with consumer–resource interactions: high δ¹³C values preceded peaks in the midge population, and δ¹³C values tended to decline after midges reached high abundance. One interpretation of this dynamic coupling is that midge isotope signatures reflect temporal variation in benthic algal δ¹³C values, which we expected to mirror primary production. Following from this explanation, high benthic production (enriched δ¹³C values) would contribute to increased midge abundance, and high midge abundance would result in declining benthic production (depleted δ¹³C values). An additional and related explanation is that midges deplete benthic algal abundance once they reach peak densities, causing midges to increase their relative reliance on other resources including detritus and associated microorganisms. Such a shift in resource use would be consistent with the subsequent decline in midge δ¹³C values. Our study adds evidence that midge–resource interactions drive T. gracilentus fluctuations and demonstrates a novel application of stable isotope time‐series data to understand consumer population dynamics.

The abundances of competing species may show positive correlations in time and space if they rely on a shared resource. Such positive correlations might obscure resource partitioning that facilitates competitor coexistence. Here, we examine the potential for resource partitioning between two ecologically similar midge species (Diptera: Chironomidae) in Lake Mývatn, Iceland. Tanytarsus gracilentus and Chironomus islandicus show large, roughly synchronized population fluctuations, implying potential reliance on a shared fluctuating resource and thereby posing the question of how these species coexist at high larval abundances. We first considered spatial partitioning of larvae. Abundances of both species were positively correlated; thus, spatial partitioning across different sites in the lake did not appear to be strong. We then inferred differences in dietary resources with stable carbon isotopes. T. gracilentus larvae had significantly higher δ¹³C values than C. islandicus, suggesting interspecific differences in resource use. Differences in resource selectivity, tube-building behavior, and feeding styles may facilitate resource partitioning between these two species. Relative to surface sediments, T. gracilentus had higher δ¹³C values, suggesting that they selectively graze on ¹³C-enriched resources such as productive algae from the surface of their tubes. In contrast, C. islandicus had lower δ¹³C values than surface sediments, suggesting reliance on ¹³C-depleted resources, which may include detrital organic matter and associated microbes that larvae selectively consume from the sediment surface or within their burrow walls. Overall, our study illustrates that coexisting and ecologically similar species may show positive correlations in space and time while using different resources at fine spatial scales.

The abundances of competing species may show positive correlations in time and space if they rely on a shared resource. Such positive correlations might obscure resource partitioning that facilitates coexistence of competitors and affects their abundances, spatial distributions, and population dynamics. Here, we examine the potential for resource partitioning between two ecologically similar midge species (Diptera: Chironomidae) in Lake Mývatn, Iceland. Tanytarsus gracilentus and Chironomus islandicus larvae coexist at high abundances in benthic habitats, and they have been previously described as feeding upon diatoms and detritus. Furthermore, both species show large, roughly synchronized population fluctuations, implying potential reliance on a shared fluctuating resource and posing the question of how these species coexist at high abundances. We first considered spatial partitioning of larvae; across multiple sites, abundances of both species were positively correlated. Thus, spatial partitioning across different sites in the lake did not appear to be strong. We then inferred differences in dietary resource use with stable carbon isotopes. T. gracilentus larvae had higher δ ¹³ C values than C. islandicus (mean difference = 5.39 ± 1.84‰), suggesting interspecific differences in resource use. Differences in resource selectivity, tube-building behavior, and feeding styles may facilitate resource partitioning between these two species. Relative to surface sediments, T. gracilentus had higher δ ¹³ C values (1.84 ± 0.96‰), suggesting that they selectively graze on ¹³ C-enriched resources such as productive algae from the surface of their tubes. In contrast, C. islandicus had lower δ ¹³ C values than surface sediments (−2.87 ± 1.95‰), suggesting reliance on isotopically depleted resources, which may include detrital organic matter and associated microbes that larvae selectively consume from the sediment surface or within their tube walls. Overall, our study illustrates that coexisting and ecologically similar species may show positive correlations in space and time while using different resources at fine spatial scales.

Characterizing the dynamics of energy flow through ecosystems requires quantifying the degree to which primary and secondary production are coupled. This coupling is expected to be tight in ecosystems with high internal production relative to external carbon and energy inputs. We experimentally quantified the dependence of aquatic insect emergence on fresh primary production, specifically for the midge population in Lake Mývatn, Iceland. Using field mesocosms, we manipulated algal primary production by reducing light availability via shading. We then used dissolved oxygen incubations to estimate fluxes of carbon through photosynthesis (i.e., gross primary production or “GPP”) over the course of the experiment. We found that elevated GPP was associated with higher emergence rates of adults, as judged both by comparison of emergence across the experimental shading treatments and estimates of in situ GPP within the mesocosms. Furthermore, larger adults emerged earlier than smaller ones, suggesting that asymmetries in resource availability among individuals affected the timing of emergence. Nonetheless, midge emergence was substantial under light-limiting conditions, indicating that while midges benefit from primary production contemporaneous with larval development, they are also capable of completing their life cycles on carbon already existing in the organic matter pool. Our results show that even in systems with limited allochthonous inputs, contemporaneous primary production may be not necessary for high secondary production and insect emergence. Instead, consumers can develop from consumption of biomass derived from past autochthonous primary production. This suggests that primary production and consumer dynamics can be partially decoupled in time in systems that depend on internal production.

Ecosystem engineers have large impacts on the communities in which they live, and these impacts may feed back to populations of engineers themselves. In this study, we assessed the effect of ecosystem engineering on density-dependent feedbacks for midges in Lake Mývatn, Iceland. The midge larvae reside in the sediment and build silk tubes that provide a substrate for algal growth, thereby elevating benthic primary production. Benthic algae are in turn the primary food source for the midge larvae, setting the stage for the effects of engineering to feed back to the midges themselves. Using a field mesocosm experiment manipulating larval midge densities, we found a generally positive but nonlinear relationship between density and benthic production. Furthermore, adult emergence increased with the primary production per midge larva. By combining these two relationships in a simple model, we found that the positive effect of midges on benthic production weakened the negative density dependence at low to intermediate larval densities. However, this benefit disappeared at high densities when midge consumption of primary producers exceeded their positive effects on primary production through ecosystem engineering. Our results illustrate how ecosystem engineering can alter density-dependent feedbacks for engineer populations.

The relative contributions of benthic and pelagic primary production affect ecosystem function, but studies documenting natural variation in the partitioning of production (i.e., autotrophic structure) are uncommon. This study examines autotrophic structure of shallow Lake Mývatn over 7 summers (2012–2018). We used routine measurements of benthic gross primary production (GPP), pelagic chlorophyll a concentrations, and pelagic production to estimate benthic and pelagic maximum productivity (P_max) across summers. With these parameters and corresponding incident light and water clarity data, we estimated in situ benthic and pelagic GPP. Our results demonstrated substantial variation in Mývatn’s autotrophic structure within and across summers. Benthic GPP often exceeded pelagic GPP. However, periods of increased phytoplankton abundance were associated with increased light attenuation and, consequently, declines in benthic GPP. These effects were strongest in 3 summers with dense cyanobacteria blooms, in which the benthic fraction of total production declined from >95% to P_max, implying that past shading by phytoplankton may decrease the photosynthetic potential of benthic producers. Moreover, variation in estimated benthic P_max values can affect the point at which autotrophic structure shifts toward pelagic-dominated conditions. Overall, our study demonstrates that the balance between benthic and pelagic production can vary greatly at intra- and interannual scales because of changes in the photosynthetic capacity of both pelagic and benthic primary producers. Understanding natural variation in lake autotrophic structure may inform how benthic and pelagic production respond to ongoing and future environmental changes.