Reconstructing midge consumer-resource dynamics using carbon stable isotope signatures of archived specimens.

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 δ13 C values reflect those of resources they consumed as larvae. We used the time series for population abundance and δ13 C 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 δ13 C values preceded peaks in the midge population, and δ13 C 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 δ13 C values, which we expected to mirror primary production. Following from this explanation, high benthic production (enriched δ13 C values) would contribute to increased midge abundance, and high midge abundance would result in declining benthic production (depleted δ13 C 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 δ13 C 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.

Numbers and distribution of the Great Cormorant in Iceland: Limitation at the regional and metapopulation level.

We studied a metapopulation of great cormorant (Phalacrocorax carbo) in Iceland, using complete aerial censuses of nests in 25 years during 1975-2015. Age composition was estimated in 1998-2014 by ground surveys in September and February. Brood size was estimated from aerial photographs in 2007-2015.Weather, food, breeding habitat, and density were considered as explanatory variables when examining numerical and distributional changes in the cormorant metapopulation.In 1975-1990 total nest numbers changed little, very low numbers about 1992 were followed by an annual increase of 3.5% in 1994-2015. Total nest numbers were positively correlated with estimates of spawning stocks of cod and saithe and inversely related to the subpolar gyre index (SPG-I).During the increase in 1994-2015, average colony size at first increased and then declined. Habitat use also changed: the proportion of nests on small rocky islets (skerries) at first declined, from 69% to 44% in 1995-2003 and then increased again to about 58% in 2012-2014. Habitat changes were probably a response to changed patterns of human disturbance.Breeding density, as nests per km2 sea <20 m deep, was rather uniform among five defined regions in 1975-1996. Thereafter, densities became much higher in two sheltered regions with kelp forests and in one mostly exposed region. A second exposed region remained low and in the third nest numbers declined markedly. Thus, carrying capacity was higher in sheltered regions where cormorant breeding had historically been depressed by human disturbance.Brood size varied little among regions but declined with the years from about 2.5 to 1.8.The proportion of juveniles in September (fecundity) declined in 1998-2015 from over 0.4 to 0.3 and was inversely correlated with year and nest numbers, if outlier years were excluded, suggesting resource limitation. Survival of juvenile cormorants in September-February was estimated at 0.471 ± 0.066 SE. Commercial fish stocks and climate indices were not correlated with the proportion of juveniles.Annual survival of adults (breeding and nonbreeding) was estimated from nest counts and age composition 1999-2014, as 0.850 ± 0.026 SE and showed no trend in 1998-2014.We conclude that the metapopulation of cormorants in Iceland was resource-limited at two levels: fecundity at the regional and winter survival at the total level.

Relationships between Long-Term Demography and Weather in a Sub-Arctic Population of Common Eider.

Effects of local weather on individuals and populations are key drivers of wildlife responses to climatic changes. However, studies often do not last long enough to identify weather conditions that influence demographic processes, or to capture rare but extreme weather events at appropriate scales. In Iceland, farmers collect nest down of wild common eider Somateria mollissima and many farmers count nests within colonies annually, which reflects annual variation in the number of breeding females. We collated these data for 17 colonies. Synchrony in breeding numbers was generally low between colonies. We evaluated 1) demographic relationships with weather in nesting colonies of common eider across Iceland during 1900-2007; and 2) impacts of episodic weather events (aberrantly cold seasons or years) on subsequent breeding numbers. Except for episodic events, breeding numbers within a colony generally had no relationship to local weather conditions in the preceding year. However, common eider are sexually mature at 2-3 years of age and we found a 3-year time lag between summer weather and breeding numbers for three colonies, indicating a positive effect of higher pressure, drier summers for one colony, and a negative effect of warmer, calmer summers for two colonies. These findings may represent weather effects on duckling production and subsequent recruitment. Weather effects were mostly limited to a few aberrant years causing reductions in breeding numbers, i.e. declines in several colonies followed severe winters (1918) and some years with high NAO (1992, 1995). In terms of life history, adult survival generally is high and stable and probably only markedly affected by inclement weather or aberrantly bad years. Conversely, breeding propensity of adults and duckling production probably do respond more to annual weather variations; i.e. unfavorable winter conditions for adults increase probability of death or skipped breeding, whereas favorable summers can promote boom years for recruitment.

High-amplitude fluctuations and alternative dynamical states of midges in Lake Myvatn.

Complex dynamics are often shown by simple ecological models and have been clearly demonstrated in laboratory and natural systems. Yet many classes of theoretically possible dynamics are still poorly documented in nature. Here we study long-term time-series data of a midge, Tanytarsus gracilentus (Diptera: Chironomidae), in Lake Myvatn, Iceland. The midge undergoes density fluctuations of almost six orders of magnitude. Rather than regular cycles, however, these fluctuations have irregular periods of 4-7 years, indicating complex dynamics. We fit three consumer-resource models capable of qualitatively distinct dynamics to the data. Of these, the best-fitting model shows alternative dynamical states in the absence of environmental variability; depending on the initial midge densities, the model shows either fluctuations around a fixed point or high-amplitude cycles. This explains the observed complex population dynamics: high-amplitude but irregular fluctuations occur because stochastic variability causes the dynamics to switch between domains of attraction to the alternative states. In the model, the amplitude of fluctuations depends strongly on minute resource subsidies into the midge habitat. These resource subsidies may be sensitive to human-caused changes in the hydrology of the lake, with human impacts such as dredging leading to higher-amplitude fluctuations. Tanytarsus gracilentus is a key component of the Myvatn ecosystem, representing two-thirds of the secondary productivity of the lake and providing vital food resources to fish and to breeding bird populations. Therefore the high-amplitude, irregular fluctuations in midge densities generated by alternative dynamical states dominate much of the ecology of the lake.