Reply to formal comment on Griffiths et al. (2017) submitted by Gajewski (2020).

Gajewski offers a formal comment on Griffiths et al. (2017), a paper that explored how microclimates and their varying ice cover regimes on lakes and ponds in Arctic regions modified the diatom assemblage responses to recent warming. One of Gajewski's main criticisms is that the microclimate classification scheme used in Griffiths et al. (2017) is merely anecdotal; a claim which ignores the value of observational evidence and misunderstands the frequency that each site was visited or surveyed. We clarify that the study sites were visited multiple times via recurrent aerial surveys and ground observations dating back to the 1970s, which supports our microclimate classification scheme. Many of Gajewski's claims regarding climate, catchment characteristics, and ice melting properties from field locations he has not visited were refuted by veteran Arctic scientists with long-term field experience in these regions. In addition, Gajewski makes several criticisms concerning radioisotopic dating, core chronology, sediment mixing, diagenesis, and preservation of bioindicators that relate more to general paleolimnological assumptions than to conclusions reached by Griffiths et al. (2017). Research from the 1980s and 1990s, when scientific consensus on these issues was first reached, readily show that the methodologies and data interpretation of Griffiths et al. (2017) are sound. We appreciate the opportunity to expound on the finer details of the Griffiths et al. (2017) paper, work based on field research by the study's co-authors spanning almost three decades, with additional observations from colleagues dating back to the 1970s. We address Gajewski's criticisms with relevant literature, expert statements, and a few clarifying comments.

Contrasting the ecological effects of decreasing ice cover versus accelerated glacial melt on the High Arctic's largest lake.

Lake Hazen, the High Arctic's largest lake, has received an approximately 10-fold increase in glacial meltwater since its catchment glaciers shifted from net mass gain to net mass loss in 2007 common era (CE), concurrent with recent warming. Increased glacial meltwater can alter the ecological functioning of recipient aquatic ecosystems via changes to nutrient budgets, turbidity and thermal regimes. Here, we examine a rare set of five high-resolution sediment cores collected in Lake Hazen between 1990 and 2017 CE to investigate the influence of increased glacial meltwater versus alterations to lake ice phenology on ecological change. Subfossil diatom assemblages in all cores show two major shifts over the past approximately 200 years including: (i) a proliferation of pioneering, benthic taxa at approximately 1900 CE from previously depauperate populations; and (ii) a rise in planktonic taxa beginning at approximately 1980 CE to present-day dominance. The topmost intervals from each sequentially collected core provide exact dates and demonstrate that diatom regime shifts occurred decades prior to accelerated glacial inputs. These data show that diatom assemblages in Lake Hazen are responding primarily to intrinsic lake factors linked to decreasing duration of lake ice and snow cover rather than to limnological impacts associated with increased glacial runoff.

Ice-cover is the principal driver of ecological change in High Arctic lakes and ponds.

Recent climate change has been especially pronounced in the High Arctic, however, the responses of aquatic biota, such as diatoms, can be modified by site-specific environmental characteristics. To assess if climate-mediated ice cover changes affect the diatom response to climate, we used paleolimnological techniques to examine shifts in diatom assemblages from ten High Arctic lakes and ponds from Ellesmere Island and nearby Pim Island (Nunavut, Canada). The sites were divided a priori into four groups ("warm", "cool", "cold", and "oasis") based on local elevation and microclimatic differences that result in differing lengths of the ice-free season, as well as about three decades of personal observations. We characterized the species changes as a shift from Condition 1 (i.e. a generally low diversity, predominantly epipelic and epilithic diatom assemblage) to Condition 2 (i.e. a typically more diverse and ecologically complex assemblage with an increasing proportion of epiphytic species). This shift from Condition 1 to Condition 2 was a consistent pattern recorded across the sites that experienced a change in ice cover with warming. The "warm" sites are amongst the first to lose their ice covers in summer and recorded the earliest and highest magnitude changes. The "cool" sites also exhibited a shift from Condition 1 to Condition 2, but, as predicted, the timing of the response lagged the "warm" sites. Meanwhile some of the "cold" sites, which until recently still retained an ice raft in summer, only exhibited this shift in the upper-most sediments. The warmer "oasis" ponds likely supported aquatic vegetation throughout their records. Consequently, the diatoms of the "oasis" sites were characterized as high-diversity, Condition 2 assemblages throughout the record. Our results support the hypothesis that the length of the ice-free season is the principal driver of diatom assemblage responses to climate in the High Arctic, largely driven by the establishment of new aquatic habitats, resulting in increased diversity and the emergence of novel growth forms and epiphytic species.

Determining diatom ecotones and their relationship to terrestrial ecoregion designations in the central Canadian Arctic Islands.

Ecotones are key areas for the detection of global change because many are predicted to move with shifts in climate. Prince of Wales Island, in the Canadian Arctic Archipelago, spans the transition between mid- to high-Arctic ecoregions. We analyzed limnological variables and recent diatom assemblages from its lakes and ponds to determine if assemblages reflected this ecotone. Limnological gradients were short, and water chemistry explained 20.0% of diatom variance in a redundancy analysis (RDA), driven primarily by dissolved organic carbon, Ca and SO4 . Most taxa were small, benthic forms; key taxa such as planktonic Cyclotella species were restricted to the warmer, southern portion of the study area, while benthic Staurosirella were associated with larger, ice-dominated lakes. Nonetheless, there were no significant changes in diatom assemblages across the mid- to high-Arctic ecoregion boundary. We combined our data set with one from nearby Cornwallis Island to expand the study area and lengthen its environmental gradients. Within this expanded data set, 40.6% of the diatom variance was explained by a combination of water chemistry and geographic variables, and significant relationships were revealed between diatom distributions and key limnological variables, including pH, specific conductivity, and chl-a. Using principal coordinates analysis, we estimated community turnover with latitude and applied piecewise linear regression to determine diatom ecotone positions. A pronounced transition was present between Prince of Wales Island and the colder, more northerly Cornwallis Island. These data will be important in detecting any future northward ecotone movement in response to predicted Arctic climate warming in this highly sensitive region.

Comparison of Freshwater Diatom Assemblages from a High Arctic Oasis to Nearby Polar Desert Sites and Their Application to Environmental Inference Models.

Arctic oases are regions of atypical warmth and relatively high biological production and diversity. They are small in area (<5 km(2) ) and uncommon in occurrence, yet they are relatively well studied due to the abundance of plant and animal life contained within them. A notable exception is the lack of research on freshwater ecosystems within polar oases. Here, we aim to increase our understanding of freshwater diatom ecology in polar oases. Diatoms were identified and enumerated from modern sediments collected in 23 lakes and ponds contained within the Lake Hazen oasis on Ellesmere Island, and compared with diatom assemblages from 29 sites located outside of the oasis across the northern portion of the island. There were significant differences in water chemistry variables between oasis and northern sites, with oasis sites having higher conductivity and greater concentrations of nutrients and related variables such as dissolved organic carbon (DOC). Taxa across all sites were typical of those recorded in Arctic freshwaters, with species from the genera Achnanthes sensu lato, Fragilaria sensu lato, and Nitzschia dominating the assemblages. A correspondence analysis (CA) ordination showed that oasis sites generally plotted separately from the northern sites, although the sites also appear to plot separately based on whether they were lakes or ponds. Canonical correspondence analysis (CCA) identified specific conductivity, DOC, and SiO2 as explaining significant (P < 0.05) and additional amounts of variation in the diatom data set. The most robust diatom-based inference model was generated for DOC, which will provide useful reconstructions on long-term changes in paleo-optics of high Arctic lakes.

Climate change and mercury accumulation in Canadian high and subarctic lakes.

Mercury (Hg) profiles were compared to profiles of climate indicators including microfossil remains and algal-derived or S2 carbon (C) in dated sediment cores from 14 lakes spanning latitudinal and longitudinal gradients across the Canadian high and subarctic. Hg fluxes increased postindustrialization (post-∼1850) in 11 of these lakes (postindustrialization Hg fluxes (ΔHgF(F)) = 2-24 µg m(-2) y(-1)). Correction of HgF(F) for catchment contributions demonstrated that Hg deposition originating from catchment-independent factors, such as atmospheric deposition, increased since industrialization in all 14 lakes. Several of these lakes also showed postindustrial shifts in algal assemblages consistent with climate-induced changes. Eleven lakes showed post-1850s increases in S2F(F), suggesting that lake primary productivity has recently increased in the majority of our sites (ΔS2F(F) = 0.1-4 g m(-2) y(-1)). Other studies have interpreted significant relationships between Hg:S2 concentrations in Arctic sediment as support for the algal scavenging hypothesis, which postulates that Hg fluxes to Arctic sediments are largely driven by S2. However, in six of our lakes we observed no Hg:S2 relationship, and in one lake a significant negative Hg:S2 relationship was observed due to increased Hg and decreased S2 C deposition during the postindustrialization period. In six of the seven lakes where a significant positive Hg:S2 relationship was observed, algal assemblages either did not change through time or the timing of the shifts did not correspond to changes in Hg deposition. Our results demonstrate that, although Arctic lakes are experiencing a myriad of changes, including increased Hg and S2 deposition or changing algal assemblages, increased lake primary productivity does not appear to be driving changes in Hg fluxes to sediments.

Trophic position influences the efficacy of seabirds as metal biovectors.

Seabirds represent a well documented biological transport pathway of nutrients from the ocean to the land by nesting in colonies and providing organic subsidies (feces, carcasses, dropped food) to these sites. We investigated whether seabirds that feed at different trophic levels vary in their potency as biovectors of metals, which can bioaccumulate through the marine foodweb. Our study site, located on a small island in Arctic Canada, contains the unique scenario of two nearby ponds, one of which receives inputs almost exclusively from upper trophic level piscivores (Arctic terns, Sterna paradisaea) and the other mainly from lower trophic level molluscivores (common eiders, Somateria mollissima). We used dated sediment cores to compare differences in diatoms, metal concentrations and also stable isotopes of nitrogen (delta(15)N), which reflect trophic position. We show that the seabirds carry species-specific mixtures of metals that are ultimately shunted to their nesting sites. For example, sediments from the tern-affected pond recorded the highest levels of delta(15)N and the greatest concentrations of metals that are known to bioaccumulate, including Hg and Cd. In contrast, the core from the eider-affected site registered lower delta(15)N values, but higher concentrations of Pb, Al, and Mn. These metals have been recorded at their greatest concentrations in eiders relative to other seabirds, including Arctic terns. These data indicate that metals may be used to track seabird population dynamics, and that some metal tracers may even be species-specific. The predominance of large seabird colonies on every continent suggests that similar processes are operating along coastlines worldwide.

An isotopic investigation of mercury accumulation in terrestrial food webs adjacent to an Arctic seabird colony.

At Cape Vera (Devon Island, Nunavut, Canada), a seabird colony of northern fulmars (Fulmarus glacialis) congregates and releases nutrients through the deposition of guano to the coastal terrestrial environment, thus creating nutrient-fertilized habitats important to insects, birds, and mammals. Here we determined whether mercury was similarly enriched in various terrestrial food web components in this High Arctic coastal ecosystem due to seabird inputs. Stable isotopes (delta(15)N, delta(13)C) were used to identify trophic linkages and possible routes of contaminant transfer in the food web. Values of delta(15)N were significantly higher in lichens and certain plants collected closer to the bird colony, demonstrating a gradient of seabird influence, and were higher at Cape Vera than our reference site at Cape Herschel, on eastern Ellesmere Island, an area relatively unaffected by seabirds. In contrast, delta(13)C showed little variation among terrestrial species, suggesting minimal influence by seabirds. Concentrations of total mercury (THg) in primary producers and phyto/zooplankton were not significantly correlated with distance from the seabird colony or delta(15)N values, and were similar to other taxa from the High Arctic. Our results provide novel data on THg in several Arctic taxa where concentrations have not been reported previously. Moreover, the analyses indicate that delta(15)N is significantly enriched in the adjacent environment by guano fertilization, but our study was unable to show an enrichment of THg and delta(13)C in the terrestrial food web near the seabird colony.

Accelerated delivery of polychlorinated biphenyls (PCBs) in recent sediments near a large seabird colony in Arctic Canada.

Polychlorinated biphenyls (PCBs) were measured in sediment cores from ponds located near a large seabird colony at Cape Vera, Devon Island, Arctic Canada. Surface sediment PCB concentrations were approximately 5x greater in seabird-affected sites relative to a nearby control pond and were correlated with independent indicators of seabird activity including, sedimentary delta(15)N and lakewater chlorophyll a and cadmium concentrations. PCB fluxes were amongst the highest recorded from the High Arctic, ranging from 290 to 2400 ng m(-2) yr(-1). Despite a widespread ban of PCBs in the mid-1970s, PCB accumulation rates in our cores increased, with the highest values recorded in the most recent sediments. Possible mechanisms for the recent PCB increases include a vertical flux step driven by seabird-delivered nutrients and/or delayed loading of PCBs from the catchment into the ponds. The high PCB levels recorded in the seabird-affected sites suggest that seabird colonies are exposing coastal ecosystems to elevated levels of contaminants.

Seabird-driven shifts in Arctic pond ecosystems.

Migratory animals such as seabirds, salmon and whales can transport large quantities of nutrients across ecosystem boundaries, greatly enriching recipient food webs. As many of these animals biomagnify contaminants, they can also focus pollutants at toxic levels. Seabirds arguably represent the most significant biovectors of nutrients and contaminants from the ocean to the land, given their sheer numbers and global distribution. However, long-term census data on seabirds are rare. Using palaeolimnological proxies, we show that a colony of Arctic seabirds has experienced climate-induced population increases in recent decades. We then document increasing concentrations of contaminants, including polychlorinated biphenyls and cadmium, in pond sediments that are linked to biotransport by seabirds. Our findings suggest that climate-related shifts in global seabird populations will have the unexpected consequence of restructuring coastal ecosystems.

Crossing the final ecological threshold in high Arctic ponds.

A characteristic feature of most Arctic regions is the many shallow ponds that dot the landscape. These surface waters are often hotspots of biodiversity and production for microorganisms, plants, and animals in this otherwise extreme terrestrial environment. However, shallow ponds are also especially susceptible to the effects of climatic changes because of their relatively low water volumes and high surface area to depth ratios. Here, we describe our findings that some high Arctic ponds, which paleolimnological data indicate have been permanent water bodies for millennia, are now completely drying during the polar summer. By comparing recent pond water specific conductance values to similar measurements made in the 1980s, we link the disappearance of the ponds to increased evaporation/precipitation ratios, probably associated with climatic warming. The final ecological threshold for these aquatic ecosystems has now been crossed: complete desiccation.

Arctic seabirds transport marine-derived contaminants.

Long-range atmospheric transport of pollutants is generally assumed to be the main vector for arctic contamination, because local pollution sources are rare. We show that arctic seabirds, which occupy high trophic levels in marine food webs, are the dominant vectors for the transport of marine-derived contaminants to coastal ponds. The sediments of ponds most affected by seabirds had 60 times higher DDT, 25 times higher mercury, and 10 times higher hexachlorobenzene concentrations than nearby control sites. Bird guano greatly stimulates biological productivity in these extreme environments but also serves as a major source of industrial and agricultural pollutants in these remote ecosystems.

Climate-driven regime shifts in the biological communities of arctic lakes.

Fifty-five paleolimnological records from lakes in the circumpolar Arctic reveal widespread species changes and ecological reorganizations in algae and invertebrate communities since approximately anno Domini 1850. The remoteness of these sites, coupled with the ecological characteristics of taxa involved, indicate that changes are primarily driven by climate warming through lengthening of the summer growing season and related limnological changes. The widespread distribution and similar character of these changes indicate that the opportunity to study arctic ecosystems unaffected by human influences may have disappeared.

Prehistoric Inuit whalers affected Arctic freshwater ecosystems.

It is commonly assumed that High Arctic lakes and ponds were not affected by direct local human activities before the arrival of Europeans, because most native peoples were primarily nomadic, maintained relatively low population densities, and practiced unintrusive hunting and gathering technologies. Our archeological and paleolimnological data show that this was not always the case. Thule Inuit whalers, whose winter settlements consisted of houses constructed from the bones of bowhead whales on Somerset Island between about anno Domini 1200 and 1600, markedly changed pond water quality and ecology. The arrival of whalers 8 centuries ago caused marked changes in water chemistry and the expansion of moss substrates. Although whalers abandoned the area >4 centuries ago, the legacy of these human disturbances is still evident in the pond's present-day limnology and is characterized by elevated nutrient concentrations and atypical biota. This is the earliest reported paleolimnological record of changes in aquatic ecology associated with local human activities in Canada or the United States, or for any circumpolar ecosystem.

Fisheries productivity in the northeastern Pacific Ocean over the past 2,200 years.

Historical catch records suggest that climatic variability has had basin-wide effects on the northern Pacific and its fish populations, such as salmon, sardines and anchovies. However, these records are too short to define the nature and frequency of patterns. We reconstructed approximately 2,200-year records of sockeye salmon abundance from sediment cores obtained from salmon nursery lakes on Kodiak island, Alaska. Large shifts in abundance, which far exceed the decadal-scale variability recorded during the past 300 years, occurred over the past two millennia. A marked, multi-centennial decline in Alaskan sockeye salmon was apparent from approximately 100 BC to AD 800, but salmon were consistently more abundant from AD 1200 to 1900. Over the past two millennia, the abundances of Pacific sardine and Northern anchovy off the California coast, and of Alaskan salmon, show several synchronous patterns of variability. But sardines and anchovies vary out of phase with Alaskan salmon over low frequency, which differs from the pattern detected in historical records. The coherent patterns observed across large regions demonstrate the strong role of climatic forcing in regulating northeastern Pacific fish stocks.