Agata Weydmann

Tipping Points in the Arctic: Eyeballing or Statistical Significance?

Arctic ecosystems have experienced and are projected to experience continued large increases in temperature and declines in sea ice cover. It has been hypothesized that small changes in ecosystem drivers can fundamentally alter ecosystem functioning, and that this might be particularly pronounced for Arctic ecosystems. We present a suite of simple statistical analyses to identify changes in the statistical properties of data, emphasizing that changes in the standard error should be considered in addition to changes in mean properties. The methods are exemplified using sea ice extent, and suggest that the loss rate of sea ice accelerated by factor of ~5 in 1996, as reported in other studies, but increases in random fluctuations, as an early warning signal, were observed already in 1990. We recommend to employ the proposed methods more systematically for analyzing tipping points to document effects of climate change in the Arctic.

Responses in Arctic marine carbon cycle processes: conceptual scenarios and implications for ecosystem function

The Arctic Ocean is one of the fastest changing oceans, plays an important role in global carbon cycling and yet is a particularly challenging ocean to study. Hence, observations tend to be relatively sparse in both space and time. How the Arctic functions, geophysically, but also ecologically, can have significant consequences for the internal cycling of carbon, and subsequently influence carbon export, atmospheric CO2 uptake and food chain productivity. Here we assess the major carbon pools and associated processes, specifically summarizing the current knowledge of each of these processes in terms of data availability and ranges of rates and values for four geophysical Arctic Ocean domains originally described by Carmack & Wassmann (2006): inflow shelves, which are Pacific-influenced and Atlantic-influenced; interior, river-influenced shelves; and central basins. We attempt to bring together knowledge of the carbon cycle with the ecosystem within each of these different geophysical settings, in order to provide specialist information in a holistic context. We assess the current state of models and how they can be improved and/or used to provide assessments of the current and future functioning when observational data are limited or sparse. In doing so, we highlight potential links in the physical oceanographic regime, primary production and the flow of carbon within the ecosystem that will change in the future. Finally, we are able to highlight priority areas for research, taking a holistic pan-Arctic approach.

Distribution of small phytoflagellates along an Arctic fjord transect

Phytoflagellates <10 μm substantially contribute to the abundance, biomass and primary production in polar waters, but information on the distribution of specific groups is scarce. We applied catalysed reporter deposition-fluorescence in situ hybridization to investigate the distribution of total phytoflagellates and of eight specific groups along a 100 km transect west off Kongsfjorden (Spitsbergen) from 29 to 31 July 2010. Phytoflagellates contributed to >75% of the depth-integrated abundance and biomass of total eukaryotes <10 μm at all stations. Their depth-integrated abundance and biomass decreased along the transect from 1.5 × 10(12)  cells m(-2) (6.6 × 10(12)  pgC m(-2) ) at the outermost station to 1.7 × 10(10)  cells m(-2) (4.7 × 10(10) pgC m(-2) ) at the innermost station. Chlorophytes contributed to the total abundance of phytoflagellates with a range from 13% to 87% (0.7-30.5 × 10(3)  cells ml(-1) ), and predominated in open waters. The contribution of haptophytes was < 1-38% (10-4500 cells ml(-1) ). The other groups represented <10%. The temperature and salinity positively correlated with the total abundance of phytoflagellates, chlorophytes, haptophytes, bolidophytes and pelagophytes. Cryptophytes, pedinellids and pavlovophytes were negatively associated with the nutrient concentrations. The community composition of phytoflagellates changed along the transect, which could have implications on food web dynamics and biogeochemical cycles between the open ocean environment and Kongsfjorden investigated here.

The effect of temperature on egg development rate and hatching success in Calanus glacialis and C. finmarchicus

The pelagic copepods Calanus glacialis and C. finmarchicus are important components of Arctic marine ecosystems. Projected climate warming may influence the roles they play in the ecosystem. Arctic C. glacialis and boreal C. finmarchicus eggs were incubated at temperatures of 0, 2.5, 5, 7.5 and 10°C to investigate the effects of increasing temperature on egg development rate and hatching success. The effect of increasing temperature on median development time, described by B[ebreve]lehrádeks temperature function, was examined using a Bayesian approach. For the studied temperature range, we observed the increase of egg development rates with the increasing temperature, although there was no change in hatching success. Calanus finmarchicus eggs hatched significantly faster than C. glacialis above approximately 2°C; the difference was progressively larger at higher temperatures. This may indicate that the boreal species have physiological advantages in areas where ambient temperatures increase, which may lead to C. finmarchicus outcompeting the Arctic species in situations where timing is important, for example, in relation to spring bloom dynamics. Development time to hatching (DH) was evaluated using B[ebreve]lehrádeks model and a set of different assumptions. The models that best fitted our data were those with species-specific parameters: DH (h)=5940 (T+9.7)−1.63 for C. finmarchicus and DH (h)=14168 (T+14)−1.75 for C. glacialis.

Ecosystem maturation follows the warming of the Arctic fjords

Two fjords in West Spitsbergen (Hornsund 77°N and Kongsfjorden 79°N) differ with regard to their exposure towards increasingly warm Atlantic water inflow. Hornsund remains in many respects cooler than Kongsfjorden (on average 2°C SST in summer) and is less influenced by warmer and more saline Atlantic waters. Reported changes in the physical environment (temperature rise, freshwater inflow, salinity drop, turbidity, fast-ice reduction, coastal change) are discussed in the context of biological observations in the pelagic and benthic realms with special reference to krill (Euphausiacea). We conclude that well-documented changes in the physical environment have had little effect on the fjord biota and that both organisms and their ecological functions in the fjords are well adapted to the scale of ongoing change. The observed changes fit the definition of ecosystem maturation, with greater diversity, a more complex food web and dispersed energy flow at the warmer site.

A transcriptome resource for the copepod Calanus glacialis across a range of culture temperatures

The copepod Calanus glacialis plays a key role in the Arctic pelagic ecosystem. Despite its ecological importance and ongoing climate changes, limited knowledge at the genomic level has hindered the understanding of the molecular processes underlying environmental stress responses and ecological adaptation. Transcriptome data was generated from an experiment with C. glacialis copepodite (CV) subjected to five different temperatures. We obtained a total of 512,352 high-quality 454 pyrosequencing reads, which were assembled into 55,562 contigs distributed in 128 KEGG pathways. Functional analysis revealed numerous genes related to diverse biological functions and processes, including members of all major conserved signaling pathways. Comparative analysis of acclimated individuals to experimental temperatures has provided information about gene variations observed in several pathways (e.g. genes involved in energy, lipid and amino acid metabolism were shown to be down-regulated with increasing temperatures). These mRNA sequence resources will facilitate further studies on genomics and physiology-driven molecular processes in C. glacialis and related species.

Microsatellite markers for the Arctic copepod Calanus glacialis and cross-amplification with C. finmarchicus

Calanus glacialis is a major component of Arctic zooplankton and a keystone species in Arctic marine ecosystems. Due to the observed climate warming, its numbers are being reduced to the advantage of a sibling Atlantic species Calanus finmarchicus. We developed and characterized the first set of microsatellite markers in this species to investigate its population genetic structure and dispersal capabilities. Nine polymorphic loci displayed an average of 7.3 alleles (range between 2 and 13) and the levels of expected heterozygosity ranged from 0.039 to 0.806. These provide a valuable tool to understand present connectivity patterns across Arctic regions, look for signatures of past climate effects and predict the response to future climate-driven environmental changes. Additionally, due to the cross-amplification with C. finmarchicus, the markers can be used to discriminate between these sibling species.

Postglacial expansion of the Arctic keystone copepod Calanus glacialis

Calanus glacialis, a major contributor to zooplankton biomass in the Arctic shelf seas, is a key link between primary production and higher trophic levels that may be sensitive to climate warming. The aim of this study was to explore genetic variation in contemporary populations of this species to infer possible changes during the Quaternary period, and to assess its population structure in both space and time. Calanus glacialis was sampled in the fjords of Spitsbergen (Hornsund and Kongsfjorden) in 2003, 2004, 2006, 2009 and 2012. The sequence of a mitochondrial marker, belonging to the ND5 gene, selected for the study was 1249 base pairs long and distinguished 75 unique haplotypes among 140 individuals that formed three main clades. There was no detectable pattern in the distribution of haplotypes by geographic distance or over time. Interestingly, a Bayesian skyline plot suggested that a 1000-fold increase in population size occurred approximately 10,000 years before present, suggesting a species expansion after the Last Glacial Maximum.

Ecosystem maturation process follows the warming of the Arctic fjords

Two fjords in West Spitsbergen (Hornsund 77°N and Kongsfjorden 79°N) differ with regard to their exposure towards increasingly warm Atlantic water inflow. Hornsund remains in many respects cooler than Kongsfjorden (on average 2°C SST in summer) and is less influenced by warmer and more saline Atlantic waters. Reported changes in the physical environment (temperature rise, freshwater inflow, salinity drop, turbidity, fast-ice reduction, coastal change) are discussed in the context of biological observations in the pelagic and benthic realms with special reference to krill (Euphausiacea). We conclude that well-documented changes in the physical environment have had little effect on the fjord biota and that both organisms and their ecological functions in the fjords are well adapted to the scale of ongoing change. The observed changes fit the definition of ecosystem maturation, with greater diversity, a more complex food web and dispersed energy flow at the warmer site.

Mitochondrial genomes of the key zooplankton copepods Arctic Calanus glacialis and North Atlantic Calanus finmarchicus with the longest crustacean non-coding regions

We determined the nearly complete mitochondrial genomes of the Arctic Calanus glacialis and its North Atlantic sibling Calanus finmarchicus, which are key zooplankton components in marine ecosystems. The sequenced part of C. glacialis mitogenome is 27,342 bp long and consists of two contigs, while for C. finmarchicus it is 29,462 bp and six contigs, what makes them the longest reported copepod mitogenomes. The typical set of metazoan mitochondrial genes is present in these mitogenomes, although the non-coding regions (NCRs) are unusually long and complex. The mitogenomes of the closest species C. glacialis and C. finmarchicus, followed by the North Pacific C. sinicus, are structurally similar and differ from the much more typical of deep-water, Arctic C. hyperboreus. This evolutionary trend for the expansion of NCRs within the Calanus mitogenomes increases mitochondrial DNA density, what resulted in its similar density to the nuclear genome. Given large differences in the length and structure of C. glacialis and C. finmarchicus mitogenomes, we conclude that the species are genetically distinct and thus cannot hybridize. The molecular resources presented here: the mitogenomic and rDNA sequences, and the database of repetitive elements should facilitate the development of genetic markers suitable in pursuing evolutionary research in copepods.