Additive decompositions for rings of modular forms

We study rings of integral modular forms for congruence subgroups as modules over the ring of integral modular forms for the full modular group. In many cases these modules are free or decompose at least into well-understood pieces. We apply this to characterize which rings of modular forms are Cohen--Macaulay and to prove finite generation results. These theorems are based on decomposition results about vector bundles on the compactified moduli stack of elliptic curves.Comment: Complete revision. Comments welcome. arXiv admin note: text overlap with arXiv:1609.0926

Functional group diversity is key to Southern Ocean benthic carbon pathways

High latitude benthos are globally important in terms of accumulation and storage of ocean carbon, and the feedback this is likely to have on regional warming. Understanding this ecosystem service is important but difficult because of complex taxonomic diversity, history and geography of benthic biomass. Using South Georgia as a model location (where the history and geography of benthic biology is relatively well studied) we investigated whether the composition of functional groups were critical to benthic accumulation, immobilization and burial pathway to sequestration–and also aid their study through simplification of identification. We reclassified [1], [2]) morphotype and carbon mass data to 13 functional groups, for each sample of 32 sites around the South Georgia continental shelf. We investigated the influence on carbon accumulation, immobilization and sequestration estimate by multiple factors including the compositions of functional groups. Functional groups showed high diversity within and between sites, and within and between habitat types. Carbon storage was not linked to a functional group in particular but accumulation and immobilization increased with the number of functional groups present and the presence of hard substrata. Functional groups were also important to carbon burial rate, which increased with the presence of mixed (hard and soft substrata). Functional groups showed high surrogacy for taxonomic composition and were useful for examining contrasting habitat categorization. Functional groups not only aid marine carbon storage investigation by reducing time and the need for team size and speciality, but also important to benthic carbon pathways per se. There is a distinct geography to seabed carbon storage; seabed boulder-fields are hotspots of carbon accumulation and immobilization, whilst the interface between such boulder-fields and sediments are key places for burial and sequestration

Assessing meiofaunal variation among individuals utilising morphological and molecular approaches: an example using the Tardigrada

<p>Abstract</p> <p>Background</p> <p>Meiofauna – multicellular animals captured between sieve size 45 μm and 1000 μm – are a fundamental component of terrestrial, and marine benthic ecosystems, forming an integral element of food webs, and playing a critical roll in nutrient recycling. Most phyla have meiofaunal representatives and studies of these taxa impact on a wide variety of sub-disciplines as well as having social and economic implications. However, studies of variation in meiofauna are presented with several important challenges. Isolating individuals from a sample substrate is a time consuming process, and identification requires increasingly scarce taxonomic expertise. Finding suitable morphological characters in many of these organisms is often difficult even for experts. Molecular markers are extremely useful for identifying variation in morphologically conserved organisms. However, for many species markers need to be developed <it>de novo</it>, while DNA can often only be extracted from pooled samples in order to obtain sufficient quantity and quality. Importantly, multiple independent markers are required to reconcile gene evolution with species evolution. In this primarily methodological paper we provide a proof of principle of a novel and effective protocol for the isolation of meiofauna from an environmental sample. We also go on to illustrate examples of the implications arising from subsequent screening for genetic variation at the level of the individual using ribosomal, mitochondrial and single copy nuclear markers.</p> <p>Results</p> <p>To isolate individual tardigrades from their habitat substrate we used a non-toxic density gradient media that did not interfere with downstream biochemical processes. Using a simple DNA release technique and nested polymerase chain reaction with universal primers we were able amplify multi-copy and, to some extent, single copy genes from individual tardigrades. Maximum likelihood trees from ribosomal 18S, mitochondrial <it>cytochrome oxidase subunit </it>1, and the single copy nuclear gene <it>Wingless </it>support a recent study indicating that the family Hypsibiidae is a non-monophyletic group. From these sequences we were able to detect variation between individuals at each locus that allowed us to identify the presence of cryptic taxa that would otherwise have been overlooked.</p> <p>Conclusion</p> <p>Molecular results obtained from individuals, rather than pooled samples, are a prerequisite to enable levels of variation to be placed into context. In this study we have provided a proof of principle of this approach for meiofaunal tardigrades, an important group of soil biota previously not considered amenable to such studies, thereby paving the way for more comprehensive phylogenetic studies using multiple nuclear markers, and population genetic studies.</p

Against the flow: evidence of multiple recent invasions of warmer continental shelf waters by a Southern Ocean brittle star

The Southern Ocean is anomalously rich in benthos. This biodiversity is native, mostly endemic and perceived to be uniquely threatened from climate- and anthropogenically- mediated invasions. Major international scientific effort throughout the last decade has revealed more connectivity than expected between fauna north and south of the worlds strongest marine barrier – the Polar Front (the strongest jet of the Antarctic Circumpolar Current). To date though, no research has demonstrated any radiations of marine taxa out from the Southern Ocean, except at abyssal depths (where conditions differ much less). Our phylogeographic investigation of one of the most ubiquitous and abundant clades at high southern latitudes, the ophiuroids (brittlestars), shows that one of them, Ophiura lymani, has gone against the flow. Remarkably our genetic data suggest that O. lymani has successfully invaded the South American shelf from Antarctica at least three times, in recent (Pleistocene) radiation. Many previous studies have demonstrated links within clades across the PF this is the first in which northwards directional movement of a shelf-restricted species is the only convincing explanation. Rapid, recent, regional warming is likely to facilitate multiple range shift invasions into the Southern Ocean, whereas movement of cold adapted fauna (considered highly stenothermal) out of the Antarctic to warmer shelves has, until now, seemed highly unlikely

Icebergs, sea ice, blue carbon and Antarctic climate feedbacks

Sea ice, including icebergs, has a complex relationship with the carbon held within animals (blue carbon) in the polar regions. Sea-ice losses around West Antarctica's continental shelf generate longer phytoplankton blooms but also make it a hotspot for coastal iceberg disturbance. This matters because in polar regions ice scour limits blue carbon storage ecosystem services, which work as a powerful negative feedback on climate change (less sea ice increases phytoplankton blooms, benthic growth, seabed carbon and sequestration). This resets benthic biota succession (maintaining regional biodiversity) and also fertilizes the ocean with nutrients, generating phytoplankton blooms, which cascade carbon capture into seabed storage and burial by benthos. Small icebergs scour coastal shallows, whereas giant icebergs ground deeper, offshore. Significant benthic communities establish where ice shelves have disintegrated (giant icebergs calving), and rapidly grow to accumulate blue carbon storage. When 5000 km2 giant icebergs calve, we estimate that they generate approximately 106 tonnes of immobilized zoobenthic carbon per year (t C yr−1). However, their collisions with the seabed crush and recycle vast benthic communities, costing an estimated 4 × 104 t C yr−1. We calculate that giant iceberg formation (ice shelf disintegration) has a net potential of approximately 106 t C yr−1 sequestration benefits as well as more widely known negative impacts

Extremes in benthic ecosystem services; Blue carbon natural capital shallower than 1000 m in isolated, small and young Ascension Island’s EEZ

Biodiversity tends to decrease with increasing isolation and reduced habitat size, and increase with habitat age. Ascension Island and its seamounts are small, isolated and relatively young, yet, within its soon-to-be designated major Marine Protected Area, harbour patchily dense life in the shallows and cold water corals in deeper waters. With few local threats, global climate related stressors (e.g. temperature and acidification) and arguably plastic pollution are key issues for its survival and continued provision of ecosystem services. We evaluate the accumulated carbon in benthos around Ascension Island’s EEZ shallower than 1000 m using data from two research cruises in 2015 and 2017 through seabed mapping, seabed camera imagery and collections of benthos using a mini-Agassiz trawl. Benthos shallower than 1000 m essentially comprises the coastal waters around Ascension Island and three seamounts (Harris-Stewart, Grattan and Un-named). There is considerable societal benefit from benthic carbon storage and sequestration through its mitigation value buffering climate change. This service is often termed ‘blue carbon’. Overall we estimate that there is at least 43,000 tonnes of blue carbon, on the 3% of Ascension Island EEZ’s seabed which is <1000 m, mainly in the form of cold coral reefs. Two thirds of that occurs around the main island of Ascension, but it is very unevenly distributed on the seabed. Seabed roughness (e.g. rocky outcrops) seems most important for the development of blue carbon hotspots. About 21% of the total blue carbon is considered to be sequestered (removed from the carbon cycle for 100+ years) = 9060 tonnes Carbon. At the 2019 Shadow Price of Carbon the proportion of CO2 considered sequestered is £29-59. As 9060 t C this is equivalent to 33,250 t CO2, which in 2019 is valued at approximately £1-2 million. With time, this increases with rising value of carbon, but also annual increment of carbon deposition, to £2-4 million by 2030. Thus even when biogeographic values of isolation, size and age are least favourable to biodiversity, the natural capital stock and future services of benthic ecosystems can be considerable and generate a quantifiable economic return on their conservation

Amundsen Sea Mollusca from the BIOPEARL II expedition

Information regarding the molluscs in this dataset is based on the epibenthic sledge (EBS) samples collected during the cruise BIOPEARL II / JR179 RRS James Clark Ross in the austral summer 2008. A total of 35 epibenthic sledge deployments have been performed at five locations in the Amundsen Sea at Pine Island Bay (PIB) and the Amundsen Sea Embayment (ASE) at depths ranging from 476 to 3501m. This presents a unique and important collection for the Antarctic benthic biodiversity assessment as the Amundsen Sea remains one of the least known regions in Antarctica. Indeed the work presented in this dataset is based on the first benthic samples collected with an EBS in the Amundsen Sea. However we assume that the data represented are an underestimation of the real fauna present in the Amundsen Sea. In total 9261 specimens belonging to 6 classes 55 families and 97 morphospecies were collected. The species richness per station varied between 6 and 43. Gastropoda were most species rich 50 species followed by Bivalvia (37), Aplacophora (5), Scaphopoda (3) and one from each of Polyplacophora and Monoplacophora

East Weddell Sea echinoids from the JR275 expedition

Information regarding the echinoids in this dataset is based on the Agassiz Trawl (AGT) and epibenthic sledge (EBS) samples collected during the British Antarctic Survey cruise JR275 on the RRS James Clark Ross in the austral summer 2012. A total of 56 (1 at the South Orkneys and 55 in the Eastern Weddell Sea) Agassiz Trawl and 18 (2 at the South Orkneys and 16 in the Eastern Weddell Sea) epibenthic sledge deployments were performed at depths ranging from ~280 to ~2060 m. This presents a unique collection for the Antarctic benthic biodiversity assessment of an important group of benthic invertebrates. In total 487 specimens belonging to six families, 15 genera, and 22 morphospecies were collected. The species richness per station varied between one and six. Total species richness represents 27% of the 82 echinoid species ever recorded in the Southern Ocean (David et al. 2005b, Pierrat et al. 2012, Saucède et al. 2014). The Cidaridae (sub-family Ctenocidarinae) and Schizasteridae are the two most speciose families in the dataset. They comprise seven and nine species respectively. This is illustrative of the overall pattern of echinoid diversity in the Southern Ocean where 65% of Antarctic species belong to the families Schizasteridae and Cidaridae (Pierrat et al. 2012)

The influence of glacial melt and retreat on the nutritional condition of the bivalve Nuculana inaequisculpta (Protobranchia: Nuculanidae) in the West Antarctic Peninsula

Due to climate change, numerous ice bodies have been lost in the West Antarctic Peninsula (WAP). As a consequence, deglaciation is expected to impact the marine environment and its biota at physiological and ecosystem levels. Nuculana inaequisculpta is a marine bivalve widely distributed around Antarctica that plays an important role for ecosystem functioning. Considering that N. inaequisculpta inhabits coastal areas under effect of glacial melt and retreat, impacts on its nutritional condition are expected due to alterations on its physiology and food availability. To test this hypothesis, biochemical composition (lipids, proteins, and fatty acids) and energy content were measured in individuals of N. inaequisculpta collected in a fjord at different distances to the retreating glacier in the WAP. Oceanographic parameters of the top and bottom-water layers (temperature, salinity, dissolved oxygen, and chlorophyll-a) were measured to investigate how the environment changes along the fjord. Results showed that surface oceanographic parameters displayed a lower temperature and dissolved oxygen, but a higher salinity and chlorophyll-a content at nearest compared to farthest sites to the glacier. In contrast, a lower temperature and chlorophyll-a, and a higher salinity and dissolved oxygen was measured in the bottom-water layer toward the glacier. N. inaequisculpta had a higher amount of lipids (17.42 ± 3.24 vs. 12.16 ± 3.46%), protein (24.34 ± 6.12 vs. 21.05 ± 2.46%) and energy content (50.57 ± 6.97 J vs. 39.14 ± 5.80 J) in the farthest compared to the nearest site to the glacier. No differences were found in total fatty acids among all sites. It seems likely that lower individual fitness related to proximity to the glacier would not be related to nutritional quality of sediment food, but rather to food quantity

Gene flow in the Antarctic bivalve Aequiyoldia eightsii (Jay, 1839) suggests a role for the Antarctic Peninsula Coastal Current in larval dispersal

The Antarctic Circumpolar Current (ACC) dominates the open-ocean circulation of the Southern Ocean, and both isolates and connects the Southern Ocean biodiversity. However, the impact on biological processes of other Southern Ocean currents is less clear. Adjacent to the West Antarctic Peninsula (WAP), the ACC flows offshore in a northeastward direction, whereas the Antarctic Peninsula Coastal Current (APCC) follows a complex circulation pattern along the coast, with topographically influenced deflections depending on the area. Using genomic data, we estimated genetic structure and migration rates between populations of the benthic bivalve Aequiyoldia eightsii from the shallows of southern South America and the WAP to test the role of the ACC and the APCC in its dispersal. We found strong genetic structure across the ACC (between southern South America and Antarctica) and moderate structure between populations of the WAP. Migration rates along the WAP were consistent with the APCC being important for species dispersal. Along with supporting current knowledge about ocean circulation models at the WAP, migration from the tip of the Antarctic Peninsula to the Bellingshausen Sea highlights the complexities of Southern Ocean circulation. This study provides novel biological evidence of a role of the APCC as a driver of species dispersal and highlights the power of genomic data for aiding in the understanding of the influence of complex oceanographic processes in shaping the population structure of marine species