Completing the hybridization triangle: the inheritance of genetic incompatibilities during homoploid hybrid speciation in ragworts (Senecio)

A new homoploid hybrid lineage needs to establish a degree of reproductive isolation from its parent species if it is to persist as an independent entity, but the role hybridization plays in this process is known in only a handful of cases. The homoploid hybrid ragwort species, Senecio squalidus, (Oxford ragwort) originated following the introduction of hybrid plants to the UK approximately 320 years ago. The source of the hybrid plants was from a naturally occurring hybrid zone between S. aethnensis and S. chrysanthemifolius on Mount Etna, Sicily. Previous studies of the parent species found evidence for multiple incompatibility loci causing transmission ratio distortion of genetic markers in their hybrid progeny. This study closes the hybridization triangle by reporting a genetic mapping analysis of the remaining two paired cross combinations between S. squalidus and its parents. Genetic maps produced from F2 mapping families were generally collinear but with half of the linkage groups showing evidence of genomic reorganization between genetic maps. The new maps produced from crosses between S. squalidus and each parent showed multiple incompatibility loci distributed across the genome, some of which co-locate with previously reported incompatibility loci between the parents. These findings suggest that this young homoploid hybrid species has inherited a unique combination of genomic rearrangements and incompatibilities from its parents that contribute to its reproductive isolation

Dissolved nitrous oxide (N2O) dynamics in agricultural field drains and headwater streams in an intensive arable catchment

Indirect nitrous oxide (N2O) emissions produced by nitrogen (N) leaching into surface water and groundwater bodies are poorly understood in comparison to direct N2O emissions from soils. In this study, dissolved N2O concentrations were measured weekly in both lowland headwater streams and subsurface agricultural field drain discharges over a two-year period (2013–2015) in an intensive arable catchment, Norfolk, UK. All field drain and stream water samples were found to have dissolved N2O concentrations higher than the water–air equilibrium concentration, illustrating that all sites were acting as a net source of N2O emissions to the atmosphere. Soil texture was found to significantly influence field drain N2O dynamics, with mean concentrations from drains in clay loam soils (5.3 µg N L-1) being greater than drains in sandy loam soils (4.0 µg N L-1). Soil texture also impacted upon the relationships between field drain N2O concentrations and other water quality parameters (pH, flow rate, and nitrate (NO3) and nitrite (NO2) concentrations), highlighting possible differences in N2O production mechanisms in different soil types. Catchment antecedent moisture conditions influenced the storm event mobilisation of N2O in both field drains and streams, with the greatest concentration increases recorded during precipitation events preceded by prolonged wet conditions. N2O concentrations also varied seasonally, with the lowest mean concentrations typically occurring during the summer months (JJA). Nitrogen fertiliser application rates and different soil inversion regimes were found to have no effect on dissolved N2O concentrations, whereas higher N2O concentrations recorded in field drains under a winter cover crop compared to fallow fields revealed cover crops are an ineffective greenhouse gas emission mitigation strategy. Overall, this study highlights the complex interactions governing the dynamics of dissolved N2O concentrations in field drains and headwater streams in a lowland intensive agricultural catchment

Mitigating river sediment enrichment through the construction of roadside wetlands

Metalled roads have been shown to act as a major pathway for land-to-river sediment transfer, but there currently exists limited research into mitigation solutions to tackle this pollution source. The aim of this study was to assess the effectiveness of three roadside constructed wetlands, installed in September 2016, at reducing sediment enrichment in a tributary of the River Wensum, UK. Two wetland designs were trialled (linear and ‘U-shaped’), both of which act as settling ponds to encourage entrained sediment to fall out of suspension and allow cleaner water to discharge into the river. Wetland efficiency was monitored through automated, high-resolution (30 min) turbidity probes installed upstream and downstream of the wetlands, providing a near-continuous record of river turbidity before (October 2011 – August 2016) and after (November 2016 – February 2018) installation. This was supplemented by lower resolution monitoring of the wetland inflows and outflows, as well as an assessment of sediment and nutrient accumulation rates within the linear wetland. Results revealed median river sediment concentrations decreased up to 14% after wetland construction and sediment load decreased by up to 82%, although this was largely driven by low river discharge post-installation. Median sediment concentrations discharging from the linear wetland (7.2 mg L-1) were higher than the U-shaped wetland (3.9 mg L-1), confirming that a longer flow pathway through wetlands can improve sediment retention efficiency. After 12 months of operation, the linear wetland had retained 7,253 kg (305 kg ha-1 y-1) of sediment, 11.6 kg (0.5 kg ha-1 y-1) of total phosphorus, 29.7 kg (1.3 kg ha-1 y-1) of total nitrogen and 400 kg (17 kg ha-1 y-1) of organic carbon. This translates into mitigated pollutant damage costs of £392 for sediment, £148 for phosphorus and £13 for nitrogen, thus giving a combined total mitigated damage cost of £553 y-1. With the linear wetland costing £3,411 to install and £145 – 182 y-1 to maintain, this roadside constructed wetland has an estimated payback time 8 years, making it a cost-effective pollution mitigation measure for tackling sediment-enriched road runoff that could be widely adopted at the catchment-scale

Indirect Nitrous Oxide Emission Factors for Agricultural Field Drains and Headwater Streams

Agriculture is a major source of nitrous oxide (N2O) emissions, a potent greenhouse gas. While direct N2O emissions from soils have been widely investigated, indirect N2O emissions from nitrogen (N) enriched surface water and groundwater bodies are poorly understood. In this contribution, indirect N2O emissions from subsurface agricultural field drains and headwater streams were monitored over a two-year period (2013–2015) in an intensive arable catchment in eastern England. Indirect N2O emission factors for groundwater (EF5g) and surface runoff (EF5r) were calculated for both field drain and streamwater samples, respectively, using two approaches: the N2O–N/NO3–N ratio and the IPCC (2006) methodology. Mean EF5g values derived from the N2O–N/NO3–N ratio were 0.0012 for field drains and 0.0003 for streamwater. Using the IPCC (2006) methodology, the mean EF5g values were 0.0011 for field drains and 0.0001 for streamwater. Thus, EF values derived from both methods were below the current IPCC (2006) default value of 0.0025 and a downward revision to 0.0012 for EF5g and 0.0002 for EF5r is recommended. Such revision would halve current estimates of N2O emissions associated with nitrogen leaching and runoff from agriculture for both the UK and globally

Diel turbidity cycles in a headwater stream: evidence of nocturnal bioturbation?

Purpose: A small number of recent studies have linked daily cycles in stream turbidity to nocturnal bioturbation by aquatic fauna, principally crayfish, and demonstrated this process can significantly impact upon water quality under baseflow conditions. Adding to this limited body of research, we use high-resolution water quality monitoring data to investigate evidence of diel turbidity cycles in a lowland, headwater stream with a known signal crayfish (Pacifastacus leniusculus) population and explore a range of potential causal mechanisms. Materials and methods: Automatic bankside monitoring stations measured turbidity and other water quality parameters at 30-min resolution at three locations on the River Blackwater, Norfolk, UK during 2013. Specifically, we focused on two 20-day periods of baseflow conditions during January and April 2013 which displayed turbidity trends typical of winter and spring seasons, respectively. The turbidity time-series, which were smoothed with 6.5 hour Savitzky-Golay filters to highlight diel trends, were correlated against temperature, stage, dissolved oxygen and pH to assess the importance of abiotic influences on turbidity. Turbidity was also calibrated against suspended particulate matter (SPM) over a wide range of values via linear regression. Results and discussion: Pronounced diel turbidity cycles were found at two of the three sites under baseflow conditions during April. Spring night-time turbidity values consistently peaked between 21:00 and 04:00 with values increasing by ~10 nephelometric turbidity units (NTU) compared with the lowest recorded daytime values which occurred between 10:00 and 14:00. This translated into statistically significant increases in median midnight SPM concentration of up to 76% compared with midday, with night-time (18:00 – 05:30) SPM loads also up to 30% higher than that recorded during the daytime (06:00 – 17:30). Relating turbidity to other water quality parameters exhibiting diel cycles revealed there to be neither any correlation that might indicate a causal link, nor any obvious mechanistic connections to explain the temporal turbidity trends. Diel turbidity cycles were less prominent at all sites during the winter. Conclusions: Considering the seasonality and timing of elevated turbidity, visual observations of crayfish activity, and an absence of mechanistic connections with other water quality parameters, the results presented here are consistent with the hypothesis that nocturnal bioturbation is responsible for generating diel turbidity cycles under baseflow conditions in headwater streams. However, further research in a variety of fluvial environments is required to better assess the spatial extent, importance and causal mechanisms of this phenomenon

Assessing the farm-scale impacts of cover crops and non-inversion tillage regimes on nutrient losses from an arable catchment

The efficacy of cover crops and non-inversion tillage regimes at minimising farm-scale nutrient losses were assessed across a large, commercial arable farm in Norfolk, UK. The trial area, covering 143 ha, was split into three blocks: winter fallow with mouldboard ploughing (Block J); shallow non-inversion tillage with a winter oilseed radish (Raphanus sativus) cover crop (Block P); and direct drilling with a winter oilseed radish cover crop (Block L). Soil, water and vegetation chemistry across the trial area were monitored over the 2012/13 (pre-trial), 2013/14 (cover crops and non-inversion tillage) and 2014/15 (non-inversion tillage only) farm years. Results revealed oilseed radish reduced nitrate (NO3¬¬-N) leaching losses in soil water by 75–97% relative to the fallow block, but had no impact upon phosphorus (P) losses. Corresponding reductions in riverine NO3¬¬-N concentrations were not observed, despite the trial area covering 20% of the catchment. Mean soil NO3¬¬-N concentrations were reduced by ~77% at 60–90 cm depth beneath the cover crop, highlighting the ability of deep rooting oilseed radish to scavenge nutrients from deep within the soil profile. Alone, direct drilling and shallow non-inversion tillage were ineffective at reducing soil water NO3¬¬-N and P concentrations relative to conventional ploughing. Applying starter fertiliser to the cover crop increased radish biomass and nitrogen (N) uptake, but resulted in net N accumulation within the soil. There was negligible difference between the gross margins of direct drilling (£731 ha-1) and shallow non-inversion tillage (£758 ha-1) with a cover crop and conventional ploughing with fallow (£745 ha-1), demonstrating farm productivity can be maintained whilst mitigating diffuse pollution. The results presented here support the wider adoption of winter oilseed radish cover crops to reduce NO3¬¬-N leaching losses in arable systems, but caution that it may take several years before catchment-scale impacts downstream are detected

Application of high-resolution telemetered sensor technology to develop conceptual models of catchment hydrogeological processes

Mitigating agricultural water pollution requires changes in land management practices and the implementation of on-farm measures to tackle the principal reasons for water quality failure. However, a paucity of robust empirical evidence on the hydrological functioning of river catchments can be a major constraint on the design of effective pollution mitigation strategies at the catchment-scale. In this regard, in 2010 the UK government established the Demonstration Test Catchment (DTC) initiative to evaluate the extent to which on-farm mitigation measures can cost-effectively reduce the impacts of agricultural water pollution on river ecology while maintaining food production capacity. A central component of the DTC platform has been the establishment of a comprehensive network of automated, web-based sensor technologies to generate high-temporal resolution empirical datasets of surface water, soil water, groundwater and meteorological parameters. In this paper, we demonstrate how this high-resolution telemetry can be used to improve our understanding of hydrological functioning and the dynamics of pollutant mobilisation and transport under a range of hydrometerological and hydrogeological conditions. Furthermore, we demonstrate how these data can be used to develop conceptual models of catchment hydrogeological processes and consider the implications of variable hydrological functioning on the performance of land management changes aimed at reducing agricultural water pollution

Hydrochemical and dual-isotope approach to the identification of denitrification in arable field drainage in the Wensum catchment, eastern England

The global pool of reactive nitrogen has doubled in the last century in response to the need to increase food production with the consequent increase in fertiliser-derived reactive nitrogen detrimentally affecting aquatic ecosystems. This study investigates the spatial distribution and significance of denitrification in the lowland, agriculturally-impacted River Wensum catchment in eastern England as a natural attenuation process. To investigate the evidence for denitrification, the hydrochemical characteristics and dual stable isotope composition of nitrate (15N and 18O) were measured over a 15-month period, 2015–2017, in 63 samples of field drainage in predominantly clay loam and sandy clay loam soils under mainly arable cultivation. Microbially-mediated denitrification in field drainage was indicated by the gradient of the linear regression of 15NNO3 and 18ONO3 compositions with a value of 0.58. Dual fractionation of the nitrate isotopes yielded enrichment factors for δ15NNO3 (−4.52‰) and δ18ONO3, (−4.51‰) within the reported ranges for denitrification in aquatic studies. Soil type influenced denitrification, with a positive relationship between percentage clay and δ15NNO3 and δ18ONO3 values. The same relationship was observed for denitrification rates calculated via a simple mass balance approach, which ranged from 11.0 to 26.3 kg N ha−1 and accounted for 30–73% of the leached soil nitrogen. Higher denitrification rates were recorded in drainage areas with a greater soil clay content (>20% by weight). Comparing calculated dentification rates for individual drain areas with median δ15NNO3 values of drain samples demonstrated that an isotopic enrichment of +1‰ is associated with a denitrification rate of 2.6 kg N ha−1. In conclusion, sustainable agricultural practices that maintain natural attenuation processes such as denitrification, for example by preserving and increasing the soil organic carbon content, are desirable to improve overall soil health to support ecosystem services that reduce nitrate pollution

Temporal hydrochemical dynamics of the River Wensum, UK: Observations from long-term high-resolution monitoring (2011–2018)

In 2010, the UK government established the Demonstration Test Catchment (DTC) initiative to evaluate the extent to which on-farm mitigation measures can cost-effectively reduce the impacts of agricultural water pollution on river ecology whilst maintaining food production capacity. A central component of the DTC platform was the establishment of a comprehensive network of automated, web-based sensor technologies to generate high-temporal resolution (30 min) empirical datasets of surface water, groundwater and meteorological parameters over a long period (2011–2018). Utilising 8.9 million water quality measurements generated for the River Wensum, this paper demonstrates how long-term, high-resolution monitoring of hydrochemistry can improve our understanding of the complex temporal dynamics of riverine processes from 30 min to annual timescales. This paper explores the impact of groundwater-surface water interactions on instream pollutant concentrations (principally nitrogen, phosphorus and turbidity) and reveals how varying hydrochemical associations under contrasting flow regimes can elicit important information on the dominant pollution pathways. Furthermore, this paper examines the relationships between agricultural pollutants and precipitation events of varying magnitude, whilst demonstrating how high-resolution data can be utilised to develop conceptual models of hydrochemical processes for contrasting winter and summer seasons. Finally, this paper considers how high-resolution hydrochemical data can be used to increase land manager awareness of environmentally damaging farming operations and encourage the adoption of more water sensitive land management practices

Sensitivity of fluvial sediment source apportionment to mixing model assumptions: A Bayesian model comparison

Mixing models have become increasingly common tools for apportioning fluvial sediment load to various sediment sources across catchments using a wide variety of Bayesian and frequentist modeling approaches. In this study, we demonstrate how different model setups can impact upon resulting source apportionment estimates in a Bayesian framework via a one-factor-at-a-time (OFAT) sensitivity analysis. We formulate 13 versions of a mixing model, each with different error assumptions and model structural choices, and apply them to sediment geochemistry data from the River Blackwater, Norfolk, UK, to apportion suspended particulate matter (SPM) contributions from three sources (arable topsoils, road verges, and subsurface material) under base flow conditions between August 2012 and August 2013. Whilst all 13 models estimate subsurface sources to be the largest contributor of SPM (median ∼76%), comparison of apportionment estimates reveal varying degrees of sensitivity to changing priors, inclusion of covariance terms, incorporation of time-variant distributions, and methods of proportion characterization. We also demonstrate differences in apportionment results between a full and an empirical Bayesian setup, and between a Bayesian and a frequentist optimization approach. This OFAT sensitivity analysis reveals that mixing model structural choices and error assumptions can significantly impact upon sediment source apportionment results, with estimated median contributions in this study varying by up to 21% between model versions. Users of mixing models are therefore strongly advised to carefully consider and justify their choice of model structure prior to conducting sediment source apportionment investigations