Winter Population Research Articles

Hanöbukten, an important wintering area for waterbirds in South Sweden

Hanöbukten in SE Sweden (including Blekinge archipelago) is an important wintering and staging area for a number of waterbird species. The inshore parts have been covered by counts from the shore during the International Waterbird Counts (IWC) since 1969. In the present study we analyze these counts for the period 1969–2014. When the counts started the total number of wintering waterbirds varied between 20,000 and 40,000 per year. However, from the early 1990s the number doubled, reaching 60,000 to 80,000 birds per year. All species with the exception of the Long-tailed Duck Clangula hyemalis showed similar increase. The area had a high proportion of the Swedish wintering population of Pochard Aythya ferina and Smew Mergellus albellus. Large numbers of Tufted Duck Aythya fuligula also wintered in the area, and some parts of the archipelago had internationally important concentrations of Tufted Duck and Smew. The increase in numbers in the study area showed the same pattern as in all Sweden and was most probably a result of milder winters in recent years.

Surviving winter: Food, but not habitat structure, prevents crashes in cyclic vole populations.

Vole population cycles are a major force driving boreal ecosystem dynamics in northwestern Eurasia. However, our understanding of the impact of winter on these cycles is increasingly uncertain, especially because climate change is affecting snow predictability, quality, and abundance. We examined the role of winter weather and snow conditions, the lack of suitable habitat structure during freeze‐thaw periods, and the lack of sufficient food as potential causes for winter population crashes. We live‐trapped bank voles Myodes glareolus on 26 plots (0.36 ha each) at two different elevations (representing different winter conditions) in southeast Norway in the winters 2013/2014 and 2014/2015. We carried out two manipulations: supplementing six plots with food to eliminate food limitation and six plots with straw to improve habitat structure and limit the effect of icing in the subnivean space. In the first winter, all bank voles survived well on all plots, whereas in the second winter voles on almost all plots went extinct except for those receiving supplemental food. Survival was highest on the feeding treatment in both winters, whereas improving habitat structure had no effect. We conclude that food limitation was a key factor in causing winter population crashes.

Validation of quantitative trait loci for grain quality-related traits in a winter wheat mapping population

Validating positions and effects of putative quantitative trait loci (QTL) is an important step before employing linked markers in marker-assisted selection (MAS) or conducting finer-scale mapping. Previously, QTL for grain quality traits were identified in a population of 185 doubled haploid (DH) winter wheat (Triticum aestivum L.) lines developed from the cross CO940610/’Platte’. Clusters of QTL for grain quality traits were detected on chromosomes 1B, 6B, and 7B. Our objectives were to (1) confirm the previously identified QTL, using recombinant inbred line (RIL, n = 186) and BC3F2:3 (n = 35) populations developed from the same parents; and (2) test the effects of combinations of alleles at three marker loci in the target chromosome regions. Field trials were conducted in two Colorado environments for the RIL population in 2009/10 and three environments for the BC3F2:3 population in 2012/13. Most QTL previously detected for grain quality traits on 1B, 6B, and 7B were repeated in either the RIL or BC3F2:3 populations, although these QTL were not robust across populations and environments. However, the QTL for grain ash concentration (Gac) on 7B was detected in all environments and populations, with Platte contributing the higher value allele. The BC3F2:3 lines with the allelic combination predicted to have higher grain protein concentration (Gpc) surpassed the lowest predicted combination in one of three environments investigated. Based on these evaluations, marker Bx7-MAR on 1B and the regions around markers Xwmc182a on 6B and Xwmc182b on 7B are recommended for further research for improving grain quality traits.

Ornithological Fauna of the Waste Water Treatment Plants in the Northern Left Bank Ukraine (Chernihiv and Kyiv Regions): Winter Populations and Ecological Structure

Abstract The article discusses winter bird populations of the waste water treatment plants (WWTP) located in the North of Left -bank Ukraine. The said population comprises 12 orders and 29 families. The most numerous are Passeriformes (37 species), Аnsеriformes (16 species) and Falconiformes (6 species). Parus major was registered at all types of facilities while most of the others house Passer montanus, Carduelis carduelis, Turdus pilaris, and Parus caeruleus. The largest number of wintering birds was registered at Bortnychi aeration station, Chernihiv municipal WWTP and Chernihiv wool processing factory - 79. 51 and 15 species respectively. The nuclear part of the bird numbers are the species residing at the facilities all year around (65.8 %); species occurring there in winter only account for 34.2 %. Dendrophilous (38 species) and hydrophilous (35 species) dominate among them. The primary role in forming the winter fauna of the waste water treatment plants belongs to the zones of water bodies and dams.

The phylogenetic relationship and cross-infection of nucleopolyhedroviruses between the invasive winter moth (Operophtera brumata) and its native congener, Bruce spanworm (O. bruceata)

Disease can affect biological invasions by acting as either a synergist or antagonist. Disease-mediated invasions have important implications for understanding the spread of invasive insects, which cost billions of dollars in damages annually. One such non-native, destructive insect is the winter moth, Operophtera brumata L. (Lepidoptera: Geometridae), which causes defoliation and mortality of deciduous trees in its introduced range. In the northeastern United States, winter moth populations overlap with a native congener, Bruce spanworm, Operophtera bruceata Hulst. Nucleopolyhedrovirus (NPV), appears to be an important natural enemy in Bruce spanworm and there is some evidence that the NPV infection found in winter moth in the northeastern U.S. may originate from Bruce spanworm. By sequencing two viral genes (the polyhedrin and p74 genes) from field-collected larvae of both species, we found that the winter moth virus (OpbuNPV) is distinct from the virus from Bruce spanworm (OpbrNPV). However, the two viruses do constitute a clade within the Alphabaculovirus Group 2 NPVs, indicating that they are more similar to each other than they are to other lepidopteran viruses, even other geometrid-derived NPVs. As far as we know, this is the first report of sequences from an NPV from Bruce spanworm. Results from cross infection trials suggest that cross infection is uncommon if it occurs at all. Our results show that these two closely related species have distinct viruses and, unlike previous suggestions, Bruce spanworm virus is not mediating the winter moth invasion.

The Influence of Environmental Factors on the Distribution of Wintering Populations of Asian Houbara Bustard Chlamydotis Macqueenii in Central Iranian Plateau

The Influence of Environmental Factors on the Distribution of Wintering Populations of Asian Houbara Bustard Chlamydotis Macqueenii in Central Iranian Plateau

Asymmetric hybridization between non-native winter moth, Operophtera brumata (Lepidoptera: Geometridae), and native Bruce spanworm, Operophtera bruceata, in the Northeastern United States, assessed with novel microsatellites and SNPs.

The European winter moth, Operophtera brumata, is a non-native pest in the Northeastern USA causing defoliation of forest trees and crops such as apples and blueberries. This species is known to hybridize with O. bruceata, the Bruce spanworm, a native species across North America, although it is not known if there are hybrid generations beyond F1. To study winter moth population genetics and hybridization with Bruce spanworm, we developed two sets of genetic markers, single nucleotide polymorphisms (SNPs) and microsatellites, using genomic approaches. Both types of markers were validated using samples from the two species and their hybrids. We identified 1216 SNPs and 24 variable microsatellite loci. From them we developed a subset of 95 species-diagnostic SNPs and ten microsatellite loci that could be used for hybrid identification. We further validated the ten microsatellite loci by screening field collected samples of both species and putative hybrids. In addition to confirming the presence of F1 hybrids reported in previous studies, we found evidence for multi-generation asymmetric hybridization, as suggested by the occurrence of hybrid backcrosses with the winter month, but not with the Bruce spanworm. Laboratory crosses between winter moth females and Bruce spanworm males resulted in a higher proportion of viable eggs than the reciprocal cross, supporting this pattern. We discuss the possible roles of population demographics, sex chromosome genetic incompatibility, and bacterial symbionts as causes of this asymmetrical hybridization and the utility of the developed markers for future studies.

Topography of the Andes Mountains shapes the wintering distribution of a migratory bird

AbstractAimIn birds, patterns of migratory connectivity are influenced by landscape structure experienced throughout the annual cycle. Assessing how landscape patterns drive the movements and distributions of migratory species is important for understanding the evolution of migration and determining conservation priorities. We determined migratory connectivity in the Canada Warbler (Cardellina canadensis), a declining Neotropical migratory bird, and evaluated the degree to which the Andean mountains influence winter population distribution in this montane forest specialist.LocationSeven study regions across the Western, Central, and Eastern Andes of Colombia.MethodsWe inferred breeding and natal origins of Canada Warblers overwintering in Colombia by analysing stable hydrogen isotope values (δ2H) in feathers and assigning wintering populations to breeding origins using an established feather δ2H isoscape for North America.ResultsWe found strong evidence for parallel migration between breeding and wintering sites, giving rise to moderately strong migratory connectivity between breeding and wintering regions. The model best explaining population distribution in the Colombian Andes implied that populations were most similar within basins or valleys. Such segregation is notable, given that basins are separated by less than 300 km on the wintering grounds, while breeding populations may be separated by more than 3000 km from east to west.Main conclusionsOur results suggest that the high spine of each Andean cordillera acts to shape the winter distribution of individuals from different breeding populations. To our knowledge, this is the first demonstration that the topography of the Andean mountains can act to shape the winter distribution of populations of a migratory bird. The strength of migratory connectivity found in Canada Warblers may inhibit evolutionary responses to environmental change, and the more rapid population declines recorded in some eastern breeding populations may be related to habitat loss and local environmental conditions in the Eastern Andes, where individuals from these populations were more likely to overwinter.

Collective Vigilance in the Wintering Hooded Crane: The Role of Flock Size and Anthropogenic Disturbances in a Human‐Dominated Landscape

AbstractVigilance achieved at the group level, known as collective vigilance, can enhance the ability to assess threats and confer benefits to gregarious prey species. Examining the factors that influence collective vigilance and exploring how individual vigilance is organized at the group level can help to understand how prey groups respond to potential threats. We quantified collective vigilance and determined its temporal pattern in a natural wintering population of the hooded craneGrus monachain the Shengjin Lake reserve in China. We examined the role of flock size and anthropogenic disturbances in the human‐dominated landscape on collective vigilance and level of synchronization. The proportion of time during which at least one individual scanned the surroundings (collective vigilance) increased with flock size and was higher in the more disturbed buffer zone of the lake. Synchronization of vigilance occurred more frequently in the smaller flocks but was not related to the risk of disturbance. Synchronization implies that individuals tend to monitor and copy the vigilance of neighbors. In the degraded wetlands, the wintering hooded crane benefits from foraging in groups and synchronizing their vigilance in response to human disturbances.

Climate constrains range expansion of an allochronic population of the pine processionary moth

AbstractAimAllochronic speciation occurs when sympatric populations sharing similar feeding regimes diverge because they mate at different times. Such speciation mode is fascinating because it permits the study of subsequent adaptations and/or geographic range shifts undergone by the populations that face new ecological constraints. Moreover, exploring whether intraspecific differentiation is accompanied by niche divergence is crucial for planning efficient biodiversity management and invasive species control. Here, we address the topic of potential climatic niche divergence between two sympatric populations of the pine processionary moth Thaumetopoea pityocampa (PPM) that diverged under allochrony. The larval development of a PPM population recently discovered in coastal regions of Portugal, referred to as the summer population (SP), occurs in the spring–summer instead of the autumn–winter as for all other populations of this species.LocationPortugal.MethodsWe carried out intensive field sampling in the area where both populations are sympatric and calibrated spread (MigClim approach) and species distribution models (ensemble forecasting approach) to depict the response of SP to environment and predict its potential range under current and future climatic conditions.ResultsSince its discovery in 1997, the distribution of SP has been expanding along the Western coasts of Portugal. Still, its establishment in inland regions failed, even though suitable hosts occur. Our models identify maximal temperatures as the main constraint explaining SP absence from inland regions. The coastal area where winter populations and SP co‐occur displays unique climatic conditions (moderate maximal and minimal temperatures), which enable the coexistence of these populations with totally different phenologies. The models predict a future decrease of SP range due to climate warming.Main conclusionsWe show here that a climatic niche shift occurred following allochronic divergence. This study highlights how climate differentially affects allochronic populations and how considering intraspecific diversity is crucial when predicting species responses to climate change.

Willet be one species or two? A genomic view of the evolutionary history ofTringa semipalmata

The Willet (Tringa semipalmata; Scolopacidae) is composed of 2 morphologically and vocally distinct subspecies with broadly disjunct breeding distributions in North America. Nominate T. s. semipalmata breeds in coastal salt and brackish marshes along the Atlantic and Gulf coasts of eastern North America and the West Indies, and T. s. inornata breeds in wet grasslands and prairies in the northwestern interior of North America. To assess divergence and test for hybridization between the 2 subspecies, we sampled breeding and wintering populations and collected morphological data, mitochondrial DNA sequences from the ND2 locus, and nuclear DNA sequences from genomic libraries enriched for ultraconserved elements (UCEs). Mitochondrial haplotypes were reciprocally monophyletic between the 2 subspecies and indicated divergence approximately 700,000 yr ago. The UCE dataset included 4,635 loci containing 19,322 single nucleotide polymorphisms (SNPs), and, based on these data, individuals clustered by subspecies with no evidence of admixture between them and no substructure within subspecies. We identified 42 nuclear loci that contained SNPs fixed for alternate alleles between the 2 subspecies. Of the 42 loci with fixed differences, a statistically disproportional 17 were Z-linked, indicating a role for sexual selection in the divergence of the 2 subspecies. Genetic, morphological, ecological, and behavioral differences suggest that the 2 Willet subspecies may merit treatment as separate species. Further studies are needed to determine the presence of pre- or post-mating reproductive isolation.

Changes in numbers and distribution of wintering Long-tailed Ducks Clangula hyemalis in Swedish waters during the last fifty years

The Long-tailed Duck Clangula hyemalis is the most common wintering waterfowl in Swedish marine waters but it is not well covered in the International Midwinter Counts due to its offshore distribution making special surveys necessary for the species. Surveys covering the entire Baltic Sea were organized for the first time in 1992/1993 with follow-up surveys in 2007–2011 and 2016. In Sweden, aerial line transect surveys covered all areas of importance for the Long-tailed Duck during the last two all-Baltic surveys. In the 1970s extensive regional surveys were done from coastguard boats and airplane. The total wintering population of Long-tailed Duck was estimated to be around 370,000 individuals in 2016 compared to 436,000 in 2009, and in the order of 1,400,000 in 1992/1993, indicating an ongoing decrease. Comparing the more recent surveys with the surveys from the 1970s, the most marked changes in wintering numbers were found east of Gotland, where only a small proportion of the numbers estimated in the 1970s remained. Decreases were also found in the waters around Öland, whereas no marked changes were noted for the smaller wintering populations of the east coast archipelagos.

Unraveling migratory connectivity of two European diving ducks: a stable isotope approach

Quantifying the strength of migratory connectivity or the degree to which breeding and wintering populations are connected is crucial for conservation and management of migratory birds. Using stable hydrogen isotope ratios of secondary flight feathers (δ 2Hf), we assessed the breeding origins/locations of 666 common pochards (Aythya ferina) and 190 tufted ducks (A. fuligula) captured in France during the 2005–2006 and 2008–2009 wintering seasons. Isotopic values of common pochards wintering on Grand-lieu Lake (western France) displayed a marked bimodal spatial distribution supportive of strong contributions of both northern (60–65° of latitude) and central European (45–55° of latitude) origins. According to the temporal distribution of δ 2Hf values, the presence of common pochards of northern origin reached a maximum at the peak of the wintering season (December–January). In contrast, tufted duck did not display a clear bimodal pattern in δ 2Hf, with individuals displaying a more northerly origin (60–70° of latitude), likely of Russian and Scandinavian origin. First-year individuals had more northern origins than adults, suggesting that either isotopic discrimination differed among age groups or, more probably, that some adults may undergo moult at latitudes lower than that of their breeding ground. Tufted ducks with northerly δ 2Hf values were observed early in the season in individuals sampled in eastern France and late in the season in those sampled in western France. The effects of winter severity on the distribution of individuals over the wintering range were supported for the tufted duck with more northerly values recorded during harsh winters. Management implications of these results are discussed.

Large‐scale climatic drivers of regional winter bird population trends

AbstractAimChanges in climate and land use practices have been found to affect animal populations in different parts of the world. These studies have typically been conducted during the breeding season, whereas the non‐breeding season (hereafter ‘winter’) has received much less attention. Changes in regional winter abundances could be caused by changes in overall population sizes and/or redistribution of populations. We tested these mechanisms for terrestrial winter bird population changes in Northern Europe and explored the role of climate change and species habitat preference.LocationThe Netherlands, Denmark, Sweden, Finland.MethodsWe used winter bird counts from four countries conducted annually between 15 December and 20 January in 1980/1981–2013/2014. We report national population trends for 50 species for which a trend could be calculated in at least three of the countries. We analysed country‐specific population growth rates in relation to species’ climatic summer and winter niches, habitat preference and migratory behaviour.ResultsSpecies breeding in colder (typically northern) areas showed more negative winter population trends than species breeding in warmer areas. Regional winter population trends were negatively correlated with characteristics of their winter climatic niche: populations in the colder part of their winter distribution increased in abundance, whereas populations in the warmer part of their winter distribution decreased. Woodland species tended to do better than farmland species. Migratory behaviour did not explain variation in population trends.Main conclusionsThe generally decreasing winter population trends of cold‐dwelling breeding species probably reflect the general decline in population sizes of these species. In contrast, increasing winter population trends for populations in the colder parts of the winter distribution indicate a redistribution of wintering individuals towards the north‐east. Both these patterns are likely caused by climate change.

Predation of wood mice (Apodemus sylvaticus) on hibernating bats

AbstractBat hibernacula with high numbers of bats can become high‐risk areas, as they attract flying and non‐flying predators. In order to protect hibernating bats effectively, more knowledge about mortality factors is needed. During the winters of 2003–2015, we found 214 dead bats in 12 hibernacula in The Netherlands province of Zuid‐Holland. Most bat remains were found in December and January, with a second peak in April. Their remains showed a typical pattern of lesions consistent with those caused by predation by the wood mouse (Apodemus sylvaticus). Trail camera surveys showed that wood mice actively searched for bats. Predation pressure seemed to vary between winters, with a peak in the winters of 2004, 2011 and 2015. The annual mortality (relative to the maximum winter population size) caused by wood mouse predation varied between 0.1 and 8.8 %, with a maximum local effect of 83.6 %. The years with high wood mouse predation pressure were characterized by a long frost period and a low mast production of common oak in the preceding autumn. The size of a hibernaculum and the population density of its bats had an effect on predation‐dependent mortality. The highest predation risk occurred near the entrance of bunkers. From these results we tentatively conclude that predation is not incidental and that wood mice actively search for and kill hibernating bats or scavenge for weakened individuals.

Habitat Use and its Implications for the Conservation of the Overwintering Populations of Bean GooseAnser fabalisand Greater White-Fronted GooseA. albifronsin South Korea

Abstract The Korean peninsula is geographically important for the migration of geese. Several thousand geese visit South Korea every winter, but currently it is unclear which habitats are preferred by the geese. Understanding patterns of habitat usage is important for establishing conservation strategies, as some habitats may be disappearing more rapidly than others making it critical to know whether or not a disappearing habitat is crucial to certain birds. In this study, we investigated the habitat use of the wintering populations of Bean Goose Anser fabalis and Greater White-fronted Goose A. albifrons. We found that the wintering populations of these two geese species in South Korea have stabilized after sudden increases during the mid and late 2000s. Both species formed larger wintering populations at coastal lakes, on reclaimed lands, and at estuaries than on freshwater habitats, which may be related to the availability and/or diversity of food items. Considering that environmental changes resulting ...

Why fly the extra mile? Using stress biomarkers to assess wintering habitat quality in migratory shorebirds.

Migratory birds make decisions about how far to travel based on cost-benefit trade-offs. However, in many cases the net effect of these trade-offs is unclear. We sought to address this question by measuring feather corticosterone (CORTf), leucocyte profile, avian malaria parasite prevalence and estimating fueling rates in three spatially segregated wintering populations of the migratory shorebird ruddy turnstone Arenaria interpres during their stay in the winter habitat. These birds fly from the high-Arctic breeding ground to Australia, but differ in that some decide to end their migration early (Broome, Western Australia), whereas others travel further to either South Australia or Tasmania. We hypothesized that the extra costs in birds migrating greater distances and overwintering in colder climates would be offset by benefits when reaching their destination. This would be evidenced by lower stress biomarkers in populations that travel further, owing to the expected benefits of greater resources and improved vitality. We show that avian malaria prevalence and physiological stress levels were lower in birds flying to South Australia and Tasmania than those overwintering in Broome. Furthermore, our modeling predicts that birds in the southernmost locations enjoy higher fueling rates. Our data are consistent with the interpretation that birds occupying more costly wintering locations in terms of higher migratory flight and thermoregulatory costs are compensated by better feeding conditions and lower blood parasite infections, which facilitates timely and speedy migration back to the breeding ground. These data contribute to our understanding of cost-benefit trade-offs in the decision making underlying migratory behaviour.

Evidence for low-level hybridization between two allochronic populations of the pine processionary moth,Thaumetopoea pityocampa(Lepidoptera: Notodontidae)

Divergence between populations sharing the same habitat can be initiated by different reproductive times, leading to allochronic differentiation. A spatially localized allochronic summer population (SP) of the pine processionary moth Thaumetopoea pityocampa, recently discovered in Portugal, occurs in sympatry with the local winter population (WP). We examined the level of genetic differentiation between the two populations and estimated the current gene flow within the spatial framework of their co-occurrence. Mitochondrial data indicated that the two sympatric populations were genetically closer than other WP populations. Conversely, microsatellite genotyping uncovered greater differentiation between the two sympatric populations than between allopatric ones. While male trapping confirmed that reproduction of SP and WP occurred at distinct times, clustering approaches demonstrated the presence of a few LateSP individuals emerging within the WP flight period, although genetically identified as SP. We also identified rare recent hybridization events apparently occurring mainly in the margins of the current SP range. The ongoing gene flow detected between the ancestral and the emerging allochronic populations revealed an incomplete reproductive isolation, which must therefore be taken into account and integrated with studies focussed on ecological drivers, so that a complete understanding of the ongoing speciation process might be achieved.

Seasonal variation in the diet of Antarctic fur seals, Arctocephalus gazella, at 25 de Mayo/King George Island, South Shetland Islands, Antarctica

Assessing the trophic ecology of fur seals is essential for defining their roles as a top predator in Antarctic marine ecosystems. To this end, we examined the diet of Antarctic fur seals, Arctocephalus gazella, in the South Shetlands. While previous studies have considered the diet of this species at this location in summer, to the best of our knowledge, this is the first study to report on the diet of this population in winter, thus adding essential information on the extent of seasonal variation. A total of 224 scats were collected in the winter 2004 (n = 118) and summer 2004/2005 (n = 106) on the coasts of Stranger Point, Isla 25 de Mayo/King George. For the total study period krill (Euphausia superba) was the main prey taxon, followed by fish, cephalopods and penguins. During winter Pleuragramma antarcticum was dominant, while myctophids were almost absent. Conversely, in summer Electrona antarctica and Gymnoscopelus nicholsi, followed by P. antarcticum, dominated the fish diet. It is concluded that fur seals centred their foraging activity on a krill community and fish associated with krill aggregations.

Effect of the Gpc‐B1 Allele in Hard Red Winter Wheat in the US Pacific Northwest

Grain protein concentration (GPC) influences end‐use quality of products made from wheat (Triticum aestivum L.) and is therefore a high priority for breeders. Most variation in GPC is due to environmental rather than genetic effects. Because of the negative correlation between yield and GPC, enhancing genetic expression is important for improving GPC in hard wheat varieties. The objective of this study was to evaluate the Gpc‐B1 allele in hard winter wheat cultivars. Near‐isogenic lines (NILs) with and without the Gpc‐B1 allele were created in four populations: ‘Farnum’ hard red winter wheat (HRW) × ‘Lassik’ hard red spring wheat (HRS), WA8061 (HRW; advanced breeding line) × Lassik, Farnum × ‘Hollis’ HRS, and Farnum by ‘Eddy’ HRW. Presence of this allele was validated using diagnostic marker Xuhw89. Results from field trials planted in Pullman and Lind, WA, indicate that inclusion of the Gpc‐B1 allele in HRW increases GPC. A negative correlation with grain yield and GPC was found, showing inclusion of the Gpc‐B1 allele significantly decreased grain yield potential. An exception was the Farnum × Hollis NIL with the Gpc‐B1 allele that showed higher GPC and grain yield potential than the NIL without this allele. End‐use quality of lines was also affected with the inclusion of the Gpc‐B1 allele, with average decreases in milling characteristics and increases in certain baking properties. These data demonstrate that in winter wheat breeding populations, lines can be selected with the Gpc‐B1 allele, resulting in increased GPC and acceptable grain yield and end‐use quality profiles.