Beyond the single index: Investigating ecological mechanisms underpinning ecosystem multifunctionality with network analysis

Ecosystems simultaneously deliver multiple functions that relate to both the activities of resident species and environmental conditions. One of the biggest challenges in multifunctionality assessment is balancing analytical simplicity with ecosystem complexity. As an alternative to index‐based approaches, we introduce a multivariate network analysis that uses network theory to assess multifunctionality in terms of the relationships between species'' functional traits, environmental characteristics, and functions. We tested our approach in a complex and heterogeneous ecosystem, marine intertidal sandflats. We considered eight ecosystem function, five macrofaunal functional trait groups derived from 36 species, and four environmental characteristics. The indicators of ecosystem functions included the standing stock of primary producers, oxygen production, benthic oxygen consumption, DIN (ammonium and NOx efflux) and phosphate release from the sediments, denitrification, and organic matter degradation at the sediment surface. Trait clusters included functional groups of species that shared combinations of biological traits that affect ecosystem function: small mobile top 2 cm dwellers, suspension feeders, deep‐dwelling worms, hard‐bodied surface dwellers, and tube‐forming worms. Environmental characteristics included sediment organic matter, %mud, %shell hash, and %sediment water content. Our results visualize and quantify how multiple ecosystem elements are connected and contribute to the provision of functions. Small mobile top 2 cm dwellers (among trait clusters) and %mud (among environmental characteristics) were the best predictor for multiple functions. Detailed knowledge of multifunctionality relationships can significantly increase our understanding of the real‐world complexity of natural ecosystems. Multivariate network analysis, as a standalone method or applied alongside already existing single index multifunctionality methods, provides means to advance our understanding of how environmental change and biodiversity loss can influence ecosystem performance across multiple dimensions of functionality. Embedding such a detailed yet holistic multifunctionality assessment in environmental decision‐making will support the assessment of multiple ecosystem services and social‐ecological values.     

Landscape variables influence taxonomic and trait composition of insect assemblages in Neotropical savanna streams

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Trait structure and redundancy determine sensitivity to disturbance in marine fish communities

Trait diversity is believed to influence ecosystem dynamics through links between organismal traits and ecosystem processes. Theory predicts that key traits and high trait redundancy—large species richness and abundance supporting the same traits—can buffer communities against environmental disturbances. While experiments and data from simple ecological systems lend support, large‐scale evidence from diverse, natural systems under major disturbance is lacking. Here, using long‐term data from both temperate (English Channel) and tropical (Seychelles Islands) fishes, we show that sensitivity to disturbance depends on communities’ initial trait structure and initial trait redundancy. In both ecosystems, we found that increasing dominance by climatically vulnerable traits (e.g., small, fast‐growing pelagics/corallivores) rendered fish communities more sensitive to environmental change, while communities with higher trait redundancy were more resistant. To our knowledge, this is the first study demonstrating the influence of trait structure and redundancy on community sensitivity over large temporal and spatial scales in natural systems. Our results exemplify a consistent link between biological structure and community sensitivity that may be transferable across ecosystems and taxa and could help anticipate future disturbance impacts on biodiversity and ecosystem functioning.     

Multi‐scale habitat modification by coexisting ecosystem engineers drives spatial separation of macrobenthic functional groups

By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large‐scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co‐occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co‐occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. This understanding can profitably be used to improve ecosystem‐based management and conservation actions.     

Functional diversity of marine macrobenthic communities from sublittoral soft-sediment habitats off northern Chile

Benthic communities show changes in composition and structure across different environmental characteristics and habitats. However, incorporating species biological traits into the analysis can provide a better understanding of system functioning within habitats. We compare the functional diversity of macrobenthic communities from a contrasting shallow (15 m) and deep (50 m) sublittoral soft-sediment habitats in northern Chile, using biological traits analysis. Our aim was to highlight the biological characteristics responsible for differences between habitats and the implications for ecosystem functioning. Trait analysis showed that the deep habitat was restricted in providing functionally important biogenic structure and bioturbation and supports less diverse feeding-related energy pathways. The shallow habitat is characterized by more diverse energy pathways and a higher potential for matter exchange through bioturbation. We provide support to the predictions of transfer of energy from the benthos to upper trophic levels in the shallow, which is characterized mainly by normoxia and little organic matter content in the sediment. In the deep habitat, characterized by hypoxia and more organic matter, energy appears to be transferred to microbial components. We suggest that trait analysis should be added to the traditional approaches based on species diversity, because it provides indicators of ecosystem stress.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

A biological trait approach to assess the functional composition of subtidal benthic communities in an estuarine ecosystem

Within transitional/estuarine environments ‘ecosystem functioning’ has been mostly investigated with “traditional” taxonomic analysis, based on the taxonomic composition of benthic invertebrate communities. However, ‘ecosystem functioning’ depends also greatly on the functional characteristics (biological traits) of organisms.It was a priori suggested that the biological traits of the subtidal benthic invertebrate communities within an estuarine environment would respond to the high variability of environmental pressures (natural and human induced) within this type of ecosystem.For this study, traditional taxonomic analysis (species richness, species density and Shannon–Wiener diversity) as well as biological trait analysis were used together for the first time to investigate the response of the subtidal benthic invertebrate communities to the environmental pressures within the Mondego estuary (Portugal).Biological trait analysis, in addition to traditional taxonomic analysis provided a more comprehensive understanding of the functioning within this type of ecosystem. Some of the most important outcomes are: (i) the trait “salinity preference” was the most important trait that distributed the species along the estuary, (ii) the central part of the estuary appeared to be under higher environmental stress levels than the other areas, as suggested by a dominance of some “opportunistic” traits (e.g. small short-lived species), (iii) the ratio between functional diversity (FD) and Shannon–Wiener diversity (H′) indicated lower functional redundancy at the upper reaches of the estuary. Our results, suggest that the ratio (FD/H′) might be a helpful tool to visualize this functional attribute and could potentially be applied to different communities from distinct environments. Using the traditional taxonomic analysis alone, this last functional aspect would not be detectable. Therefore, the inclusion of biological traits analysis is recommendable for estuarine ecological studies.     

Spatio-temporal dynamics of multi-trophic communities reveal ecosystem-wide functional reorganization

Large-scale alterations in marine ecosystems as a response to environmental and anthropogenic pressures have been documented worldwide. Yet, these are primarily investigated by assessing abundance fluctuations of a few dominant species, which inadequately reflect ecosystem-wide changes. In addition, it is increasingly recognized that it is not species identity per se, but their traits that determine environmental responses, biological interactions and ecosystem functioning. In this study, we investigated long-term, spatio-temporal variability in trait composition across multiple organism groups to assess whether functional changes occur in a similar way across trophic levels and whether shifts in trait composition link to environmental change. We combined extensive trait datasets with long-term surveys (30–40 yr) of four organism groups (phytoplankton, zooplankton, benthic invertebrates and fish) in three environmentally distinct areas of a large marine ecosystem. We found similar temporal trajectories in the community weighted mean trait time-series of the different trophic groups, revealing ecosystem-wide functional changes. The traits involved and their dynamics differed between areas, concurrent with climate-driven changes in temperature and salinity, as well as more local dynamics in nutrients and oxygen. This finding highlights the importance of considering both global climate, as well as local external drivers when studying ecosystem changes. Using a multi-trophic trait-based approach, our study demonstrates the importance of integrating community functional dynamics across multiple trophic levels to capture ecosystem-wide responses which could, ultimately, help moving towards a holistic understanding, assessment and management of marine ecosystems.     

Dominance determines fish community biomass in a temperate seagrass ecosystem

Biodiversity and ecosystem function are often correlated, but there are multiple hypotheses about the mechanisms underlying this relationship. Ecosystem functions such as primary or secondary production may be maximized by species richness, evenness in species abundances, or the presence or dominance of species with certain traits. Here, we combine surveys of natural fish communities (conducted in July and August 2016) with morphological trait data to examine relationships between biodiversity and ecosystem function (quantified as fish community biomass) across 14 subtidal eelgrass meadows in the Northeast Pacific (54°N, 130°W). We employ both taxonomic and functional trait measures of diversity to investigate whether ecosystem function is best predicted by species diversity (complementarity hypothesis) or by the presence or dominance of species with particular trait values (selection or dominance hypotheses). After controlling for environmental variation, we find that fish community biomass is maximized when taxonomic richness and functional evenness are low, and in communities dominated by species with particular trait values, specifically those associated with benthic habitats and prey capture. While previous work on fish communities has found that species richness is often positively correlated with ecosystem function, our results instead highlight the capacity for regionally prevalent and locally dominant species to drive ecosystem function in moderately diverse communities. We discuss these alternate links between community composition and ecosystem function and consider their divergent implications for ecosystem valuation and conservation prioritization.     

Macroinvertebrate community responses to land use: a trait-based approach for freshwater biomonitoring in Mongolia

Land-use practices in Mongolia can lead to environmental degradation and consequently affect the structure and function of biological communities. The main aim of this study was to determine land-use effects on freshwater macroinvertebrate communities based on their response to grazing and mining, using a trait-based approach (TBA). The functional structure of macroinvertebrate communities was examined using 86 categories of 16 traits. A total of 13 physical and chemical variables were significantly different among the levels of land-use intensity. Significant declines in functional diversity were observed with increased land-use intensity. The community weighted mean of 19 trait categories for 11 traits varied significantly among different levels of land-use intensity. Traits were significantly explained by environmental variables across a land-use intensity gradient. Water temperature, gravel, nitrate, silt, and cobble were the main predictor variables and explained 28% of the total variance of the trait variation. The functional structure of the macroinvertebrate community was strongly related to environmental conditions. The TBA is an important method in assessing disturbance responses in freshwater communities of steppe and taiga regions, such as in Mongolia and other countries in Central Asia and will be useful in finding best management practices for conserving aquatic ecosystems.

     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.     

Meta‐community theory and stream restoration: evidence that spatial position constrains stream invertebrate communities in a mine impacted landscape

Mining activities, particularly acid mine drainage, often result in adverse effects on stream diversity and ecosystem functioning, and increased concern about these effects has generated a focus on restoration of mine‐impacted waterways. However, many stream restoration projects have not led to increased stream diversity and ecological recovery. One reason for this failure may be that restoration practitioners focus on local environmental conditions and fail to consider the importance of dispersal as a driver of stream invertebrate composition. To test this hypothesis, we used a meta‐community analysis to compare the influence of the local stream conditions with the regional supply of colonists. Invertebrate communities and physico‐chemical conditions were sampled in 37 streams across a mine‐impact gradient on the Stockton Plateau, West Coast of New Zealand's South Island. We found that pH, temperature, dissolved metals, and sediment significantly influenced invertebrate community composition. Furthermore, the spatial location of streams was a good predictor of stream diversity and invertebrate communities, independent of local environmental conditions. This result indicates an important role for regional dispersal barriers in determining stream invertebrate communities. Consequently, consideration of both the locations and strategic preservation of future colonist source streams and potential dispersal barriers during mine planning would enhance post‐mining restoration.     

Plant trait responses to the environment and effects on ecosystem properties

A combined analysis of plant trait responses to the environment, and their effects on ecosystem properties has recently been proposed. In this study, we related the trait composition of plant communities to soil nutrients and disturbance as environmental drivers and to productivity, decomposition and soil carbon as ecosystem properties. We surveyed two sites, one comprising intensively grazed and fertilized grasslands, the other consisting of semi-natural grassland and open heathland. Species abundance and trait values of 49 species were recorded in 69 plots, as well as parameters describing soil resources, land-use disturbances, and ecosystem properties. Our main goal was to test whether the average or the diversity of the trait values of the vegetation had stronger effects on ecosystem properties (mass ratio vs. diversity hypothesis). Structural equation modeling was used to perform a simultaneous analysis of trait responses and effects. Specific leaf area and leaf nutrient contents were always negatively correlated with stem dry matter content and canopy height, indicating greater investments in supportive and nutrient-conserving tissue as plants increased in size. In the agricultural site, disturbance was the single most important factor decreasing plant height, while leaf traits such as specific leaf area and leaf nutrient contents increased with soil resources in heathlands. Productivity was directly or indirectly driven by leaf traits, and investments in structural tissue increased standing biomass and soil carbon. Different environmental drivers in the two sites produced opposing leaf trait effects on litter decomposition. Ecosystem properties were explained by the community mean trait value as predicted by the mass ratio hypothesis. Evidence for effects of functional diversity on productivity and other ecosystem properties was not detected, suggesting that diversity–productivity relationships depend on the length of the investigated environmental gradients. We conclude that changes in community composition and dominance hierarchies deserve the most attention when ecosystem properties must be maintained.     

Mycorrhizal suppression and phosphorus addition influence the stability of plant community composition and function in a temperate steppe

Nutrient enrichment can reduce ecosystem stability, typically measured as temporal stability of a single function, e.g. plant productivity. Moreover, nutrient enrichment can alter plant–soil interactions (e.g. mycorrhizal symbiosis) that determine plant community composition and productivity. Thus, it is likely that nutrient enrichment and interactions between plants and their soil communities co-determine the stability in plant community composition and productivity. Yet our understanding as to how nutrient enrichment affects multiple facets of ecosystem stability, such as functional and compositional stability, and the role of above–belowground interactions are still lacking. We tested how mycorrhizal suppression and phosphorus (P) addition influenced multiple facets of ecosystem stability in a three-year field study in a temperate steppe. Here we focused on the functional and compositional stability of plant community; functional stability is the temporal community variance in primary productivity; compositional stability is represented by compositional resistance, turnover, species extinction and invasion. Community variance was partitioned into population variance defined as community productivity weighted average of the species temporal variance in performance, and species synchrony defined as the degree of temporal positive covariation among species. Compared to treatments with mycorrhizal suppression, the intact AM fungal communities reduced community variance in primary productivity by reducing species synchrony at high levels of P addition. Species synchrony and population variance were linearly associated with community variance with the intact AM fungal communities, while these relationships were decoupled or weakened by mycorrhizal suppression. The intact AM fungal communities promoted the compositional resistance of plant communities by reducing compositional turnover, but this effect was suppressed by P addition. P addition increased the number of species extinctions and thus promoted compositional turnover. Our study shows P addition and AM fungal communities can jointly and independently modify the various components of ecosystem stability in terms of plant community productivity and composition.     

A gradient of nutrient enrichment reveals nonlinear impacts of fertilization on Arctic plant diversity and ecosystem function

Rapid environmental change at high latitudes is predicted to greatly alter the diversity, structure, and function of plant communities, resulting in changes in the pools and fluxes of nutrients. In Arctic tundra, increased nitrogen (N) and phosphorus (P) availability accompanying warming is known to impact plant diversity and ecosystem function; however, to date, most studies examining Arctic nutrient enrichment focus on the impact of relatively large (>25x estimated naturally occurring N enrichment) doses of nutrients on plant community composition and net primary productivity. To understand the impacts of Arctic nutrient enrichment, we examined plant community composition and the capacity for ecosystem function (net ecosystem exchange, ecosystem respiration, and gross primary production) across a gradient of experimental N and P addition expected to more closely approximate warming‐induced fertilization. In addition, we compared our measured ecosystem CO₂ flux data to a widely used Arctic ecosystem exchange model to investigate the ability to predict the capacity for CO₂ exchange with nutrient addition. We observed declines in abundance‐weighted plant diversity at low levels of nutrient enrichment, but species richness and the capacity for ecosystem carbon uptake did not change until the highest level of fertilization. When we compared our measured data to the model, we found that the model explained roughly 30%–50% of the variance in the observed data, depending on the flux variable, and the relationship weakened at high levels of enrichment. Our results suggest that while a relatively small amount of nutrient enrichment impacts plant diversity, only relatively large levels of fertilization—over an order of magnitude or more than warming‐induced rates—significantly alter the capacity for tundra CO₂ exchange. Overall, our findings highlight the value of measuring and modeling the impacts of a nutrient enrichment gradient, as warming‐related nutrient availability may impact ecosystems differently than single‐level fertilization experiments.     

Responses of foraminifera communities to aquaculture‐derived organic enrichment as revealed by environmental DNA metabarcoding

Current monitoring methods to assess benthic impacts of marine finfish aquaculture are based on complex biological indices and/or geochemistry data. The former requires benthic macrofauna morpho‐taxonomic characterization that is time‐ and cost‐intensive, while the latter provides rapid assessment of the organic enrichment status of sediments but does not directly measure biotic impacts. In this study, sediment samples were collected from seven stations at six salmon farms in British Columbia, Canada, and analyzed for geochemical parameters and by eDNA metabarcoding to investigate linkages between geochemistry and foraminifera. Sediment texture across farm sites ranged from sand to silty loam, while the maximum sediment pore‐water sulphide concentration at each site ranged from 1,000 to 13,000 μM. Foraminifera alpha diversity generally increased with distance from cage edge. Adonis analyses revealed that farm site explained the most variation in foraminifera community, followed by sediment type, enrichment status, and distance from cage edge. Farm‐specific responses were observed in diversity analyses, taxonomic difference analyses, and correlation analyses. Results demonstrated that species diversity and composition of foraminifera characterized by eDNA metabarcoding generated signals consistent with benthic biodiversity being impacted by finfish farming activities. This substantiates the validity of eDNA metabarcoding for augmenting current approaches to benthic impact assessments by providing more cost‐effective and practicable biotic measures than traditional morpho‐taxonomy. To capitalize on this potential, further work is needed to design a new nomogram that combines eDNA metabarcoding data and geochemistry data to enable accurate monitoring of benthic impacts of fish farming in a time‐ and cost‐efficient way.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Whole genome scan to detect quantitative trait loci for bovine milk protein composition

The objective of this study was to perform a whole genome scan to detect quantitative trait loci (QTL) for milk protein composition in 849 Holstein–Friesian cows originating from seven sires. One morning milk sample was analysed for the major milk proteins using capillary zone electrophoresis. A genetic map was constructed with 1341 single nucleotide polymorphisms, covering 2829 centimorgans (cM) and 95% of the cattle genome. The chromosomal regions most significantly related to milk protein composition ( P _genome < 0.05) were found on Bos taurus autosomes (BTA) 6, 11 and 14. The QTL on BTA6 was found at about 80 cM, and affected α_S1-casein, α_S2-casein, β-casein and κ-casein. The QTL on BTA11 was found at 124 cM, and affected β-lactoglobulin, and the QTL on BTA14 was found at 0 cM, and affected protein percentage. The proportion of phenotypic variance explained by the QTL was 3.6% for β-casein and 7.9% for κ-casein on BTA6, 28.3% for β-lactoglobulin on BTA11, and 8.6% for protein percentage on BTA14. The QTL affecting α_S2-casein on BTA6 and 17 showed a significant interaction. We investigated the extent to which the detected QTL affecting milk protein composition could be explained by known polymorphisms in β-casein , κ -casein , β-lactoglobulin and DGAT1 genes. Correction for these polymorphisms decreased the proportion of phenotypic variance explained by the QTL previously found on BTA6, 11 and 14. Thus, several significant QTL affecting milk protein composition were found, of which some QTL could partially be explained by polymorphisms in milk protein genes.