Title : Functional traits of carabid beetles reveal seasonal variation in community assembly in 1 annual crops

24 1. Trait-based community assembly studies have mostly been addressed along spatial 25 gradients, and do not consider explicitly a fundamental dimension governing community 26 assembly, the time. Nevertheless, such consideration seems particularly necessary in systems 27 in which organisms have to face regular disturbances and rapid changes in vegetation 28 phenology, such as in intensively managed farmlands. 29 2. In this study, we aimed at understanding how the functional diversity of carabid beetle 30 communities varied across the growing season in response to crop type. We tested three 31 alternative hypotheses on mechanisms underlying the community assembly. 32 3. We used data from a long-term monitoring conducted over nine years in an intensively33 managed farmland in central western France, in a total of 625 fields. First, we measured 34 morphological traits related to body size, dispersal mode, and resource acquisition on the 13 35 dominant carabid species (> 85 % of all trapped individuals) and identified three independent 36 dimensions of functional specialization within our species pool along axes of a PCA and 37 highlighted key traits for community assembly. Second, we evaluated the community 38 assembly temporal dynamics and the impact of habitat filtering and niche differentiation in 39 the different crop types with time, using linear mixed-effects models. 40 4. We showed that functional species assembly of carabid beetle communities occurring in 41 crop fields varies importantly intra-annually, with strong variations in these dynamics 42 depending on crop type and crop phenology. Each crop acted as a filter on carabid 43 communities for body size and resource-acquisition traits, and functional differentiation 44 between crops increased with time. We did not find any evidence of habitat filtering on traits 45 related to dispersal mode. 46 5. Our results emphasize the major role of crop phenology but also disturbances involved by 47 agricultural practices such as crop harvesting on changes in community assembly, likely due 48 . CC-BY-NC-ND 4.0 International license available under a (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint this version posted February 5, 2021. ; https://doi.org/10.1101/2021.02.04.429696 doi: bioRxiv preprint

[1]  C. Violle,et al.  Grassland-to-crop conversion in agricultural landscapes has lasting impact on the trait diversity of bees , 2020, Landscape ecology.

[2]  J. Recasens,et al.  Increasing crop heterogeneity enhances multitrophic diversity across agricultural regions , 2019, Proceedings of the National Academy of Sciences.

[3]  V. Bretagnolle,et al.  The consumption pattern of 28 species of carabid beetles (Carabidae) to a weed seed, Viola arvensis. , 2018, Bulletin of entomological research.

[4]  V. Bretagnolle,et al.  Spatiotemporal dynamics of the agricultural landscape mosaic drives distribution and abundance of dominant carabid beetles , 2017, Landscape Ecology.

[5]  José Paulo Sousa,et al.  Handbook of protocols for standardized measurement of terrestrial invertebrate functional traits , 2017 .

[6]  Ashley P. G. Dowling,et al.  Pitfalls and preservatives: a review , 2016 .

[7]  Frédéric Jiguet,et al.  Rapid adjustment of bird community compositions to local climatic variations and its functional consequences , 2015, Global change biology.

[8]  Ben Collen,et al.  Global effects of land use on local terrestrial biodiversity , 2015, Nature.

[9]  Carsten Thies,et al.  Species’ traits influence ground beetle responses to farm and landscape level agricultural intensification in Europe , 2014, Journal of Insect Conservation.

[10]  Gregor Hagedorn,et al.  Fauna Europaea – all European animal species on the web , 2014, Biodiversity data journal.

[11]  D. Roy,et al.  National patterns of functional diversity and redundancy in predatory ground beetles and bees associated with key UK arable crops , 2014 .

[12]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[13]  C. Leifert,et al.  Crop, field boundary, productivity and disturbance influences on ground beetles (Coleoptera, Carabidae) in the agroecosystem , 2013 .

[14]  M. Pärtel,et al.  Functional species pool framework to test for biotic effects on community assembly. , 2012, Ecology.

[15]  Katharine N. Suding,et al.  Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes , 2012 .

[16]  H. Turin,et al.  Forty years of carabid beetle research in Europe – from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation , 2011, ZooKeys.

[17]  Lenore Fahrig,et al.  Functional landscape heterogeneity and animal biodiversity in agricultural landscapes. , 2011, Ecology letters.

[18]  P. Legendre,et al.  A distance-based framework for measuring functional diversity from multiple traits. , 2010, Ecology.

[19]  M. Evans,et al.  A comparison of adaptations to running, pushing and burrowing in some adult Coleoptera: especially Carabidae , 2009 .

[20]  T. G. Forsythe Locomotion in ground beetles (Coleoptera carabidae): an interpretation of leg structure in functional terms , 2009 .

[21]  K. Barton MuMIn : multi-model inference, R package version 0.12.0 , 2009 .

[22]  Felix L. Wäckers,et al.  Hibernation of predatory arthropods in semi-natural habitats , 2009, BioControl.

[23]  D. Moser,et al.  Ground-dwelling predators can affect within-field pest insect emergence in winter oilseed rape fields , 2009, BioControl.

[24]  J. Holland,et al.  Contrasting the farm-scale spatio-temporal dynamics of boundary and field overwintering predatory beetles in arable crops , 2009, BioControl.

[25]  Nathan J B Kraft Assembly in an Amazonian Forest Functional Traits and Niche-Based Tree Community , 2008 .

[26]  A. Matalin Typology of life cycles of ground beetles (Coleoptera, Carabidae) in Western Palaearctic , 2007, Entomological Review.

[27]  D. K. Weaver,et al.  Grasshoppers and locusts , 2007 .

[28]  Stephen P. Hubbell,et al.  Neutral theory in community ecology and the hypothesis of functional equivalence , 2005 .

[29]  Ann-Christin Weibull,et al.  Species richness in agroecosystems: the effect of landscape, habitat and farm management , 2003, Biodiversity & Conservation.

[30]  M. Loreau,et al.  On testing temporal niche differentiation in carabid beetles , 1989, Oecologia.

[31]  M. Nelemans Possibilities for flight in the carabid beetle Nebria brevicollis (F.) , 1987, Oecologia.

[32]  T. V. Huizen The significance of flight activity in the life cycle of Amara plebeja Gyll. (Coleoptera, Carabidae) , 2004, Oecologia.

[33]  R. O’Hara,et al.  Species decline—but why? Explanations of carabid beetle (Coleoptera, Carabidae) declines in Europe , 2003, Oecologia.

[34]  Ann-Christin Weibull,et al.  Species composition in agroecosystems: The effect of landscape, habitat, and farm management , 2003 .

[35]  Garth N. Foster,et al.  Effect of land disturbance and stress on species traits of ground beetle assemblages , 2001 .

[36]  Keith E. MasJeus,et al.  Quantifying the Impact of , 2000 .

[37]  Bernhard Kromp,et al.  Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation impacts and enhancement , 1999 .

[38]  T. New The role of ground beetles (Coleoptera: Carabidae) in monitoring programmes in Australia , 1998 .

[39]  Eyre,et al.  Effects of habitat type and grassland management practices on the body size distribution of carabid beetles , 1994, Pedobiologia.

[40]  Paul A. Keddy,et al.  Assembly and response rules: two goals for predictive community ecology , 1992 .

[41]  G. Ball,et al.  The mandibles of some adult ground beetles: structure, function, and the evolution of herbivory (Coleoptera: Carabidae) , 1991 .

[42]  H. Turin,et al.  Loss of habitats and changes in the composition of the ground and tiger beetle fauna in four West European countries since 1950 (Coleoptera: Carabidae, cicindelidae) , 1989 .

[43]  P. J. Boer,et al.  Changes in the distribution of carabid beetles in The Netherlands since 1880. II. Isolation of habitats and long-term time trends in the occurence of carabid species with different powers of dispersal (Coleoptera, Carabidae)☆ , 1988 .

[44]  Madalynne Schoenfeld Seasonal changes , 1983 .

[45]  H. Thiele Carabid Beetles in Their Environments: A Study on Habitat Selection by Adaptations in Physiology and Behaviour , 1977 .

[46]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[47]  Fitting linear mixed-effects models , 2022 .