Data and code from: Sex-dependent carry-over effects of non-breeding habitat use on breeding performance in the Eurasian Curlew
Data files
Oct 01, 2026 version files 591 MB
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Curlew_COE_habitat.zip
590.99 MB
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README.md
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Abstract
In migratory birds, the effects of habitat use during the non-breeding period can carry over to body condition in subsequent stages of the annual cycle and affect the breeding performance of individuals. We investigated whether the habitat types used on the non-breeding grounds and at spring stopover sites influenced nest success through carry-over effects in the Eurasian Curlew, a migratory species with declining populations. We also investigated the association between habitat use on the breeding grounds and nest success. Using a Europe-wide dataset of 124 individually GPS-tracked Curlews, comprising 233 periods covering non-breeding, spring migration, and breeding periods collected between 2016 and 2025, we determined habitat use on non-breeding grounds, spring stopovers, and breeding grounds and estimated nest success of each individual. We related Curlew nest success to habitat use, while considering sex-dependent responses, differences between breeding grounds, and the time from arrival to incubation onset, using principal component regression. We found weak carry-over effects of habitat use gradients on nest success, which appeared to be stronger in females than in males. Nest success increased with the use of mudflats and wetlands but declined with the use of salt marshes on the non-breeding grounds in females. Use of grasslands at stopover sites was associated with decreased nest success in females. Use of grasslands and wetlands on the breeding grounds was associated with decreased and increased nest success, respectively, in males. Our results suggest that conserving intertidal mudflats on the non-breeding grounds and maintaining high-quality stopover habitats in agricultural landscapes may contribute to improved breeding success. The identified links between non-breeding habitat use and breeding success emphasise the importance of conservation throughout the annual cycle.
Data and R script to reproduce the modelling in the accompanying journal article (Part I). The models are stored for convenience. Furthermore, the code for the assessment of nest success based on the analysis of GPS tracking data is provided (Part II). All files are available in Curlew_COE_habitat.zip
Description of the data and file structure
File tree
- data
- models
- scripts
Part I – habitat use effects on nest success
Data
dt_hab_Curlew.csv dataset includes the raw data, arranged into the following variables:
ID_seas= bird ID and year of breedingBird_ID= ID of the individualSex= male (M) of female (F)Non_gr= non-breeding grounds, ATL-S = southern Atlantic coast, ATL-C = central Atlantic coast, ATL-N = northern Atlantic coast, NSea = North Sea coastBr_gr= breeding grounds, W = western Europe, NE = northeastern EuropeNest_succ= nest was successfully incubated (1) or not (0)X_br= mean longitude of the breeding grounds (redacted as sensitive data)Y_br= mean latitude of the breeding grounds (redacted as sensitive data)X_non= mean longitude of the non-breeding grounds (redacted as sensitive data)Y_non= mean latitude of the non-breeding grounds (redacted as sensitive data)Dist= shortest migration distance from non-breeding grounds to breeding grounds (km)Prebr= length of the pre-breeding period, i.e. a period between arrival at the breeding grounds and the onset of incubation (day)non_arable= arable land use on non-breeding grounds (proportions of use)non_grass= grasslands use on non-breeding grounds (proportions of use)non_marsh_salt= salt marshes use on non-breeding grounds (proportions of use)non_mudflats= intertidal mudflats use on non-breeding grounds (proportions of use)non_salines= salines use on non-breeding grounds (proportions of use)non_wetland= inland wetlands use on non-breeding grounds (proportions of use)st_arable= arable land use at stopover sites (proportions of use)st_grass= grasslands use at stopover sites (proportions of use)st_marsh_salt= salt marshes use at stopover sites (proportions of use)st_mudflats= intertidal mudflats use at stopover sites (proportions of use)st_wetland= inland wetlands use at stopover sites (proportions of use)br_arable= arable land use on breeding grounds (proportions of use)br_grass= grasslands use on breeding grounds (proportions of use)br_mudflats= intertidal mudflats use on breeding grounds (proportions of use)br_wetland= inland wetlands use on breeding grounds (proportions of use)
In the variables describing proportions of use of particular habitat classes, NA means the absence of a habitat class in a given breeding ground, non-breeding ground, or stopover site.
land_use_classes.xlsx Habitat use classes on sheets LULC_Coastal_Zones, LULC_CLCplus_backbone, LULC_CGLS as defined in the original data sources, and the habitat classes used for analyses. Detailed metadata is included in a separate sheet within the file.
summary_table_prop_dirich.csv A table including proportions of habitat use derived from Dirichlet regression, including the 95% credible interval of the estimates. Data is used for plotting.
summary_table_reg_prop_dirich.csv A table including proportions of habitat use in non-breeding and breeding areas, derived from Dirichlet regression, including the 95% credible interval of the estimates. Data is used for plotting.
Models
Mod_PCR.rds Stored model testing sex-specific effects of habitat use on non-breeding and breeding grounds on nest success, including the length of the pre-breeding period and breeding grounds (categorical predictor) as covariates. The model provides estimates of the fixed effects of the following variables:
Sex= male (M) of female (F)Prebr= length of the pre-breeding period, i.e. a period between arrival at the breeding grounds and the onset of incubation (day)Br_gr= breeding grounds, W = western Europe, NE = northeastern EuropePCX_non= gradient of habitat use on the non-breeding grounds derived from a principal component analysis; X denotes particular principal componentPCX_br= gradient of habitat use on the breeding grounds derived from a principal component analysis; X denotes particular principal componentSex:PCX_non= sex-specific effects of a gradient of habitat use on the non-breeding grounds derived from a principal component analysis; X denotes particular principal componentSex:PCX_br= sex-specific effects of a gradient of habitat use on the breeding grounds derived from a principal component analysis; X denotes particular principal component
The model also provides estimate of the variance of Bird_ID random effect (ID of the individual).
Mod_PCR_stop.rds Stored model testing sex-specific effects of habitat use at stopover sites on nest success, including the length of the pre-breeding period and breeding grounds (categorical predictor) as covariates. The model provides estimates of the effects of the following variables:
Sex= male (M) of female (F)Prebr= length of the pre-breeding period, i.e., a period between arrival at the breeding grounds and the onset of incubation (day)Br_gr= breeding grounds, W = western Europe, NE = northeastern EuropePCX_st= gradient of habitat use on the stopover sites derived from a principal component analysis; X denotes the number of a principal componentSex:PCX_st= sex-specific effects of a gradient of habitat use on the stopover sites derived from a principal component analysis; X denotes particular principal component
The model also provides estimate of the variance of Bird_ID random effect (ID of the individual).
Mod_hab_dirich_all.rds Stored model investigating the proportions of habitat types used on non-breeding grounds, stopovers, and breeding grounds. The model provides estimates of the proportions of habitat use:
munonHAB_propmatrixnon_Intercept= log odds of habitat use proportions (HAB) on non-breeding grounds, where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), salines, or wetlands (inland wetlands)mubrHAB_propmatrixbr_Intercept= log odds of habitat use proportions (HAB) on breeding grounds, where HAB is grass (grasslands), mudflats (intertidal mudflats), or wetlands (inland wetlands)mustHAB_propmatrixst_Intercept= log odds of habitat use proportions (HAB) on stopover sites, where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), or wetlands (inland wetlands)
Mod_hab_dirich_reg.rds Stored model investigating the proportions of habitat types used on non-breeding grounds, stopovers, and breeding grounds, grouped by the non-breeding areas and breeding areas. The model provides grouped estimates of the proportions (log odds) of habitat use:
munonHAB_propmatrixnon_Intercept= habitat use proportions (log odds) on the non-breeding grounds for Curlews using the reference non-breeding (central Atlantic coast) and breeding areas (northeastern Europe), where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), salines, or wetlands (inland wetlands)mubrHAB_propmatrixbr_Intercept= habitat use proportions (log odds) on the breeding grounds for Curlews using the reference non-breeding (central Atlantic coast) and breeding areas (northeastern Europe), where HAB is grass (grasslands), mudflats (intertidal mudflats), or wetlands (inland wetlands)mustHAB_propmatrixst_Intercept= habitat use proportions (log odds) on the stopover sites for Curlews using the reference non-breeding (central Atlantic coast) and breeding areas (northeastern Europe), where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), or wetlands (inland wetlands)munonHAB_propmatrixnon_Non_grAREA= difference in habitat use proportions (log odds) of a given non-breeding habitat type (HAB) between Curlews using a given (AREA) and the reference non-breeding area (central Atlantic coast), where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), salines, or wetlands (inland wetlands), and AREA is ATLMN (northern Atlantic coast), ATLMS (southern Atlantic coast), or NSea (North Sea coast)munonHAB_propmatrixnon_Br_grW= difference in habitat use proportions (log odds) of a given non-breeding habitat type (HAB) between Curlews using western Europe and the reference breeding area (northeastern Europe), where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), salines, or wetlands (inland wetlands), and W is western European breeding groundmunbrHAB_propmatrixbr_Non_grAREA= difference in habitat use proportions (log odds) of a given breeding habitat type (HAB) between Curlews using a given (AREA) and the reference non-breeding area (central Atlantic coast), where HAB is grass (grasslands), mudflats (intertidal mudflats), or wetlands (inland wetlands), and AREA is ATLMN (northern Atlantic coast), ATLMS (southern Atlantic coast), or NSea (North Sea coast)mubrHAB_propmatrixbr_Br_grW= difference in habitat use proportions (log odds) of a given breeding habitat type (HAB) between Curlews using western European and the reference breeding area (northeastern Europe), where HAB is grass (grasslands), mudflats (intertidal mudflats), or wetlands (inland wetlands), and W is western European breeding groundmustHAB_propmatrixst_Non_grAREA= difference in habitat use proportions (log odds) of a given stopover habitat type (HAB) between Curlews using a given (AREA) and the reference non-breeding area (central Atlantic coast), where HAB is grass (grasslands), marshsalt (salt marshes), mudflats (intertidal mudflats), or wetlands (inland wetlands), and AREA is ATLMN (northern Atlantic coast), ATLMS (southern Atlantic coast), or NSea (North Sea coast)mustHAB_propmatrixst_Br_grW= difference in habitat use proportions (log odds) of a given stopover habitat type (HAB) between Curlews using western European and the reference breeding area (northeastern Europe), where HAB is grass (grasslands), mudflats (intertidal mudflats), or wetlands (inland wetlands), and W is western European breeding ground
Scripts
R-script_COE_habitat_v3.R Code for preparing the datasets, fitting the models, and plotting the figures for the analysis of the effects of habitat use on nest success in the Eurasian Curlew.
Scripts below are used for the assessment of nest success based on the analysis of GPS tracking data
Nest_succ_user.R Code to run the workflow for the assessment of nest success based on the analysis of GPS tracking data: loading packages and data, setting parameters for the analyses, running all the analyses, and saving the outputs of the analyses and plots for inferences.
Nest_succ_dt_dbscan.R Data preparation and DBSCAN.
Nest_succ_helpers.R Helper functions and code for creating folders with the outputs
Nest_succ_main.R Main code for the assessment of nest success
Nest_succ_dist.R Nest distance analysis
Part II - assessment of nest success based on the analysis of GPS tracking data
Analysis of bird GPS tracking data in the breeding season, allowing to identify nesting attempts and assess nest success. Birds are analyzed one by one because the user needs to adjust the parameters based on the outcomes of individual steps of the workflow.
Documentation
The workflow run by Nest_succ_user.R can export bird tracking data with identified clusters of possible nest locations, summary table of the nest cluster analysis, and coordinates of the possible nests. Plots depicting the results of the analysis of nest clusters and distance to nest analysis are exported. The following parameters are defined before running the analyses:
bird_name = species name (shortened for usage in file names and plot titles)
bird_ID = ID of the analysed individual, usually loaded from a list of the assessed bird species
dt_nest = dataset including tracking data. Created by the classification application "Migration and stationary sites", available at "https://www.moveapps.org/apps/browser/164b6064-206a-4e44-9e6c-8516a288e18a"
Year_br = breeding season for the analysis, a single year
thin_time = thinning interval for the data (in min), used when getRecursions() function has not enough memory (default = 10)
cl_eps = epsilon (in m), the maximum distance between two points to be considered neighbours in DBSCAN, usually values of 2, 5, 10, or 20 m
cl_pts = minimum number of points in the cluster (default = 100)
tot_w = total weight for DBSCAN weighted by residence time (in hours), usually between 1 and 30 hours
bcp_thres = minimum posterior probability to consider given change points in the analysis (default = 0.3)
radius = radius (in m) used for calculation of the pattern by recurse() (default = 20)
run_recurse = run trajectory and revisits analysis, disable it if speed of computation is preferred (default TRUE)
dbscan_time = time-weighted DBSCAN is run (TRUE), or a classical DBSCAN (FALSE)
first_date = the first date of an assessed period, format YYYY-MM-DD
date_range = length of the assessed period (in days), starting from the first_date
cl_no = number of the cluster assessed, see "clus_summary" created within the workflow
Interpretation of the plots
See the Supporting Information related to the article for details on the interpretation of the plots generated by the algorithm.
Usage notes
R is required to run all the scripts, as well as to load the stored models (.rds files). The models can be loaded using the base R function readRDS(), but we recommend following the code provided in the R scripts in order to assign names to the models anticipated within the workflow.
Microsoft Excel or a similar spreadsheet editor can be used to open the data sets. All CSV files are semicolon-delimited.
