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Breeding in an agricultural landscape 243


et al., 2006) date of hatching (Mayfield, 1975; Kubelka et al., 2018). For predated nests, we calculated exposure from the day of nest finding until the midpoint between the last positive and the first negative visit to the nest. For all other outcomes (i.e. unknown, abandoned, flooded, de- stroyed by agricultural machinery), we defined exposure time from the day of nest finding until the last positive nest visit, following standard protocols (Kubelka et al., 2018). We focused trail cameras on 116 nests to identify species of nest predators and the date of hatching.


Statistical analyses


PLATE 2 Protection zone around collared pratincole nest, marked temporarily by 1.5 m-tall wooden poles. This photograph was taken using a telephoto lens that distorts the actual distance between the two wooden poles.


The survival and productivity of ground-nesting birds


are influenced by predation (Martin, 1993; Roodbergen et al., 2012; Kubelka et al., 2018). To investigate relation- ships between pratincole breeding success and number of nest predators removed from the study area, we collected data from hunters. Potential huntable nest predators include mammals (red fox Vulpes vulpes, golden jackal Canis aur- eus, European badger Meles meles) and birds (Eurasian magpie Pica pica, hooded crow Corvus cornix). We re- quested hunting bag data from professional hunters for the period 2017–2021 and we aggregated the number of individuals killed for each year. Hunting bag data relate to the regional hunting district (c. 26,000 ha), which covers c. 65% of collard pratincole nest sites (National Game Management Database, 2022).


Estimating daily and total nest survival


To investigate the effects of year and agricultural habitat on nesting success, we estimated daily and total nest survival rates using calculations provided by Mayfield (1975). We applied this method to estimate the chances of a clutch surviving daily and for the full nesting period (egg laying + incubation period) by defining the daily nest survival rate as the number of failed nests divided by the sum of exposure days (Johnson, 1979). We calculated total nest survival using Mayfield’s formula: (daily nest survival)nesting period in days, where the total nesting period is egg laying period + incubation (20 days in total). We defined exposure time as the number of days from find- ing a nest to the confirmed (or expected) last day of the breeding attempt. The incubation period started when the complete clutch had been laid. For those nests that hatched (i.e. at least one chick hatched), we calculated the exposure time from the day of nest finding to the confirmed (or predicted, using the egg floating methodology; Székely


We performed the statistical analyses in R 3.3.3 (R Core Team, 2021). To identify relationships amongst individual- level reproduction success metrics (i.e. nests hatched or failed and the number of hatched chicks), we used general- ized linear models (GLMs) with factors of year, habitat type, Julian day of egg laying, distance from the closest field boundary, distance from the closest water body and colony density. The latter factor represented the mean distance from the focal nest to the three closest nests within the same breeding colony. As nesting success was a binary response variable (hatched or failed), we used a logistic regression GLM with the logit link error function. Field boundaries and water bodies were available in shape files. We computed the mean distance from the three neighbouring nests using the nndist spatial neighbour- hood function available in the spatstat package in R.For the start of incubation, we used the Julian day of clutch completion at a given nest. We used ANOVA, imple- mented using the lm function, to analyse the associations between (1) clutch size and habitat type, (2) timing of hatching and habitat type, and (3)daily nest survival (aggregated for years and habitats).


Results


Timing of breeding and breeding success During 2012–2021, we found 315 nests, for 212 of which we also determined the hatching date of the first chick. Egg laying started in late April and terminated in mid July (c. 2.5 months duration), with most nests (60%) laid dur- ing 25 May–15 June. The first eggs hatched on 16 May, which implies that the clutch was completed on 29 April. The latest hatch date was recorded on 3 August. The mean hatching date was 15 June ± SE 1 day (n = 212 nests; Table 1, Supplementary Fig. 1). Collared pratincoles bred in three agricultural habitats:


most nests were in row crops (48%), followed by fallow lands (29%) and spring cover crops (23%, n = 315 nests; Table 1). The timing of breeding was different between


Oryx, 2024, 58(2), 240–249 © The Author(s), 2023. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605323000911


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