Evidence That Population Increase and Range Expansion by Eurasian
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bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.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 available under aCC-BY-NC-ND 4.0 International license. 1 Evidence that population increase and range expansion by 2 Eurasian Sparrowhawks has impacted avian prey populations 3 4 Christopher Paul Bell bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.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 available under aCC-BY-NC-ND 4.0 International license. 2 5 Abstract 6 The role of increased predator numbers in the general decline of bird populations in the late 7 20th century remains controversial, particularly in the case of the Eurasian Sparrowhawk, for 8 which there are contradictory results concerning its effect on the abundance of potential 9 prey species. Previous studies of breeding season census data for Sparrowhawks and prey 10 species in Britain have measured predator abundance either as raw presence-absence data 11 or as an estimate derived from spatially explicit modelling, and have found little evidence of 12 association between predator and prey populations. Here, a predator index derived from 13 site-level binary logistic modelling was used in a regression analysis of breeding census data 14 on 42 prey species, with significant effects emerging in 27 species (16 positive, 11 negative). 15 The frequency of significant positive associations may indicate the tracking of prey 16 abundance by Sparrowhawks, which would tend to cancel out any negative predation effect 17 on prey populations, rendering it difficult to detect using census data. If so, the negative 18 effects that have emerged for some species may underestimate the impact of increased 19 Sparrowhawk numbers on the prey populations concerned. Nevertheless, estimates of the 20 effect on national populations of prey species, obtained by combining effect sizes with a 21 measure of the increase in site occupancy by the predator, suggest the possibility of a 22 substantial impact. It cannot, therefore, be ruled out that the increase in abundance and 23 range expansion of Sparrowhawks made a significant contribution to contemporaneous 24 declines in the populations of many of its prey species. 25 bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (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 available under aCC-BY-NC-ND 4.0 International license. 3 26 Introduction 27 The marked population decline that occurred across a wide range of bird species in the 28 British Isles during the late 20th Century is an event of major conservation concern that 29 remains imperfectly understood. Consequently, it is still liable to generate controversy, 30 particularly over the possibility that increased predator populations may have played a role 31 (Gibbons et al. 2007, Roos et al. 2018). Declines occurred chiefly among songbirds, of which 32 the major predator in western Europe is the Eurasian Sparrowhawk Accipiter nisus 33 (henceforth ‘Sparrowhawk’), a species that increased rapidly in range and abundance during 34 the same period, following a pesticide-related crash in the 1940s and 50s (Newton 1986). 35 The main source of evidence for population declines in Britain is the Common Birds Census 36 (CBC), which ran from 1962 to 2000, and involved intensive field survey designed to 37 estimate the number of breeding territories in a census plot (Marchant 1983). Previous 38 analyses of CBC data have concluded that there is little evidence for any connection 39 between increased Sparrowhawk numbers and songbird population declines (Thomson et 40 al. 1998, Newson et al. 2010), but analyses of data from another census scheme, the Garden 41 Bird Feeding Survey (GBFS), have consistently found evidence of a negative association 42 between Sparrowhawks and House Sparrows (Bell et al. 2010, Swallow et al. 2015, 2016, 43 Jones-Todd 2018). 44 GBFS data is derived from weekly maximum counts made at bird feeding stations over the 45 winter months (Chamberlain et al. 2005), which can be averaged to derive an annually 46 varying index of species abundance, with that for Sparrowhawk being effectively the 47 proportion of weeks during which attacks on a feeding station were observed. The GBFS- 48 derived Sparrowhawk index may thus be taken to be a continuous measure of predation bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (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 available under aCC-BY-NC-ND 4.0 International license. 4 49 pressure at a point location, resulting in commensurate declines in the use of the feeder at 50 that location by potential prey species. CBC data are recorded during the breeding season, 51 and refer to a much wider area, typically 20-150 ha depending on the habitat, and can 52 therefore be more reliably used to infer population effects of a predator. However, they 53 provide a much less precise measure of predator activity, effectively devolving to presence 54 or absence in any given year. 55 Newson et al. (2010) addressed this problem by using smoothed relative abundance 56 surfaces to produce year-specific predator indices for each CBC site within their focal area 57 (England), effectively using broad-scale spatio-temporal trends to estimate variation in local 58 abundance. However, this approach may fail to capture variation in predator activity at the 59 narrower scale represented by the area of a typical CBC census plot, accurate 60 representation of which requires modelling at the individual site level. Here, therefore, I 61 reprise the approach used by Newson et al. (2010) for a wider range of potential prey 62 species, using indices of Sparrowhawk activity derived from site-level modelling. 63 Materials and Methods 64 Model structure 65 The approach used retains the basic structure of the model used by Newson et al. (2010) 66 (see appendix), and implements a regression of change in log prey population against 67 change in a log predator index: 푡−1 µ푖,푡 푃푖,푡 + 1 68 ln ( ) = ∑ 푟푗 + 훼. ln ( ) Equation 1 µ푖,1 푃푖,1 + 1 푗=1 bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (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 available under aCC-BY-NC-ND 4.0 International license. 5 69 where µi,t is estimated prey population and Pi,t an estimate of predator activity in site i and 70 year t, rj is the instantaneous rate of change in global prey population caused by all factors 71 other than predator activity in year j, and α is the effect of the change in the log predator 72 index (ln(Pi,t + 1)) between the first year with site data (t=1) and year t. Change in the log 73 predator index underestimates change in log predator activity due to the need to 74 accommodate instances of zero activity by adding one, but remains highly correlated with it. 75 The model was implemented in R version 3.1.2 (R core team 2014) with prey population as 76 the response variable, using Poisson errors and a loge link: 77 푡−1 푃푖,푡 + 1 78 µ푖,푡 = exp [∑ 푟푗 + 훼. ln ( ) + ln µ푖,1] Equation 2 푃푖,1 + 1 푗=1 79 80 Estimates of rj emerge as year effects with each data year represented in the linear model as 81 a binary variable with the value 1 corresponding to all higher values of t at site level (i.e. the 82 effect of previous years is ‘present’) and otherwise as 0, while ln µi,1 is estimated as a site 83 effect, with each site represented as a level within a categorical variable (cf. Freeman & 84 Newson 2008), or in some cases as more than one level to take account of changes in the 85 size of the area censused. Variance in change in prey population between the first and any 86 subsequent year with site data is therefore partitioned between the effect of change in 87 predator activity over the same period, and the effect of conditions in each of the 88 intervening years. The implementation of Newson et al. (2010) included a number of 89 additional covariates, which are excluded here for reasons set out in the appendix. bioRxiv preprint doi: https://doi.org/10.1101/491522; this version posted July 25, 2019. The copyright holder for this preprint (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 available under aCC-BY-NC-ND 4.0 International license. 6 90 Derivation of predator indices 91 CBC data were supplied by volunteer surveyors as part of a national scheme covering 92 around 250 plots each year. There was relatively high turnover and a policy of replacing a 93 lapsed plot with another of similar character, so the number of years surveyed at individual 94 sites varied considerably.