Daniel Takahashi

Detlef Weigel

eLife assessment
This study offers a useful discussion of the well-accepted abundance-occupancy relationship in macroecology. While using the ebird large dataset to revisit the theme is interesting, multiple unresolved confounding factors exist, leaving the results inadequate to overturn the repeatedly confirmed abundancy-occupancy relationship.

anonymous

Reviewer #1 (Public Review):
Summary:
This article presents a meta-analysis that challenges established abundance-occupancy relationships (AORs) by utilizing the largest known bird observation database. The analysis yields contentious outcomes, raising the question of whether these findings could potentially refute AORs.
Strengths:
The study employed an extensive aggregation of datasets to date to scrutinize the abundance-occupancy relationships (AORs).
Weaknesses:
While the dataset employed in this research holds promise, a rigorous justification of the core assumptions underpinning the analytical framework is inadequate. The authors should thoroughly address the correlation between checklist data and global range data, ensuring that the foundational assumptions and potential confounding factors are explicitly examined and articulated within the study's context.

Reviewer #2 (Public Review):
Summary:
The goal is to ask if common species when studied across their range tend to have larger ranges in total. To do this the authors examined a very large citizen science database which gives estimates of numbers, and correlated that with the total range size, available from Birdlife. The average correlation is positive but close to zero, and the distribution around zero is also narrow, leading to the conclusion that, even if applicable in some cases, there is no evidence for consistent trends in one or other direction.
Strengths:
The study raises a dormant question, with a large dataset.
Weaknesses:
This study combines information from across the whole world, with many different habitats, taxa, and observations, which surely leads to a quite heterogeneous collection.
First, scale. Many of the earlier analyses were within smaller areas, and for example, ranges are not obviously bounded by a physical barrier. I assume this study is only looking at breeding ranges; that should be stated, as 40% of all bird species migrate, and winter limitation of populations is important. Also are abundances only breeding abundances or are they measured through the year? Are alien distributions removed?
Second, consider various reasons why abundance and range size may be correlated (sometimes positively and sometimes negatively) at large scales. Combining studies across such a large diversity of ecological situations seems to create many possibilities to miss interesting patterns. For example:
(1) Islands are small and often show density release.
(2) North temperate regions have large ranges (Rapoport's rule) and higher population sizes than the tropics.
(3) Body size correlates with global range size (I am unsure if this has recently been tested but is present in older papers) and with density. For example, cosmopolitan species (barn owl, osprey, peregrine) are relatively large and relatively rare.
(4) In the consideration of alien species, it certainly looks to me as if the law is followed, with pigeon, starling, and sparrow both common and widely distributed. I guess one needs to make some sort of statement about anthropogenic influences, given the dramatic changes in both populations and environments over the past 50 years.
(5) Wing shape correlates with ecological niche and range size (e.g. White, American Naturalist). Aerial foraging species with pointed wings are likely to be easily detected, and several have large ranges reflecting dispersal (e.g. barn swallow).
Third, biases. I am not conversant with ebird methodology, but the number appearing on checklists seems a very poor estimate of local abundance. As noted in the paper, common species may be underestimated in their abundance. Flocking species must generate large numbers, skulking species few. The survey is often likely to be in areas favorable to some species and not others. The alternative approach in the paper comes from an earlier study, based on ebird but then creating densities within grids and surely comes with similar issues. 
Biases are present in range as well. Notably, tropical mountain-occupying species have range sizes over-estimated because holes in the range are not generally accounted for (Ocampo-Peñuela et al Nature communications). These species are often quite rare too.
Fourth, random error. Random error in ebird assessments is likely to be large, with differences among observers, seasons, days, and weather (e.g. Callaghan et al. 2021, PNAS). Range sizes also come with many errors, which is why occupancy is usually seen as the more appropriate measure.
If we consider both range and abundance measurements to be subject to random error in any one species list, then the removal of all these errors will surely increase the correlation for that list (the covariance shouldn't change but the variances will decrease). I think (but am not sure) that this will affect the mean correlation because more of the positive correlations appear 'real' given the overall mean is positive. It will definitely affect the variance of the correlations; the low variance is one of the main points in the paper. A high variance would point to the operation of multiple mechanisms, some perhaps producing negative correlations (Blackburn et al. 2006).
On P.80 it is stated: "Specifically, we can quantify how AOR will change in relation to increases in species richness and sampling duration, both of which are predicted to reduce the magnitude of AORs" I haven't checked the references that make this statement, but intuitively the opposite is expected? More species and longer durations should both increase the accuracy of the estimate, so removing them introduces more error? Perhaps dividing by an uncertain estimate introduces more error anyway. At any rate, the authors should explain the quoted statement in this paper.
It would be of considerable interest to look at the extreme negative and extreme positive correlations: do they make any biological sense?
Discussion:
I can see how publication bias can affect meta-analyses (addressed in the Gaston et al. 2006 paper) but less easily see how confirmation bias can. It seems to me that some of the points made above must explain the difference between this study and Blackburn et al. 2006's strong result.
Certainly, AOR really does seem to be present in at least some cases (e.g. British breeding birds) and a discussion of individual cases would be valuable. Previous studies have also noted that there are at least some negative and some non-significant associations, and understanding the underlying causes is of great interest (e.g. Kotiaho et al. Biology Letters).

Reviewer #3 (Public Review):
Summary:
This paper claims to overturn the longstanding abundance occupancy relationship.
Strengths:
(1) The above would be important if true. 
(2) The dataset is large.
Weaknesses:
(1) The authors are not really measuring the abundance-occupancy relationship (AOR). They are measuring abundance-range size. The AOR typically measures patches in a metapopulation, i.e. at a local scale. Range size is not an interchangeable notion with local occupancy.
(2) Ebird is a poor dataset for this. The sampling unit is non-standard. So abundance can at best be estimated by controlling for sampling effort. Comparisons across space are also likely to be highly heterogenous. They also threw out checklists in which abundances were too high to be estimated (reported as "X"). As evidence of the biases in using eBird for this pattern, the North American Breeding Bird Survey, a very similar taxonomic and geographic scope but with a consistent sampling protocol across space does show clear support for the AOR.
(3) In general, I wonder if a pattern demonstrated in thousands of data sets can be overturned by findings in one data set. It may be a big dataset but any biases in the dataset are repeated across all of those observations.
Overturning a major conclusion requires careful work. This paper did not rise to this level.

An illusion of a macroecological law, abundance-occupancy relationships

Figures and data

A conceptual overview of our methods.

Funnel plots.

The relationship between species average (mean) density and species range size.

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