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Open Access Publications from the University of California


Frontiers of Biogeography (FoB) is the scientific magazine of the International Biogeography Society (IBS,, a not-for-profit organization dedicated to promotion of and public understanding of the biogeographical sciences.  IBS launched FoB to provide an independent forum for biogeographical science, with the academic standards expected of a journal operated by and for an academic society.

Cover Caption: A couple of Scarabaeus laticollis rolling a brood ball near Monte Tiscali, Sardinia, Italy. See Milotić et al. article in this issue of Frontiers of Biogeography (10.1-2, e37289) for the results of a set of large-scale experiments of dung beetle ecosystem function throughout the Western Palearctic. Picture by Rafael Brix, available at under a CC BY-SA 3.0 license.

Research Letters

Where on Earth are the “tropics”?

The tropics have long been a focal point of interest in ecology and evolutionary biology - but where actually are the tropics? Classically, the tropics have been defined as all areas lying between 23.4° North and South as these zones receive direct overhead solar radiation at some point during the year. However, a suite of different environmental and climatic characteristics have also been employed to classify regions as tropical or not. The aims of this paper are to 1) briefly review some of the different criteria commonly used to define the tropics, 2) map the extent and distribution of tropical land areas according to these different criteria, and 3) asses the concordance between these criteria and a sample of recent “tropical” studies. More specifically, we review eight criteria that are frequently used (implicitly or explicitly) to define the tropics. We then map the location and extent of land areas that are “definitely tropical” (i.e., the core tropics) and areas that are “tropical by most definitions”. Finally, we examine how the different classifications apply to tropical research through an analysis of the study locations of >200 recent tropical biology articles. Depending on the definition, the extent of the terrestrial tropics ranges from 23 million to 66 million km2 – a nearly threefold difference. Likewise, the classification of many areas as being tropical vs. non-tropical depends on the specific criterion employed. Of the tropical studies reviewed here, only 44% were based on data collected from the core tropics, and 12% of tropical studies were based on data collected from sites outside of the geographic tropics but with tropical climates. Many different criteria are used to classify areas as tropical vs. non-tropical, leading to inconsistencies when estimating the extent of tropical areas and variation in the classification of ecosystems and species as being tropical vs. non-tropical.

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Frontiers of Biogeography: taking its place as a journal of choice for the publication of high quality biogeographical research articles

Executive summary. Through this editorial we seek your support and engagement as authors, readers and reviewers as we take the next steps in developing Frontiers of Biogeography as a leading international journal of biogeography and related subdisciplines. Here we make the case for submitting your next contribution to this journal: affordable, gold libre, open access, with the support of a disciplinarily-informed editorial and review team, which returns benefits to the biogeography community.

Research Articles

Changes in biodiversity and species associations along a latitudinal gradient

Biodiversity is currently threatened at local, regional, and global scales, and identifying the species that are vulnerable to these changes is essential for conservation efforts. For example, the breadth of species’ ranges may offer important clues to their susceptibility to loss, as widespread species may be more resistant to loss than species with a narrower range breadth. To determine the potential for shifts in community interactions along the latitudinal geographic ranges of species, we examined pair-wise associations between narrow-range endemic and widespread rocky intertidal species. We surveyed rocky intertidal species composition at eight sites along the California coast from San Diego to Cape Mendocino. Four sites were south of Point Conception, and four sites were north of Point Conception. Point Conception is a major biogeographic feature for coastal marine species, where sea surface temperatures transition from cool temperate waters in the north to warm temperate waters in the south. To determine whether pair-wise species associations were significant, we compared the observed communities’ standardized effect size to a null model to determine which species occurred together more or less often than by chance. Across all sites, widespread species were considerably more abundant than narrow-range endemic species, and the majority of species were widespread. However, total species richness was unrelated to the number of widespread species, and was, instead, determined by the number of narrow-range endemic species present at a site. Our analyses suggest that species are more aggregated than segregated south of Point Conception, but the opposite is true north of Point Conception. Additionally, we found that species associations between narrow-range endemics drove the overall patterns in species associations. One possible explanation for these patterns is that positive interactions, especially those involving narrow-range endemic species, are more important in southern California’s more thermally stressful intertidal habitats.

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Opinions, Perspectives & Reviews

The smartphone fallacy – when spatial data are reported at spatial scales finer than the organisms themselves

Thankfully, the days when specimen localities could be described in extremely vague terms such as “Peru” or “Indochina” are long gone. But the pendulum has swung too far the other way. Latitude and longitude data of specimens and study areas (such as small nature reserves) are nowadays commonly reported to the 0.000001 of a degree (or 0.01 of a second) or even more “precisely”. This is done either because of converting across measurement systems or because hand-held devices and internet sources provide this kind of precision. We probably report this degree of precision because we are reluctant to round – feeling it would make the data better and more “scientific”. I point out the scale referred to by different degrees of geographic precision (e.g., ~10cm for 6 decimal places) and argue that such degree of precision is false for two reasons: first, it is finer than actually achievable by hand held devices such as smartphones and GPS receivers (and much finer than we can tell from a map). Second, for large animals, such precision can refer to one part of the organism, and not another. I urge scientists to use simple reality checks when reporting latitude and longitude data and report precision at meaningful scales.


Dung beetle assemblages, dung removal and secondary seed dispersal: data from a large-scale, multi-site experiment in the Western Palaearctic

By manipulating faeces during feeding and breeding, dung beetles (Coleoptera: Scarabaeidae) fulfil important ecosystem functions in terrestrial ecosystems throughout the world. In a pan-European multi-site experiment (MSE), we estimated the ecosystem functions of dung removal and secondary seed dispersal by differing combinations of dung beetle functional groups. Therefore, we classified dung beetles into five functional groups according to their body size and dung manipulation method: dwellers, large and small tunnelers, and large and small rollers. Furthermore, we set up a dung beetle sampling database containing all sampled dung beetles during the project. By identifying dung beetle specimens to the species level, we obtained a detailed insight into the dung beetle communities at each study location.

By establishing experimental plots allowing and inhibiting specific combinations of functional groups in the local dung beetle assemblage from removing dung and seeds, we estimated the role of each group in dung removal and secondary seed dispersal during a 4-week period. We performed all experiments in grazed (semi-)natural grasslands, and used different dung types (cattle, horse, sheep, goat or red deer) to match the herbivore species grazing in close vicinity of each of the study areas. Simultaneously, we sampled dung beetle assemblages by using pitfalls baited with the same dung types as used in the experiments.

This data paper documents two datasets collected in the framework of this MSE project. All the experiments took place between 2013 and 2016 at 17 study sites in 10 countries and 11 biogeographic zones. The entire dung beetle sampling dataset was published as a sampling event dataset at GBIF. The dataset includes the sampling results of all 17 study sites, which contain 1,050 sampling events and 4,362 occurrence records of 94 species. The second dataset contains the results of the dung removal and secondary seed dispersal experiments in which we used 11 experimental treatments and the five dung types mentioned above. This experimental results dataset holds all experimental results of the MSE project (11,537 records), and was published in the online data repository Zenodo.


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