ECR feature: Vicente García-Navas on diversification in dasyurid marsupials

Vicente is an evolutionary ecologist and is currently a postdoc at the University of Zurich. He is largely interested in patterns of lineage diversity and eco-morphologies. Vicente shares his recent work on the interplay between competition, divergence time, and geographic range overlap on the diversification of dasyurid marsupials in Australia.

Name. Vicente García-Navas

Personal links. Twitter | ResearchGate | Website

Institute. University of Zurich, Switzerland (UZH)

Academic life stage. Postdoc

Major research interests. My ongoing research lies at the interface between ecology and evolution, addressing how temporal and spatial patterns of eco-morphological and lineage diversity are influenced by biotic and abiotic factors and their interactions. I try to incorporate contemporary (local) processes into “tree thinking” allowing joint testing of ecological and evolutionary processes. I am interested in how phenotypic traits evolve, how species richness accumulates, how the rate of evolution differs among lineages and over evolutionary time, and the mechanisms driving these varying rates of change.

Pictures of Triodia (spinifex) clumps and sandy substrates along the Canning Stock route in Western Australia. Triodia is a large genus of hummock-forming bunchgrass endemic to Australia. Triodia hummocks constitute the main habitat of some Sminthopsis species like the sandhill dunnart S. psammophila (pictures: Mike Westerman).

Current study system. I use vertebrate radiations as model systems with special emphasis on taxonomic groups that diversified in Australia and nearby islands. This island-continent constitutes a very suitable scenario for studying diversification dynamics at a regional scale. Marsupials account for over half of Australia’s land mammals and are well represented in the totality of environments that can be found across this continent. Consequently, this taxonomic group has prompted a large body of research. However, little is known about the role of biotic interactions in shaping coexistence patterns in marsupials. There is paucity of studies focused on interspecific competition among members of this speciose group.

Recent publication in Journal of Biogeography. García-Navas, V., Kear, B.P., Westerman, M. (2020) The geography of speciation in dasyurid marsupials. Journal of Biogeographydoi/abs/10.1111/jbi.13852 [Access here]

Motivation behind this work. Unlike other more striking and iconic Australasian radiations, dasyurid marsupials have received relatively little attention. This lack of research might (at least partially) be due to low morphological specialization in this mammalian taxon, which makes it less intuitively appealing for diversification studies. However, we have previously showed that patterns of both phenotypic disparity and speciation in dasyurids exhibited an early burst followed by a slowdown. It is likely that the extinction of thylacinids and the spread of arid habitats spurred this radiation of insecto-carnivorous mammals giving rise to that early burst signal. In this paper, we were interested in the effects of competition and divergence time on morphological dissimilarity (in terms of body mass and molar row length) and geographic range overlap between species at different scales. Most community phylogenetic studies do not consider biogeographic history and regional-scale processes despite growing recognition that these influence contemporary community patterns. Consequently, incorporating perspectives on clade-level evolution into studies of community assembly constitutes a fundamental challenge for the progress of this field.

One of the most emblematic Australian species; two Tasmanian devils Sarcophilus harrisii at Traunna Wildlife Park in Tasmania (picture: Mike Westerman).

Key methodologies. We examined the relationship between species co-occurrence patterns and morphological similarity at two spatial scales. Although local and regional scale analyses should ideally be applied in tandem, few studies have addressed both perspectives simultaneously. Specifically, we first tested whether speciation in this group has been largely allopatric at regional scale, as previously shown in other mammalian families. Secondly, where species coexist, we tested whether this sympatry might have been facilitated by morphological divergence at local scale through character displacement. For this purpose, we used a novel approach developed by M. Borregaard, node-based analysis (, which quantifies the distributional divergence between daughter lineages descending from the same node and allows obtaining a better insight of the phylogenetic structure of assemblages at large scale. At local scale, we used co-occurrence data from 83 communities in order to test whether co-existing species are more morphologically divergent that those that do not coexist or do so not preferentially. 

Major results. Our study reveals that geographic isolation arising from niche conservatism (as opposed to biotic interactions including competitive exclusion) has played a pivotal role in shaping the speciation patterns of this endemic mammal radiation. The level of sympatry observed in our sample of sister species pairs was slightly higher than that generally reported amongst placental mammals. However, despite moderate levels of sympatry through time, our results indicate low rates of spatial co-occurrence between dasyurids. It supports the idea that, under certain circumstances a high degree of range overlap may not translate into real coexistence at a local scale. That is, in many cases sympatric species (in terms of broad-scale spatial overlap) are not syntopic and do not interact ecologically. By integrating approaches used to infer broad-scale, evolutionary processes and those commonly used to study ecological interactions at fine scales, we show that it is possible to obtain a comprehensive understanding of factors driving species distributions.

Unexpected challenges. I remember writing this paper while recovering from a surgery. So, I have a folder in my laptop devoted to this study called “dasyurid speciation” but, for me, this paper will always be the gallbladder paper. There is no better plan for the summer than to stay at home writing while your scars dry and doctors remove the stitches. Apart from this, the process was quite smooth. What took us the longest was to collect data on length of the lower molar row for all species. In several cases, we had to dig into grey literature to find morphological data for some poorly-known taxa (e.g. Antechinus) or request data from experts, which indicates that even in some charismatic groups like marsupials there is still a need for more natural history studies. We still lack basic information on morphology and ecology for a high percentage of mammalian species. Consequently, there is no doubt that fieldwork is still crucial for science research.  

A Northern quoll, Dasyurus hallucatus, also known as the northen native cat, in Darwin (NT). This species was first described in 1842 by the English ornithologist John Gould (picture: Mike Westerman).

Next steps of this research. This study is part of a project on diversification and phenotypic evolution in vertebrate Australasian radiations. One of the main aims of this project is trying to reconcile macroevolutionary processes acting at a continental scale with ecological processes, like competition, that are generally interpreted at smaller spatial scales. We plan to continue interpreting local coexistence patterns in light of historical processes, taking advantage of traditional field surveys and state-of-the-art techniques. We are interested in examining how local assemblages are structured over time by explicitly considering range dynamics, speciation, colonization, and local extinction rates. In this way, we will be able to ask if there is a link between diversification rates and species co-occurrence.  

If you could study any organism on Earth, what would it be? Certainly no one will find me chasing raptors or large ungulates. I don’t know why, but I have a strong preference for rodents (and rodent-like marsupials) and small birds. So, I would definitely continue studying songbirds and small mammals. They are probably easier to study than other organisms, which allows obtaining large sample sizes and even carrying out experiments in the field. In addition, they represent some of the most striking vertebrate radiations and have successfully colonized a great variety of environments exhibiting a high variability in terms of morphological and/or behavioural adaptations. Beyond this, I think it goes without saying that they are amazing creatures and, most of them, overwhelmingly cute.

Anything else you would like to share?

I did my PhD on behaviour in blue tits and then I started to apply molecular tools to address questions at a larger scale (e.g. gene flow and local adaptation between colonies). The need to answer broader questions pushed me to move from working at the individual or population level to adopt a more comprehensive approach, one that considers macroevolution using modern phylogenetic comparative methods. Thus, I have progressively expanded my interests from individual decisions and taxon-specific questions to broad-scale processes that act as biodiversity engines. I think that this winding path has provided me with a background that allows me to approach my research with a broader and more integrative perspective.

Regarding my little obsession with the Australian fauna, I remember that I enjoyed a lot watching “Taz-mania”, an animated sitcom produced by Warner Bros and starring Taz, the Tasmanian Devil, when I was a kid. This “Looney Tunes” character was popular enough for the potential of a spin-off series focused on his adventures and those of other furry creatures including wallabies, koalas and dingoes. I really think cartoons have great potential to convey respect and interest for animals!

ECR feature: Edmund Basham on tree-top frogs

Edmund Basham is a community ecologist and biogeographer who is currently studying toward his PhD at the University of Florida. He has deep love and interest in the frog communities of tropical rainforests. Edmund shares his recent research on the seasonal shifts and vertical stratification of tree-dwelling frog species in Panama.

Personal links. Instagram | Website

Institute. University of Florida

Academic life stage. PhD

Major research themes and interests. Amphibian community ecology, vertical stratification, biogeography, climate change.

Current study system. I am interested in the vast diversity of species that make up the amphibian communities found in the rainforests of Central Panama. Such species include fossorial caecilians that are found below the leaf litter, to the rare frogs that breed in tree holes found high in the canopy. Some of the trees at the site are upward of 50 m tall, and finding frogs at this height always gives a rush of wonder and excitement. Talking to non-ecologists about my research, a regular comment is an incredulous “do frogs really live up in tops of trees?!”. This is partially what I think makes this research so interesting, there is a hidden world above our heads that often goes amiss.

Recent paper in JBI. Basham, E. W. and Scheffers, B. R. 2020. Vertical stratification collapses under seasonal shifts in climate. Journal of Biogeography: 1–11. [Access here]

A Sylvias tree frog (Cruziohyla sylviae), found 23 m up sitting close to a tree hole filled with tadpoles and and overhanging eggs ready to drop in.

Study motivation. The canopies of rainforests still hold so many secrets. I wanted to be one of the few researchers who truly sampled across the whole range of habitats within the forest, including across the year to sample seasonal climatic variation. Looking at how animals respond to seasonal climates can also give us information about how climate change could affect them, which was a major motivator of this study. Furthermore, there is scant information on the lives of canopy dwelling frogs, so collecting data on where and how these species live is valuable in its own right.

Key methodologies. This paper is one of the very few amphibian focused papers that incorporates arborist tree access. To sample amphibians high in the tree tops, we would shoot a lead weight and line into the canopy, hooking it over a branch so that we could haul up a climbing rope. I would then climb up at night, searching for frogs on the leaves, moss, vines, epiphytes, and all the microhabitats occurring along the vertical transect. This meant that I could find many illusive species that may be missed during ground surveys. Indeed, many of the largest and most beautiful frogs are the rare tree frogs found only in the canopy.

Edmund ascending a huge Espavé (Anacardium excelsum) tree during a daytime survey, on the hunt for frogs.

Major results. We found that the species that normally live up into the canopy, descend towards the ground in the dry season. One striking example was the yellow-bellied poison frog, which shifted some 25 m from its home in the canopy down to the roots at ground level. The overall downward community shift left a canopy depauperate of frogs, where only a small number of individuals from select species are found toughing out the heat in the dry season. With frogs forced to live in a smaller area of habitat near the ground, there may be greater risk of disease, competition, and predation from ground-based hunters such as snakes.

Most importantly, climate change is set to lengthen and strengthen seasonal droughts across the tropics. This research suggests a change is coming in the vertical organisation of frogs in tropical forests, with species that require above-ground habitat to live and reproduce losing out.

A Rosenberg’s gladiator frog (Hypsiboas rosenbergi), found 12 m up calling from a large palm. 

A Palmers tree frog (Hyloscirtus palmeri), found 5 m up calling during a rain storm.

The very illusive tree-hole breeding Ecnomiohyla miliaria, the fringe-limbed treefrog, listed as Vulnerable on the IUCN Redlist. Found near the ground during the dry season in Sierra Llorona, Panama.

Challenges. Conducting field work in mountainous tropical forests is challenging, but when you add a climbing component it can become extra tough. Bullet ants and eyelash pit vipers abound, not to mention the physicality of climbing huge trees. Nonetheless, I wouldn’t have it any other way. The challenging nature of this field work means that there is still so much to explore and learn about these canopy environments. For example, I found that one species of poison frog only lived on the biggest wild cashew trees at the site in Central Panama, which led me to target this interesting relationship for further surveys.

I would implore budding scientists to immerse themselves in nature, because the best inspiration often comes from witnessing something that makes you scratch your head and wonder “what is going on?”.

Next steps. To fully understand the generality of these patterns across the tropics, further field seasons using the same method need to be conducted in forests representative of major ecoregions, for example, South-East Asia, Central Africa, and India. The communities of amphibians in these far-removed forests are only distantly related, for example, frogs in Madagascar have been isolated for ~80 million years. Thus, the processes of evolution and selection may have shaped communities to react to seasonal climate in unique ways. Equally, we may find that a downward descent during dry seasons is a convergent mechanism adopted by all communities. This would then suggest that climate change will affect communities similarly across the tropics. To answer these questions, we must climb more trees and find more frogs.

If you could study any organism on Earth, what would it be? Frogs, especially the colourful tree-frogs. They exist in such incredible variety and beauty, and they have an innocence and charisma which I think is so endearing (for the most part!).

Anything else to share? Only that all are welcome to contact me, whether it be to discuss science, have a friendly chat, or form ideas for future research collaborations.

If you want to see some of this tree climbing/frog catching/fun having field work, check out this short video I made about the research!

Small grants for global colloquia in biogeography

The Journal of Biogeography is pleased to announce the second of three new opportunities.

The Journal of Biogeography invites applications for funding to facilitate one or more global colloquia.  The event may be stand-alone, or may be staged in association with a larger meeting, it may be in-person or virtual. The topic may be on any aspect of biogeography.  A goal of the colloquium should be to publish a synthesis paper and/or a series of papers that represent the range of topics discussed.

The Journal wishes these colloquia to become a regular activity that helps biogeographers develop, exchange, and explore ideas globally that advance biogeography through consolidation of fragmentary knowledge, synthesis across disciplines, and innovation.  Thus, the funds up to $2000 are offered by the journal primarily with the intention of facilitating involvement of people who might not otherwise be able to participate, or to stage an event that, because of its nature, draws in people of diverse backgrounds and with varied perspectives.   

The awardees will take care of all organization and are responsible to the journal only in terms of meeting any prior agreement on publication, promotion, and staging the proposed event.  The journal is responsible to the local committee only in terms of promoting the event via journal social media and providing the funding agreed. 

Proposals should take the following format (as a single PDF):

1. Title and description of the topical focus (e.g., early career conference, a regional focus, or a disciplinary focus). ≤0.5 page.

2. Relevance, context, and how the colloquium and publications therefrom will advance the discipline of biogeography. ≤0.5 page.

3. The target number of attendees, and distribution across career stages, countries of origin/habitation, gender. ≤0.5 page.

4. The proposed format: general organization (e.g. number of keynote speakers in plenary sessions, number of concurrent sessions, talks, posters). ≤0.5 page.

5. The actual dates and details of the colloquium: a) in-person/virtual, b) facilities/technology, c) list of organizers, d) list of committed participants. ≤0.5 page.

6. The uses of and substantial difference that will be made by the support from Journal of Biogeography. ≤0.5 page.

7. Outline of the proposed publications, which will first be submitted for consideration at Journal of Biogeography. ≤0.5 page.

8. Other budgetary considerations, partners, and obligations therein.

We aim to fund 1-2 symposia during the coming year.  Proposals on any subject in biogeography are welcome; in 2020, we encourage, but do not limit responses to this request for proposals, to colloquia exploring the following areas: Biodiversity–geodiversity; Comparative phylogeography and geo-genomics; Functional biogeography; Cross-scale biogeography & biodiversity (considering biological, spatial, and/or temporal hierarchies); Marine-terrestrial comparisons and contrasts (also with aerial, freshwater, and subterranean realms); Biogeography in the Anthropocene.

Submit proposals to: *upload only*

The deadline for submissions is: 01 November 2020.  

Address enquiries (Subject line: “Enquiry: Journal of Biogeography Colloquia sponsorship”) to the Editor-in-Chief at

JBI aims to foster inclusive science that reflects the disciplinary, human, and geographic diversity of biogeography and biogeographers. Submissions are welcomed from applicants of all ethnicities, races, colors, religions, sexes, sexual orientations, gender identities, national origins, disabilities, ages, or other individual status.

ECR: Jordan A. Hollarsmith

Jordan is a postdoc at Simon Fraser University. She is an ecologist with an interest in resource management. Jordan shares how she and her colleagues have used a remotely operated vehicle to survey marine biodiversity in the Mexican Pacific.

Jordan Hollarsmith.

Personal links. Website | Twitter

Institute. Simon Fraser University

Academic life stage. Postdoc

Major research interests. Resource management and ecology as it relates to multiple stressors.

Current study system. My research currently focuses on how we manage resources given the cocktail of threats facing nearshore marine ecosystems. In addition to comparing decision-making frameworks used across North America, I am using kelp forest ecosystems of the Salish Sea as a case study into modelling threats using expert opinion instead of more traditional data sources. This work is beautifully collaborative and very urgent given the pace of ecosystem decline we are observing in the Salish Sea and around the world.

Jordan Hollarsmith, Kyle Neumann, and Tallulah Winquist, prepare for a dive in large swell of the coast of Socorro Island in the Revillagigedo Archipelago. Photo taken by Tamara Arenovich with the Sea Shepherd Conservation Society.

Jordan Hollarsmith and Georgina Ramírez Ortiz with the ROV. The photo was taken by Arturo Bocos at El Bajo in the Bay of La Paz.

Recent paper. Hollarsmith JA, Ramírez-Ortiz G, Winquist T, Velasco-Lozano M, DuBois K, Reyes-Bonilla H, Neumann KC, Grosholz ED. 2020. Habitats and fish communities at mesophotic depths in the Mexican Pacific. Journal of Biogeography 47(7), [content link here]

Motivation for this paper? Ecosystems found below 30 m depth in the ocean are some of the least known in the world, in part because it is so difficult to access them. However, advances in ROV (remotely operated vehicle) technology means they are now smaller and cheaper than ever before. Colleagues and I decided to seize this opportunity. With the help of National Geographic, the Explorers’ Club, the Sea Shepherd Conservation Society, and other generous partners, we built our own small ROV to survey continental and oceanic islands in the Mexican Pacific. We chose these locations based on the high biodiversity and endemism found in shallow areas around the islands. Moreover, the complex mixing of water masses in the region provides increased potential for highly diverse habitats and fish communities below 30m.

Key methodologies. This paper is among the first to use small and economic ROV technology to survey fish communities at mesophotic depths. Our approach was highly collaborative, involving early career researchers and students from the United States and Mexico, multiple academic institutions in both countries, non-profits, funding sources, and direct-action environmental organizations. Thanks to the combination of diverse collaborations and affordable technology, we were able survey a wide range of islands, depths, and habitats for very little money.

The ROV.

Unexpected challenges. Driving the ROV from small boats in huge open-ocean swell under a stifling tarp surrounded by curious boobies and sharks was difficult, to say the least! It was hard to see many details when driving the ROV, so often we would only realize what we had surveyed when we reviewed the footage later. Thankfully, despite the difficult field conditions and the constant battle with sea sickness, we were able to identify the majority of fish we observed and elucidate ecological and biogeographical patterns from our data.

A soft coral reef at mesophotic depths around the Revillagigedo Archipelago.

Major results. Out of the 81 species we identified in our surveys, we observed nine fish species deeper than they’ve ever been recorded and one fish species, the Galapagos snake eel, thousands of kilometers away from anywhere it had previously been observed. Our surveys included large undocumented rhodolith beds (free-living coralline algae) and mesophotic algal communities, in addition to diverse communities of soft corals and sponges. These findings increase our knowledge of the natural history of these fish species, and increase our understanding of the ecosystems within the two Mexican National Parks that we surveyed.

A mesmerizing school of hunting jurel (Seriola rivoliana) at Clarion Island.

Next steps for this research. One of the undergraduate students involved in the video analysis is using these data for his senior thesis at the Universidad Autónoma de Baja California Sur. Watch out for his paper on the differences in fish functional diversity at mesophotic depths across continental and oceanic islands, currently in revision at Ciencias Marinas!

If you could study any organism on Earth, what would it be? I would study kelp! Kelp is a beautiful and diverse family of brown algae (Laminariaceae) that forms critical habitat in temperate rocky reefs around the world. Kelp is also found in the mesophotic zone in tropical and subtropical regions where a clear surface layer allows light to penetrate deep into the ocean where it illuminates cold, nutrient-rich water, thus providing the perfect conditions for kelp. These habitats have been observed in the Galapagos, New Zealand, and the Mediterranean. We were hoping to find another one in the Revillagigedo Archipelago, but we will have to save that for another mission!

How did ant communities assemble on reservoir islands?

From curiosity to community assembly: how a birder’s frustration opened the door to a new journey of discovery about taxonomic, functional, and phylogenetic diversity of ants on islands.

When I started my Ph.D. career and went to the Thousand Island Lake (TIL) to do fieldwork for the first time in 2014, I was attracted to the natural landscape immediately. Before then, I had never been to any reservoir islands. TIL was formed by dam construction in 1959, and is one of the largest human-made reservoirs in China. The lake area is approximately 580 km2 and includes more than 1,000 islands (as its name suggests). Each island was previously a hilltop before flooding.

Above. Thousand Island Lake. Photo credit: Wande Li.

EDITORS’ CHOICE (Read for free until Aug 2022): Zhao, Y,  Dunn, RR,  Zhou, H,  Si, X,  Ding, P.  Island area, not isolation, drives taxonomic, phylogenetic and functional diversity of ants on land‐bridge islands. J Biogeogr.  2020; 00: 1– 11.

Since 2003, a research group led by Prof. Ping Ding at Zhejiang University has been conducting bird surveys in TIL. I was trained as a “birder” too. However, I was surveying birds on a tiny island one day, and was frustrated that there seemed to be no birds to watch on that island. As I walked around searching for birds, I noticed that the island was covered by a massive number of black ants (years later I would realize that this black ant was Brachyponera chinensis, known as Asian needle ants), these ants walked all around me, and I stared at them for a long time just like a little child will do. That was the very moment I wondered how many and which ants can be found on each land-bridge island, and why.

I returned home and began to search for articles about ants. The more I learned, the more I was attracted by these little creatures. I realized that many ecologists around the world had used ants as the target taxon. Among them, Edward O. Wilson, who was inspired by his travels and ant collecting in the South Pacific, together with Robert H. MacArthur to develop one of the most iconic theories in ecology: the Theory of Island Biogeography (TIB). TIB predicts that larger and less isolated islands should host a larger number of species than smaller and more isolated islands. However, the classic TIB model does not consider the evolutionary and functional differences among species (i.e., phylogenetic diversity and functional diversity). I began to think that there was a project in the making with ants, maybe even a Ph.D. I could, I began to think, study not only the number of kinds of ants on the different islands, but also their evolutionary and functional diversity.

In 2017, after a discussion with my supervisor Prof. Ping Ding, along with other co-authors, we decided to survey ants on different islands in TIL using different capture methods. Meanwhile, we also planned to sequence the DNA and measure the functional traits of all ant species to infer the phylogenetic and functional diversity of ants.

Different capture methods: Winkler extraction (left), pitfall traps (upper right), and hand collection (lower right). Photo credit: Yuhao Zhao.

After two years of sampling, we captured over 90,000 individual ants across 33 islands, belonging to 97 species in 44 genera and eight subfamilies. The counting process was daunting as sometimes a trap can collect thousands of ant individuals, and we need to count them one by one. However, the results were promising. We found larger islands possessed more species and higher phylogenetic and functional diversity than smaller islands, which is consistent with the prediction from the classic theory of island biogeography. However, isolation did not significantly influence the taxonomic, phylogenetic, and functional diversity of ants.

To move a further step, we wanted to know if the underlying processes of community assembly on different islands are the same or not? Based on previous research of bird studies in the same system (Si et al. 2017, link:, we hypothesized that communities on smaller and more remote islands might be highly clustered (with more closely related ant species than expected by chance) because the subset of species either shares similar functional traits or similar evolutionary histories. Conversely, communities on larger and less isolated islands may be overdispersed (with fewer closely related ants than expected by chance) because these islands provide more potential habitats and niches, and closely related species would compete for resources, resulting in close relatives being competitively excluded from the community. Using a null modelling approach, we found that the structure of ant communities shifted from phylogenetic and functional clustering on smaller islands to phylogenetic and functional overdispersion on larger islands. That is to say, environmental filtering is the dominant process structuring ant communities on smaller islands, and that competitive exclusion becomes more important on larger islands.

More interestingly, we also found incongruencies between phylogenetic and functional dispersion patterns. Overall, phylogenetic structure of ant communities tended to be clustered, whereas functional structure of ant communities tended to be overdispersed. These results suggest that distinct mechanisms may influence phylogenetic and functional structure of ant communities, i.e., environmental filtering influences phylogenetic community structure of ants, whereas competitive exclusion influences functional community structure of ants. Our findings, again, highlight the need to examine both phylogenetic and functional diversity to understand the mechanisms that govern the assembly of natural communities on islands.

The Asian needle ants (Brachyponera chinensis) attacking a termite. Photo credit:Benoit Guénard.

During our ant sampling in the field, we inevitably noticed the interaction between ants and termites, as well as ants and aphids, and ants and plants. Although these interactions have been well documented, few works have investigated the variations of these interactions among islands. Based on our knowledge of ants in the TIL region, we now have multiple projects to understand the influence of ant on other species, and the functional roles of ants in the land-bridge island ecosystem.

Written by:
Yuhao Zhao – Postdoctoral fellow, School of Ecological and Environmental Sciences, East China Normal University.

Additional information: ; ;

I would like to thank Xingfeng Si and Rob Dunn, who provided feedback on this blog post, and Wande Li and Benoit Guénard to provide the photos.

Figures: the Art of Science

How to prepare figures to make an impression on editors, reviewers, and readers.

Figures are, perhaps not quite literally, worth a thousand words, but they are invaluable: try explaining in text all the details in anything but the most basic image. Yet a reasonable fraction of manuscripts submitted to Journal of Biogeography (JBI) contain too few or too many figures, figures that are difficult to interpret, figures that are poor quality, incorrectly sized, that don’t conform to JBI’s style requirements, are formulaic, and/or don’t convey the research in a compelling way. These are all things that can slow down review or publication of a manuscript or increase the likelihood it will be rejected, especially if there are also shortcomings in the text, so best to avoid them if at all possible.

By contrast, an appropriate number of nicely prepared, easily interpretable, information rich figures will emphasize the positive and can to some extent compensate for shortcomings elsewhere during review, because a series of good figures can alone tell most of the story in a paper, and reviewers and editors consider manuscripts holistically. (This is not to say other things will be ignored; to ensure the best possible chance of publication, all aspects of a paper should be prepared to publishable standard before submission.)

Moreover, creative and thoughtful illustrations will make your paper stand out from the crowd during review, when being considered for the Editors’ Choice or Cover Article, and when readers browse the journal or find your paper among tens of others in a literature search. As such, consistent with this column’s intention to help authors’ manuscripts be more successful at JBI, we here provide a primer on preparing figures for successful review and publication in JBI. This is intended as a quick reference guide to some key ‘good practices’ in preparing your figures and so it emphasizes aspects of, but does not replace, information provided in the journal’s author guidelines and Wiley instructions. We hope this is helpful!

1. Include a conceptual or synthesizing figure
We are all familiar with using the Introduction of a paper to establish the background for our study, the questions that background raises or leaves unanswered, and so the hypotheses our study is going to test. But far less often do we also present that information in a powerful visual graphic. Yet such graphics can quickly provide readers with an holistic view of your study and a visual expectation of what your data would look like if it was consistent with or refuted your hypotheses, which they can then pattern-match to your actual results. This approach can therefore also guide how you set up your Results and can simplify your Discussion. In a way, it’s really an element of good study design, showing the study has been thought all the way through from the start. An alternative (or addition), if you wish to provide a powerful take-home message that others will use in their talks and redraw in their papers, is to provide an integrative synthetic figure emphasizing the key points in the Discussion. Similarly, it has been noted that if your methods are complex, it can help to have a (numbered) flow-diagram which can both clarify and shorten your textual description..

2. Figure size and proportions
Create your figures at the size and proportions they’ll be printed. Figures are reproduced in the final article in three basic sizes: single column width (79 mm), two-thirds page width (110 mm), and full page width (168 mm). If a figure is designed for one of these sizes it’s usually obvious and the production office will make it look pretty. If it is not designed for one of these sizes, it will be shrunk or expanded leading to images with detail that is hard to read — in production, figures will be smaller than R output, so letters have to be proportionally bigger, lines thicker, and points larger to be read appropriately — or an overly simplistic and large kindergarten book kind of feel.

All graphics software these days allows sizes to be set and/or images scaled, so when designing figures, keep the final sizes in mind, and imagine what they’ll look like on the ‘printed’ page, either one full-page or fit-to-width view online or in a PDF; if you’re relying on people zooming in/out, you probably need another panel or figure. If you’ve multiple panels, consider how to arrange them so they fit down one column, or across the page, or in an # rows x # columns composite figure; if your panels are graphs and they have an axis in common, use that as your guide. Try to avoid arrangements or sizing that will lead to a block of empty ‘white space’ around the figure. Make sure to scale the text on your page to fit the final printed size (ideally use 8 pt after any reduction), don’t have too many different point sizes in one figure, and try to keep text size consistent across figures in your manuscript. N.B. The maximum height of figure is 230mm.

3. Resolution
Use vector graphics, if at all possible, which will ensure your image looks sharp at either one full-page or fit-to-width view on the screen and everything in-between and beyond. Vector graphics should be in .eps or .pdf format. If using photographic images save them in .tif format at 300 d.p.i. If figures combine line art and photographic images and cannot be saved in vector graphics format, use .tif format at high resolution (i.e. 600–800 d.p.i.).

4. Figure captions
Captions, a.k.a. legends, should be concise, comprehensive, and ‘stand-alone’ – i.e. the figure and its caption must be understandable without reference to the text. To this end both the geographical region/s and the taxon/taxa should be mentioned in each caption. Include definitions of any symbols, abbreviations, and units of measurement.

5. Colour
Use colour wisely. As colour images are free in the PDF and online (which is where everyone is going to read them) it may be tempting to use colour often and sometimes a lot of it. But unless colour actively increases readability of your figure (i.e. in figures with many components to be compared, or when >2 dimensions are needed) think twice about using it. Also, avoid heavily coloured base maps (e.g. captured from Google Earth) are a poor background on which to plot important data because they have a lot of superfluous information and don’t allow for sufficient contrast. Often it can be equally or more effective to combine different types of symbols (e.g. solid and dash lines or areas, or round versus square points) than adding many colors. This kind of presentation also works on a grey-scale printed page and for the many people who have different perceptions of colour. Colour figures should be saved in CYMK

6. Maps
(a) Include a map that provides regional context as well as details of your study. JBI has a global readership so many people might not immediately recognize your study area without broader context. Provide details of your study in your main figure and consider also an inset or additional panel(s) showing country-or-higher level context. In each panel include a scale bar in ‘km’ within the figure.
(b) Continental and global maps should usually use an equal area projection. All biogeography addresses life on an approximately spherical planet, but we draw them on a flat page. Much biogeography addresses processes influenced by area and/or distance, so patterns are best represented on a map that reproduces these attributes with as much fidelity as possible. For area, Mollweide or Aitoff’s projections are recommended choices. If distance is more important than area in your study perhaps choose an equidistant map instead (one pertinent to your area of study). If dealing with area and distance, consider different projections for different maps within the same study or a ‘compromise’ map. Regardless, maps based on, e.g. the Mercator projection provide misleading visual cues close-to or far-away from the equator and should not be used. Given the variety of maps and that it can be difficult to distinguish between some visually, it is important to state the projection employed and the reason (e.g. “equidistant” or “equal area”) in the figure caption. Two lists and descriptions of projections are available at and
(c) Show the range of the species with which you’re working as well as the extent sampled if these are not the same. There is good reason to expect that biogeographic processes vary spatially and in regard to the relative position within a species range (e.g. periphery versus center, ‘leading’ versus ‘trailing’ edge), so to the extent it may be relevant to the goals of a study, sampling location(s) should be shown relative to the species’ range.
(d) An additional consideration. It can be more convenient to summarize sample sizes and other relevant background information in a map than in the Methods text; too often papers do neither and bury this important information in supplementary documentation.

7. Phylogenies
Must include edge/node support values. A list of names in the leaves is not always most helpful for readers. Consider complementing the basic tree with clade-level annotation, images, maps, and traits.

8. A few final tips from the Production Office
Ensure (i) all figures are cited, in order, in the main text of your article; (ii) each individual figure file less than 10 MB, and if not then remove excess white space surrounding figures for smaller file sizes; (iii) figure files are named with their figure number. A checklist for basic figure requirements is here for initial peer review as well as production.

A few examples:

Effective single column black and white figures

Nice map showing global location, regional sampling, and detailed study design

Good use of a conceptual figure

Effective arrangement in 1r * 3c to facilitate comparison of species richness; also appropriate use for colour (Fig. 2)

A variety of stacked, 2/3rds width, and multi-panel figures that also support adjacent placement of captions

Some additional resources:

McInerny et al. 2014. Information visualisation for science and policy: engaging users and avoiding bias. Trends in Ecology & Evolution29:148-157

ECR feature: Himalayan wolves with Geraldine Werhahn

Geraldine Werhahn is a research associate at Oxford University. She is a conversation biologist who uses a multidisciplinary approach to study the ecology and evolution of carnivores in the Himalayas and the Tibetan Plateau. Geraldine shares her recent work on the taxonomy and adaptive evolution of the Himalayan wolf.

Geraldine in the Himalayas, collecting wolf scat for population genetic analyses (Photo credit: Naresh Kusi).

Personal links. Twitter | Instagram

Institute. Wildlife Conservation Research Unit (WildCRU), Department of Zoology, Oxford University, UK

Academic life stage. Research associate.

Research themes and interests. My research focuses on the conservation and ecology of carnivores in the high-altitude ecosystems of the Himalayas and the Tibetan Plateau of Asia. My work is driven by the need to maintain healthy carnivore populations and conserving ecosystem integrity. I take a multidisciplinary research approach (from genetics, to ecology, to social science) to understand my study system and develop science-based conservation strategies.

Current study system. My current study system is the Himalayan wolves and coexisting carnivores in the high altitudes of Asia. These high-altitude ecosystems are vast wilderness regions, also sometimes called the third pole, where the harsh climate and low oxygen levels shape a unique fauna and flora. The Himalayan wolf is found in habitats above 4000 m elevation and is unique to the Himalayas and the Tibetan Plateau. As a top predator alongside the Snow leopard it plays an important role in ecosystem function, warranting its conservation.

A Himalayan wolf pup in the Transhimalayas of Nepal (Photo credit: Geraldine Werhahn).

A herd of Kiang (Equus kiang) in the Transhimalayas of Nepal. These wild equines share their habitat with the Himalayan wolf (Photo credit: Geraldine Werhahn).

Recent paper in Journal of Biogeography. Werhahn et al. (2020) Himalayan wolf distribution and admixture based on multiple genetic markers. Journal of Biogeography. 47(6): 1272–1285.

Motivation for this recent paper. The motivation for this research was to investigate the genetics and ecology of the Himalayan wolf to inform its pending taxonomic classification and advance its conservation. Prior to our work, little research had been conducted on the Himalayan wolf. Most data were derived from captive animals at zoological gardens, or from museum collections, and a range of different names were used for this one wolf lineage across different publications. We focused especially on the evolutionary history, foraging ecology, and distribution of the Himalayan wolf. We wanted to determine if the Himalayan wolf was a distinct lineage that occurs alongside the Holarctic grey wolf, and to explore its phylogeny in relation to contemporary canids from around the globe. For conservation purposes, we also investigated whether hybridization occurs between the Himalayan wolf, the grey wolf, and domestic dogs.

Key methodologies. We developed a conservation genetics toolkit specifically tailored for non-invasive scat sampling of the Himalayan wolf. Our work was conducted in close collaboration with our genetics partner, the RZSS WildGenes lab in Edinburgh. We worked with non-invasive samples obtained from scat. Thereby we could cover many packs and individuals without interfering with the animals. Our methods allowed us to identify wolf individuals and determine their phylogenetic relationship with other canids. We also searched for genetic differences at functional genes known to be involved in adaptation to low environmental oxygen levels. Such genetic adaptations to low oxygen have been described for Tibetan mastiff dogs (the typical dog breed of the Tibetan Plateau) and human populations on the Plateau. So, we were interested to investigate if such adaptations might be important in the ecology of the Himalayan wolf. We analysed 280 wolf scat samples collected from the Tibetan Plateau in western China, and the Central Asian mountains in Kyrgyzstan and Tajikistan, and we included canid reference sequences from around the globe to build the phylogenies based on mitochondrial DNA sequences.

Major results. The Himalayan wolf forms a genetic clade that is distinct from other canids based on the tested mitochondrial and nuclear markers. We found consistent differentiation and unique alleles in the Himalayan wolf samples at functional genes known to be involved in adaptation to low oxygen levels (the hypoxia pathway).  

The distinctness of the Himalayan wolf may be related to the uplifting of the Himalayas and the Tibetan Plateau that produced a unique ecological niche in the high altitudes to be filled. Our preliminary research suggests that hybridisation is limited between the Himalayan wolf and the Holarctic grey wolf.

We hypothesise that the Himalayan wolf has a fitness benefit over the Holarctic grey wolf in the high altitudes due to its hypoxia adaptation. Our paper concludes that the Himalayan wolf merits taxonomic recognition and designation as an Evolutionary Significant Unit (ESU). Establishing the Himalayan wolf as an ESU will allow for the much-needed conservation to advance on the ground while the taxonomy is being conclusively decided on.

Geraldine observing a young wolf family from afar (Photo credit: Naresh Kusi).

Major challenges. Finding elusive carnivores like wolves in remote and harsh high-altitude habitats is challenging. Wolves inhabit large territories and leave few signs, usually in form of scats at characteristic landscape features, such as ridgelines and crossing points. Hence as researchers trying to find wolves in such vast landscapes we had to learn to read and navigate the valleys and mountains like a wolf pack does. With enough territory covered our team managed to collect substantial data of multiple wolf packs across the different study regions. We could provide a first estimation of the distribution range based on our landscape scale data and found that the Himalayan wolf is more widely distributed than previously thought: this wolf is found across the Himalayas and the Tibetan Plateau, which present a continuous high-altitude ecosystem. The distribution appears to be influenced by elevation, presumably in connection to its genetic adaptation to cope with the low oxygen levels in these high altitudes.

Pack mules, used for transporting scientific gear and equipment, grazing in the evening light on fresh grasses (a welcome nutritious forage for them after the long exhaustive weeks of expedition). Buddhist stupas in the background are characteristic landmarks of worship of the region (Photo credit: Geraldine Werhahn).

Net steps. Our team is now working on piloting conservation action in close collaboration with Himalayan mountain communities to mitigate human-carnivore conflict. The goal is to facilitate a long-term sustainable coexistence of humans and carnivores in these vast, pristine high-altitude habitats. We continue to monitor wolf populations in our study area and are collecting more ecological and behavioural data.

An important question is how many Himalayan wolves exist in wild populations today. Our study provides a first estimate of the population range. But a thorough understanding of their distribution on regional level is required to advance conservation of this species because hunting pressure seems high due to human-wildlife conflict and illegal wildlife trade.

If you could study any organism on Earth, what would it be? It would be canids, especially wolves, followed by snails. I have a profound interest for canids due to their complex social behaviour, their role in trophic cascades and importance in ecosystem functioning, their intelligence, and skilful adaptation to live in remarkably diverse habitats from urban spaces to high mountains and deserts. I would also be interested to study snails, for their immense diversity and occurrence in very diverse habitats and because I find the aesthetics of their “houses” fascinating. They are remarkable in their capability to seemingly disappear when conditions are unfavourable and then reappear in large numbers when conditions improve.

An adult Himalayan wolf watching over the valley below. Looking for his mates or potential prey? (Photo credit: Geraldine Werhahn)

Anything else? This research would not have been possible without a dedicated team that has become my family on the long expeditions. We discovered many new insights around other mammals besides the wolves, birds and plants during these expeditions. For example, my team rediscovered the charismatic wild yak for Nepal during one of our expeditions. This was breaking news for the country of Nepal and eventually our team’s photograph of the wild yak ended up on the Nepalese 5-rupee bill.

Three new initiatives at JBI

In addition to providing a choice venue for publishing the full range of biogeographical research, the Journal of Biogeography (JBI) has also long supported our scientific community in other ways too, e.g. organizing workshops for early career researchers (ECRs) to demistify the publishing process, funding symposia and special issues, providing travel grants to meetings, and supporting conference events. More recently we introduced this Journal of Biogeography blog and more social media to increase the reach of your research. We now add three major new initiatives to support ECRs and beyond, which will be rolled out in the coming six months.

First, we are delighted to announce a new Editorial Academy. The aim of the academy is to help early career biogeographers who are interested in scientific publishing to learn more about the process and to gain experience with the guidance and support of an experienced mentor. There will be multiple positions, each gaining first hand experience as an Associate Editor (with reduced workload) at JBI while working alongside one of the senior editorial team (Michael Dawson, Rosemary Gillespie, Holger Kreft, Richard Ladle, Christine Meynard, Jon Sadler). This partnership will give insight into the way a journal operates, illuminate the sometimes challenging decisions to be made by the editorial team, and allow the academy member to explore their potential to become a full-fledged member of the editorial board. We hope it also provides insight more broadly, beyond the academy, into the qualities of successful publishing as we increasingly build bridges between the editorial team and community of authors; in our view, these are ultimately one and the same. The more transparent and inclusive the process, the more rapid — i.e. fewer rounds of review — and straightforward it will be for all to support publication of creative and high quality biogeographic research. More information on this opportunity and guidelines on how to apply are available on this blog and at the journal’s website.

Second, we will soon open a call for proposals for one or more Global Colloquia.  The event may be stand-alone, or may be staged in association with a larger meeting, it may be in-person or virtual. The topic may be on any aspect of biogeography.  A goal of the colloquium should be to publish a synthesis paper and/or a series of papers that represent the range of topics discussed. We intend these colloquia to become a regular activity that helps biogeographers develop, exchange, and explore ideas that advance biogeography through consolidation of fragmentary knowledge, synthesis across disciplines, and innovation.  This call is not solely for ECRs, but we encourage proposals by and/or including ECRs, as well as biogeographers from our global audience. More information on this opportunity and guidelines on how to apply will be available in a month or two. [UPDATED 15 Aug: see announcements at the blog or the journal].

Third, in October 2020, we will invite submissions of manuscripts by Early Career Researchers for consideration for publication and JBI Awards for Innovation.  This opportunity is modeled after the early career awards first introduced when our current editor-in-chief was on the senior editorial team at Frontiers of Biogeography, and subsequently mirrored by Ecography, and is intended to provide a premium venue for emerging and innovative synthetic biogeography. Our general thinking here is that there is so much good biogeography out there that one award per year is insufficient, and it’s not always well aligned with your publication schedules, so we’ll provide more opportunities at the other end of the year. There will be awards in categories for several article types and all accepted manuscripts will be published free full access for one year. More information on this opportunity and guidelines on how to submit manuscripts will be released in October.

Our goal with each of these three JBI initiatives is to support and provide opportunity for the biogeography community to share the research about which you’re excited and that will advance and shape our discipline in the coming years.

In all these initiatives, following our updated vision and scope, which now includes an equity-diversity-inclusion statement, JBI aims to foster outstanding science that reflects the disciplinary, human, and geographic diversity of biogeography and biogeographers. Submissions will be welcomed from applicants of all ethnicities, races, colors, religions, sexes, sexual orientations, gender identities, national origins, disabilities, ages, or other individual status.

Invitation for applications: JBI Editorial Academy

The Journal of Biogeography is pleased to announce the first of three new opportunities for Early Career Researchers: the Journal of Biogeography’s Editorial Academy.

The Editorial Academy is aimed to help early career biogeographers who are interested to learn more about the publishing process to gain experience with the guidance and support of an experienced mentor.

Editorial Academy members will be partnered one-to-one with a chief editor of the journal.  Academy members will have the same role as a regular member of the Editorial Board, but will work closely with the relevant chief editor throughout the process.  This partnership will illuminate decisions by the editorial team, give insight into the way a journal operation runs, and allow the academy member to explore their potential to become a full-fledged member of the editorial board.  The partnership will convey appropriate procedures and consistent standards of the journal.

Appointments to the Academy are for two years.  The workload will be for 4 manuscripts per year.  Academy members may then be invited to be a continuing member of the editorial board with at least a half-load (6 manuscripts per year) for up to 2 years.  

To be considered for appointment to the Editorial Academy, please submit the following materials as a single PDF:

1. Your CV, including contact information

2. A one page statement explaining:
   – Your interest in and qualifications for the position
               Applicants should have 
                    .. ≥3 years of postdoctoral (or equivalent) experience.
                    .. published ≥6 papers (≥1/2 should be as first/corresponding/senior author)      – Your commitment to the discipline of biogeography
     – The role that being an Academy member will play in your development as a biogeographer
     – Your experience reviewing and, if appropriate, editing
     – Your philosophy on publishing, reviewing, and editing
     – Up to 8 keywords describing your disciplinary interests
     – The name of the chief editor(s) with whom you would prefer to be partnered.

Submit application: 01 August 2020
Decisions announced: 01 September 2020
Appointment begins: 01 October 2020 (negotiable)

Upload applications as a single PDF with the filename “LASTNAME_FIRSTNAME_EditorialAcademy.pdf” only to: *upload only*

Address enquiries (Subject line: “Enquiry: Editorial Academy”) to the Editor-in-Chief at

JBI aims to foster inclusive science that reflects the disciplinary, human, and geographic diversity of biogeography and biogeographers. Submissions are welcomed from applicants of all ethnicities, races, colors, religions, sexes, sexual orientations, gender identities, national origins, disabilities, ages, or other individual status.

A deep dive on ecoregions

Ecoregions are central to global modeling of earth systems & development of conservation plans. There is great variability across taxonomic groups and regions of the world in how strongly ecoregions described community composition.

Two years ago, a team of us published a study entitled ‘A global test of ecoregions’ (Smith et al., 2018). In that paper, we set out with a relatively simple goal, to test whether or not maps of ecoregions, which are popular with both global modelers and conservation practitioners, are actually reflective of the distribution of unique groups of species around the globe? Or are they artifacts of a bygone era when we lacked the data on both the variability in environmental conditions and the distribution of species to properly assess biogeographic patterns?

Above. Our results show that we should expect ecoregions to be much more dissimilar from one another in the tropics than they are in the temperate zone.

EDITORS’ CHOICE: Smith, JR, Hendershot, JN, Nova, N, Daily, GC. (2020) The biogeography of ecoregions: Descriptive power across regions and taxa. J Biogeogr.; 47: 1413– 1426.

Over the course of that project we believed that we would find that ecoregions did not describe communities of species, but instead that species were distributed more or less individualistically across space. We were continually surprised at how wrong we were. We found compelling evidence that ecoregions described clusters of species across space much better than we would expect by chance alone. As we analyzed and discussed our results, we focused on this high level finding that ecoregions did in fact delineate unique communities of species.

It was not until we got comments back from our first round of revisions that we realized that maybe we missed the more interesting story. Our results showed that there was a large amount of variability, both across taxonomic groups and regions of the world, in how strongly ecoregions described community composition. In the short format of that original paper we were unable to dive into the nuance of these environmental and biological mechanisms, but are now thrilled that the Journal of Biogeography has given us the opportunity to do so in our new paper, “The biogeography of ecoregions: Descriptive power across regions and taxa” (Smith et al., 2020).

However, in this expanded format we were able to revisit some of the most famous biogeographers, and test decades or centuries old hypotheses that would never have been possible before the modern data renaissance. We could test if as Humboldt suggested, “associations of the same species of plants [in the tropics] are less consistently extensive, less numerous, than in temperate climates” (Humboldt & Bonpland, 1807). Or if, as Dan Janzen suggested, that “Mountain passes are higher in the tropics” (Janzen, 1967).

A tree line in the Colombian Andes showing sharp transitions between tree and shrub habitats (photo credit Christopher B. Anderson)

What we quickly realized was that climate played a dominant role in shaping how distinct the communities within neighboring ecoregions were from one another. Those ecoregions with higher mean temperatures and more stable temperatures across the year were more distinct from one another. Similarly, steep slopes led ecoregions to be more distinct from one another. Together, this leads us to believe that it is in fact Janzen, and not Humboldt, who was correct in hypothesizing that communities are more distinct from one another in tropical climates.

We were also able to test out more recent theories on how biological traits, such as body size and functional guild, led animals to be more restricted to specific ecoregions. We were able to add evidence to a growing body of literature suggesting that larger species that feed higher on the food chain are more likely to be found across a large number of ecoregions. Sadly, given the state of trait data available today we were only able to do this for four vertebrate taxa: birds, mammals, reptiles, and amphibians.

Despite the fact that we hoped to use this paper to follow up on questions posed by our initial piece, we find ourselves with just as many questions as we started with. How might we extend our findings to other taxonomic groups that we didn’t consider here, such as plant, fungi, and arthropods? How might climate change and land use change affect the robustness of ecoregion classification schemes moving forward? How can we incorporate data on the distribution of species a priori into ecoregional classification schemes (sensu Kreft & Jetz, 2010). We are incredibly excited to continue diving into these topics, along with all the readers of Journal of Biogeography.

Written by:
Jeffrey R. Smith, J. Nicholas Hendershot, Nicole Nova, & Gretchen C. Daily – Stanford University

Additional information:
@JeffreySmithJRS, @Appalachianary, @NicoleNovaBio, @CCBatStanford, @NatCapProject, @StanfordWoods

A beach in Northwestern Costa Rica showing sharp transitions between different types of ecosystems, with the beach quickly giving way to mangroves, which give way to upland shrublands (photo credit Jeffrey Smith)


Humboldt, A. von, & Bonpland, A. (1807). Essay on the Geography of Plants (S. T. Jackson, Ed.; S. Romanowski, Trans.; Reprint edition). University of Chicago Press.

Janzen, D. H. (1967). Why Mountain Passes are Higher in the Tropics. The American Naturalist, 101(919), 233–249.

Kreft, H., & Jetz, W. (2010). A framework for delineating biogeographical regions based on species distributions. Journal of Biogeography, 37(11), 2029–2053.

Smith, J. R., Hendershot, J. N., Nova, N., & Daily, G. C. (2020). The biogeography of ecoregions: Descriptive power across regions and taxa. Journal of Biogeography, 47: 1– 14

Smith, J. R., Letten, A. D., Ke, P.-J., Anderson, C. B., Hendershot, J. N., Dhami, M. K., Dlott, G. A., Grainger, T. N., Howard, M. E., Morrison, B. M. L., Routh, D., Juan, P. A. S., Mooney, H. A., Mordecai, E. A., Crowther, T. W., & Daily, G. C. (2018). A global test of ecoregions. Nature Ecology & Evolution, 2(12), 1889.