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Frontiers of Biogeography

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Frontiers of Biogeography (FoB) is the scientific journal 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.

Issue cover
Image of the Upland Burrowing Tree Frog Smilisca dentata (Smith, 1957) during its period of activity in the natural grasslands south of the state of Aguascalientes in Mexico. In this issue, Encarnación-Luévano and colleagues explore the seasonal niches of this endangered species and the burrowing species Smilisca fodiens to test the importance of behavior in overcoming the environmental barriers conserving their niches of tropical origin. Photo by Gustavo E. Quintero-Díaz.


Opinions, Perspectives & Reviews

A framework linking biogeography and species traits to plant species vulnerability under global change in Mediterranean-type ecosystems

We review the roles that plant species traits and biogeography play in species’ exposure and vulnerability to decline or extinction under global change, focusing on separate and combined impacts of multiple threats – climate change, land-use change, and altered disturbance regimes. We establish a conceptual framework and research agenda for identifying the spatial characteristics of species ranges, as well as the life history and functional traits, that are associated with extinction risk for plant species with functional attributes emblematic of fire-prone, winter-precipitation Mediterranean-type ecosystems (MTEs). MTEs worldwide are characterized by their high plant diversity and unique floras, historical and contemporary high rates of land use change, and strong interactions between climate, fire, and land use. We focus on the California Floristic Province (CFP), an MTE that is a global plant diversity hotspot, and show how our framework can be used to understand the relationships between vulnerability to multiple global change drivers, species traits, and biogeography. Vulnerability can be assessed across species using linked distribution and population models that forecast plant responses to global change scenarios. Our overarching hypothesis is that species-specific vulnerability to global change in MTEs is a function of interactions between species and spatial traits: the nature of this interaction will depend on the type of global change process.

Globally important plant functional traits for coping with climate change

The last decade has seen a proliferation of studies that use plant functional traits to assess how plants respond to climate change. However, it remains unclear whether there is a global set of traits that can predict plants’ ability to cope or even thrive when exposed to varying manifestations of climate change. We conducted a systematic global review which identified 148 studies to assess whether there is a set of common traits across biomes that best predict positive plant responses to multiple climate changes and associated environmental changes.  Eight key traits appear to best predict positive plant responses to multiple climate/environmental changes across biomes: lower or higher specific leaf area (SLA), lower or higher plant height, greater water-use efficiency (WUE), greater resprouting ability, lower relative growth rate, greater clonality/bud banks/below-ground storage, higher wood density, and greater rooting depth. Trait attributes associated with positive responses appear relatively consistent within biomes and climate/environmental changes, except for SLA and plant height, where both lower and higher trait attributes are associated with a positive response depending on the biome and climate/environmental change considered.  Overall, our findings illustrate important and general trait-climate responses within and between biomes that help us understand which plant phenotypes may cope with or thrive under current and future climate change.

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The interface between Macroecology and Conservation: existing links and untapped opportunities

Human activities are altering the structure of ecosystems, compromising the benefits they provide to nature and people. Effective conservation actions and management under ongoing global change rely on a better understanding of socio-ecological patterns and processes across broad spatiotemporal scales. Both macroecology and conservation science contribute to this improved understanding and, while they have different scopes, these disciplines have become increasingly interconnected over time. Here we describe examples of how macroecology has contributed to conservation science, and how conservation science can motivate further macroecological developments and applications. We identify challenges and untapped potential to further strengthen the links between these two disciplines. Major macroecological contributions include developing ecological theory, providing methodologies useful for biodiversity assessments and projections, making data more accessible and addressing knowledge gaps. These contributions have played a major role in the development of conservation science, and have supported outreach to policy makers, media, and the public. Nonetheless, a pure macroecological lens is limited to inform conservation decisions, particularly in local contexts, which frequently leads to the misuse of macroecological analyses for conservation applications, misunderstandings of research outputs, and skepticism among conservation practitioners and scientists. We propose possible solutions to overcome these challenges and strengthen links between macroecology and conservation science, including a stronger focus on ecological mechanisms and predictive approaches, and the creation of hybrid journals and meetings. Finally, we suggest new avenues for macroecological research that would further benefit conservation science.

The two South American dry diagonals

South American drylands roughly form two diagonals both termed in the biogeographical literature as the "South American dry diagonal" (SADD). However, they correspond to two different geographical areas. One comprises the Caatinga, Cerrado and Chaco domains, thus encompassing the areas between northeastern Brazil and northwestern Argentina. The other stretches from Patagonia in southern Argentina to the Pacific deserts of northern Chile and Peru, thus also including the Monte, Prepuna and dry Puna domains. I termed them the eastern and western SADDs, respectively (i.e., eSADD and wSADD). In this mini review I attempt to summarize the major climatic features of the two South American dry diagonals, their possible origins, biogeographical patterns within and around them and to explore possible interconnections. The eSADD is generally more humid than the wSADD and has more pronounced rainfall seasonality, with precipitation concentrated in summer, while the wSADD tends to be less seasonal due to year-round aridity, with little precipitation largely occurring in winter. The origin of both diagonals appears to go back to the middle Miocene, associated with global cooling. Biogeographical studies show that these diagonals are important in structuring South American distribution patterns south of the Equator, both acting as barriers for humid-adapted lineages and corridors for arid-adapted taxa. Remarkably, the two diagonals appear to have few plant and animal taxa in common, which may explain why biogeographers speaking about one diagonal seem to ignore the existence of the other.

Research Articles

Geographic and ecological segregation in an extinct guild of flightless birds: New Zealand’s moa

The nine currently recognized species of moa (Order – Dinornithiformes; Bonaparte 1853) suffered extinction soon after New Zealand was settled by humans.  They were the result of an evolutionary radiation that produced a unique guild of birds – giant, and totally wingless species that evolved in the absence of non-volant mammals.   Recent advances in dating and paleoclimatology, and compilations of data on distributions of the nine species of moa, along with information on the geographic, topographic, climatic and edaphic characteristics of sites from which moa remains have been recovered, enabled us to test whether their evolutionary radiation truly was ‘adaptive’, producing ecologically distinct species.  Randomization, resampling analyses of moa distributions across North and South Islands revealed highly significant geographic and ecological segregation, with different species tending to occupy different islands, regions within islands, or elevations within regions.  Quadratic Discriminant Analyses demonstrated niche segregation at even finer scales, including that based on vegetation-defined habitats and on local climatic, topographic and edaphic conditions.  Moa distributions also appear to have been dynamic over time, shifting in their upper elevational limits as climatic conditions changed and vegetative zones shifted upward during the Holocene Epoch.  Our ongoing studies are building on the results presented here to explore the temporal dynamics of moa distributions, assess differential responses of moa species to natural and anthropogenic drivers, and determine how these forces may have combined to cause the extinction of moa just a few centuries ago.

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Hierarchical analyses of community biogeography in the Afromontane highlands

The Afromontane mountains are a complex series of highlands that have intermittently been connected by habitat corridors during climatic cycles, resulting in a mosaic of range disjunctions and allospecies complexes in the present day. Patterns of community relatedness between geographic regions are often determined through single-species analyses or spatial analyses of diversity and nestedness at the species level. To understand patterns of Afromontane community evolution and to assess the effects of taxonomy on our understanding of biogeographic patterns, I concatenated three different lists of Afromontane bird taxa divided into five different taxonomic hierarchies. These lists were converted into a presence-absence matrix across 42 different montane regions, and analyzed using multiple different clustering techniques using a replicable coding pipeline. I use these lists and methods to determine patterns of relatedness between montane blocks, to assess the consistency with which biogeographic regions are recovered, and to shed more light on patterns of connectivity within the Afromontane region. Results reaffirm the distinctiveness of many different biogeographic regions (i.e., the Cameroon Highlands) while also clarifying regional relationships and the presence of ‘transition zones’ between regions. Differences between lists illustrate how our understanding of taxonomy and distribution in the Afromontane highlands can also change our understanding of Afromontane biogeography. Most notably, I find evidence for an Expanded Eastern Arc that includes the Eastern Arc Mountains and highlands in Malawi, Mozambique, and Zimbabwe. This study presents a rigorous yet easily adjustable pipeline for studying regional biogeography from multiple perspectives with classical and novel approaches.

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Species–area relationships of the Aegean, a comparative approach between six taxa

Islands occupy a proportionately small area on Earth, however they play a crucial role in Ecology and Biogeography, as they constitute “natural laboratories”. The increased number of species, with increasing island area, is such a commonly observed pattern that it has been labelled as one of the few laws of ecology. The Aegean archipelago is of broad biogeographical interest, as it has a considerable number of islands in addition to a rich paleogeographical and geological history, while being divided among three continents (Europe, Asia, Africa). As a result, the composition of life in the Aegean is dominated by species of European, Asian, African origin as well as species endemic in the archipelago. In this framework, we approached the species–area relationship (SAR) of the Aegean islands for six different organismic groups (birds, herptiles, snails, isopods, tenebrionids and chilopods) and 20 different models. The aim was to determine which model(s) perform better for each taxon and also to compare the z and C parameters of the power model between animal groups, which are the only model parameters to date that have been linked with biological processes. We compared the relationship across different taxa for the entire archipelago and for the exact same islands, in two subgroups with similar paleogeographic history and environmental conditions in the central and eastern Aegean. For the taxonomic groups that were examined a strong correlation between the number of species and area was found, except for chilopods and herptiles. Although there is no universal best model for the SAR of the Aegean, the power model performed better for invertebrates, whereas concerning vertebrates there was more ambiguity in the shape of the relationship.

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Cryptic diversity of Rhinolophus lepidus in South Asia and differentiation across a biogeographic barrier

Peninsular India is an important region for mammalian diversification and harbors major biogeographic barriers. However, little is known about the role of this region in the diversification of bats though it harbors high chiropteran diversity. In this study, we used phenotypic, acoustic, and genetic markers to assess the diversification of Rhinolophus lepidus bats in South Asia. We first investigated if peninsular India is associated with speciation of R. lepidus. Further, we tested if the Palghat Gap acts as a biogeographic barrier to gene flow in this species. Our results revealed cryptic genetic diversity in peninsular India suggesting that this region holds at least one endemic species level lineage of the R. lepidus species complex.  Analyses of populations of R. lepidus across the Palghat Gap in the Western Ghats revealed clinal variation in phenotype, with bats south of this barrier being bigger and emitting echolocation calls of higher frequency. We also observed that populations on either side of the Palghat Gap have remained genetically isolated since the mid-Holocene.

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Burrowing habit in Smilisca frogs as an adaptive response to ecological niche constraints in seasonally dry environments

As environmental conditions change over time, some species can follow the spatial footprint of their ecological niches or can adapt physiologically to the new conditions; modifying behavior can offer an alternative means of adapting to novel environments. The burrowing habit allows organisms to avoid adverse climatic conditions during part of the year by remaining inside burrows. Smilisca fodiens and S. dentata are two burrowing hylid frogs that inhabit areas beyond the northernmost distributional limits of the other six arboreal species of their genus, and indeed beyond of most American hylids. In this study, we tested whether burrowing habit allows these species to adapt to drier conditions while conserving the climatic niche of the arboreal species. We compared the annual niches of the arboreal species to those of the burrowing species under two assumptions: true seasonal niches and full annual niches. Through ecological niche similarity tests, we performed 24 comparisons in both geographic and environmental spaces. In geographic space, when considering burrowing annual niches, only five of 24 tests indicated similarity, yet as regards seasonal niche, 18 of 24 tests indicated similarity. In environmental space, all tests failed to reject null hypotheses. The analyses showed clearly that burrowing and arboreal species were closer in environmental space when seasonal niches of the burrowing species were used, rather than annual niches. That is, climatic conditions in seasonal niches of burrowing species resemble the annual niches of arboreal species, supporting the proposition that reduction of activity to certain periods of the year is a strategy in burrowing species to conserve their tropical niches while living in dry regions.

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A simulation-based method for selecting calibration areas for ecological niche models and species distribution models

Ecological niche models and species distribution models (ENM and SDM, respectively) are tools that have seen massive use and considerable improvement during the last twenty years. The choice of calibration areas for such models has strong effects on model outcomes and model interpretation, as well as on model transfer to distinct environmental settings. However, approaches to selecting these areas remain simple and/or unlinked to biological concepts. Such models should be calibrated within areas that the species of interest has explored throughout its recent history, the accessible area (M). In this paper, we provide a simulation approach for estimating a species’ M considering processes of dispersal, colonization, and extinction in constant current climate or glacial-interglacial climate change frameworks, implemented within a new R package we developed called grinnell. Using the avian genus Aphelocoma, we explored different parameterizations of our simulation, and compared them to current approaches for M selection, in terms of model performance and risk of extrapolation using the algorithm Maxent and mobility-oriented parity analyses. Model calibration exercises from all approaches resulted in at least one model meeting optimal performance criteria for each species; however, we noted high variability among taxa and M selection methods. More importantly, M hypotheses derived directly from simulations of key biological processes, rather than being based on simple proxies of those processes, and as such are better suited to erecting biologically appropriate contrasts in model calibration, and to characterizing the potential for model extrapolation more rigorously. Major factors in our simulations were environmental layer resolution, dispersal kernel characteristics, and the inclusion of a changing framework of climatic conditions. This contribution represents the first simulation-based method for selecting calibration areas for ENM and SDM, offering a quantitative approach to estimate the accessible area of a species while considering its dispersal ability, along with patterns of change in environmental suitability across space and time.

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Invaders in waiting? Non-equilibrium in Southern Hemisphere seaweed distributions may lead to underestimation of Antarctic invasion potential

Bioinvasions pose a major threat to global biodiversity. Correlative Ecological Niche Models (ENMs) can be a valuable tool to identify invaders and invasion sites. However, in cases when species are in non-equilibrium with their native environment (i.e. do not fill their niche), correlative approaches have limited power and invasions lead to shifts of the realized niche. In recent years, several new seaweed species have been reported in Antarctica. It is impossible to unequivocally identify which of these species are truly non-natives, however, here, we provide literature-based evidence that seaweed species have been introduced to Antarctica. Under this assumption, we reconstruct pre- and post-introduction niches of these species, calculate relative niche sizes and overlap between pre-Antarctic and Antarctic sites, and evaluate increase in niche size due to inclusion of Antarctic habitats. In seven species, the absolute occupied temperature range is dramatically enlarged, with minimum sea surface temperature (SST) being 2-5°C lower than in the pre-Antarctic ranges. In all species except one, summer SST is 5-20°C lower than in the pre-Antarctic ranges. As a result, several species’ niches increase dramatically. We hypothesize that species from the Southern Hemisphere do not cover their whole abiotically suitable range due to lack of settling substrate in cold-water regions while species from the Northern Hemisphere tend to fill their niches to a greater degree due to higher connectivity between tropic and polar regions along coastlines. Thus, while correlative ENMs for Northern Hemisphere species will probably be successful in predicting Antarctica as a suitable habitat, such models will likely be insufficient to do so for Southern Hemisphere species. From a precautionary standpoint, we argue that not only species from climatically matching regions pose an invasion threat for Antarctica, but that also species from other, climatically non-matching regions, might be potential invaders. In light of higher connectivity of the Antarctic continent with other continents this finding significantly increases invasion risk for Antarctica.

Climatic drivers of Sphagnum species distributions

Peat mosses (genus Sphagnum) dominate most Northern mires and show distinct distributional limits in Europe despite having efficient dispersal and few dispersal barriers. This pattern indicates that Sphagnum species distributions are strongly linked to climate. Sphagnum-dominated mires have been the largest terrestrial carbon sinks in Europe over the last few millennia. Understanding the climatic drivers of Sphagnum species distributions is important for predicting the future functionality of peatlands. We used MaxEnt, with biologically relevant climatic variables, to model and clarify the current distributions of 45 Sphagnum species in Europe. We used a dataset of 238 316 records from across Europe (30° to 90° N, -30° to 63° E; Sahara to the Arctic, Azores to Ural mountains). We used annual degree-days, annual water balance and their monthly standard deviations (i.e. seasonality) as climatic predictors over a range of spatial resolutions (from 10 to 200 km pixel size). With these climatic predictors, we produced reasonably accurate projections of the distribution of 45 species (overall AUC >0.8). Large pixels (100 and 200 km) resulted in loss of detail, but smaller pixels (10-50 km) performed well across fit measurements. Projected distributions at the 50 × 50 km resolution showed the largest resemblance to published distribution maps. Suitable climate for many Sphagnum species was associated with the northern, western and mountainous parts of Europe. We found that annual water balance was an important indicator of Sphagnum presence. Limits in relation to annual water balance were the same as reported by bioclimatic peatland models from North America. Most Sphagnum species were limited to annual degree-days between -5000 °C y-1 and 5000 °C y-1. Seasonality in both climate variables separated species, with degree-day seasonality having a stronger influence than water balance seasonality. High degree-day seasonality as a consequence of cold temperature sets a northern distribution limit to some species. The results suggest that the future of Sphagnum diversity in Europe is most strongly dependent on changes in water availability and in seasonal temperature variation.

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