Ecological partitioning among parapatric cryptic species

Geographic range differences among species may result from differences in their physiological tolerances. In the intertidal zone, marine and terrestrial environments intersect to create a unique habitat, across which physiological tolerance strongly influences range. Traits to cope with environmental extremes are particularly important here because many species live near their physiological limits and environmental gradients can be steep. The snail Melampus bidentatus occurs in coastal salt marshes in the western Atlantic and the Gulf of Mexico. We used sequence data from one mitochondrial (COI) and two nuclear markers (histone H3 and a mitochondrial carrier protein, MCP) to identify three cryptic species within this broad‐ranging nominal species, two of which have partially overlapping geographic ranges. High genetic diversity, low population structure, and high levels of migration within these two overlapping species suggest that historical range limitations do not entirely explain their different ranges. To identify microhabitat differences between these two species, we modelled their distributions using data from both marine and terrestrial environments. Although temperature was the largest factor setting range limits, other environmental components explained features of the ranges that temperature alone could not. In particular, the interaction of precipitation and salinity likely sets physiological limits that lead to range differences between these two cryptic species. This suggests that the response to climatic change in these snails will be mediated by changes to multiple environmental factors, and not just to temperature alone.     

THE EVOLUTION OF ENVIRONMENTAL TOLERANCE AND RANGE SIZE: A COMPARISON OF GEOGRAPHICALLY RESTRICTED AND WIDESPREAD MIMULUS

The geographic ranges of closely related species can vary dramatically, yet we do not fully grasp the mechanisms underlying such variation. The niche breadth hypothesis posits that species that have evolved broad environmental tolerances can achieve larger geographic ranges than species with narrow environmental tolerances. In turn, plasticity and genetic variation in ecologically important traits and adaptation to environmentally variable areas can facilitate the evolution of broad environmental tolerance. We used five pairs of western North American monkeyflowers to experimentally test these ideas by quantifying performance across eight temperature regimes. In four species pairs, species with broader thermal tolerances had larger geographic ranges, supporting the niche breadth hypothesis. As predicted, species with broader thermal tolerances also had more within‐population genetic variation in thermal reaction norms and experienced greater thermal variation across their geographic ranges than species with narrow thermal tolerances. Species with narrow thermal tolerance may be particularly vulnerable to changing climatic conditions due to lack of plasticity and insufficient genetic variation to respond to novel selection pressures. Conversely, species experiencing high variation in temperature across their ranges may be buffered against extinction due to climatic changes because they have evolved tolerance to a broad range of temperatures.     

Evaluating the potential causes of range limits of birds of the Colombian Andes

Aim To evaluate how factors acting at different spatial scales influence range limits in bird species of the Colombian Andes. Location Andes Mountains of Colombia. Methods We used Maxent , a climate envelope model (CEM), and environmental and geographic information to study range‐filling (i.e. the extent to which a species occurs in all the areas in which it is predicted to occur) in 70 range‐restricted bird species of the Colombian Andes. Environmental data were taken from the WorldClim database, and species occurrence data were taken from museum data collated by the BioMap project, an observational database, and the literature. We evaluated how climate and geographic barriers may shape range limits at two scales. Results At a broad extent (i.e. across the three main cordilleras within the Colombian Andes), we find that CEMs predict there to be suitable environmental conditions for particular species in regions where the species is absent, possibly as a result of dispersal limitation or biotic interactions. In contrast, at a finer scale (within a given cordillera), species generally occur across the entire area predicted to be suitable by a given CEM. Geographic discontinuities within cordilleras do not generally correspond to range limits; instead, range limits correspond to changes in environmental conditions. Main conclusions Our results suggest that different mechanisms influence the presence of species at different scales. Dispersal limitation, potentially combined with species interactions, may influence range limits at a broad extent (the entire Colombian Andes), while strong environmental gradients correspond to range limits at a finer scale (within a cordillera).     

Structural equation modelling analysis of evolutionary and ecological patterns in Australian Banksia

Evolutionary history of species, their geographic ranges, ecological ranges, genetic diversity, and resistance to pathogen infection, have been viewed as being mutually linked through a complex network of interactions. Previous studies have described simple correlations between pairs of these factors, while rarely separated the direct effects among multiple interacting factors. This study was to separate the effect of multiple interacting factors, to reveal the strength of the interactions among these factors, and to explore the mechanisms underlying the ecological and evolutionary processes shaping the geographic range, genetic diversity and fitness of species. I assembled comparative data on evolutionary history, geographic range, ecological range, genetic diversity, and resistance to pathogen infection for thirteen Banksia species from Australia. I used structural equation modelling to test multivariate hypotheses involving evolutionary history, geographic range, genetic diversity and fitness. Key results are: (1) Species with longer evolutionary times tend to occupy larger geographic ranges; (2) higher genetic diversity is directly associated with longer flowering duration in Banksia; and (3) species with higher genotypic diversity have higher level of resistance to infection caused by the pathogen Phytophthora cinnamomi, whereas heterozygosity has the opposite relationship with capacity of resistance to the infections caused by this pathogen. These results revealed a mutually linked and complex network of interactions among gene, species, environment and pathogen in evolutionary and ecological scales. These findings also have great practical significance and help to provide preemptive management options in pathogen control.     

Mutualist‐mediated effects on species' range limits across large geographic scales

Understanding the processes determining species range limits is central to predicting species distributions under climate change. Projected future ranges are extrapolated from distribution models based on climate layers, and few models incorporate the effects of biotic interactions on species' distributions. Here, we show that a positive species interaction ameliorates abiotic stress, and has a profound effect on a species' range limits. Combining field surveys of 92 populations, 10 common garden experiments throughout the range, species distribution models and greenhouse experiments, we show that mutualistic fungal endophytes ameliorate drought stress and broaden the geographic range of their native grass host Bromus laevipes by thousands of square kilometres (~ 20% larger) into drier habitats. Range differentiation between fungal‐associated and fungal‐free grasses was comparable to species‐level range divergence of congeners, indicating large impacts on range limits. Positive biotic interactions may be underappreciated in determining species' ranges and species' responses to future climates across large geographic scales.     

Strong genetic differentiation but not local adaptation toward the range limit of a coastal dune plant

All species have limited geographic distributions; but the ecological and evolutionary mechanisms causing range limits are largely unknown. That many species’ geographic range limits are coincident with niche limits suggests limited evolutionary potential of marginal populations to adapt to conditions experienced beyond the range. We provide a test of range limit theory by combining population genetic analysis of microsatellite polymorphisms with a transplant experiment within, at the edge of, and 60 km beyond the northern range of a coastal dune plant. Contrary to expectations, lifetime fitness increased toward the range limit with highest fitness achieved by most populations at and beyond the range edge. Genetic differentiation among populations was strong, with very low, nondirectional gene flow suggesting range limitation via constraints to dispersal. In contrast, however, local adaptation was negligible, and a distance‐dependent decline in fitness only occurred for those populations furthest from home when planted beyond the range limit. These results challenge a commonly held assumption that stable range limits match niche limits, but also raise questions about the unique value of peripheral populations in expanding species’ geographical ranges.     

Climatic and historical factors controlling horizontal and vertical distribution patterns of two sympatric beech species, Fagus crenata Blume and Fagus japonica Maxim., in eastern Japan

The Japanese Archipelago is unique when viewed in terms of beech flora, since two native species, Fagus crenata Blume and Fagus japonica Maxim., occur sympatrically there. In order to examine the most important environmental or historical factors restricting the geographical ranges of these beech species in eastern Japan (34–44°N and 137–143°E), horizontal and vertical distributions were comparatively examined in detail. The study used two kinds of data sets: (1) DS1, constructed by assembling data from literature, herbarium specimens, etc., and (2) DS2, based on a mesh vegetation database. The upper range limit was expected to be in equilibrium with the current climatic conditions for both species (correlated with temperature factors for F. crenata and with snow depth for F. japonica). The lower range limit was also expected to be in equilibrium with the present climatic condition for both species, but the importance of competition with evergreen trees was also suggested. Dispersal limitation and a topo-geological barrier (for F. japonica) were expected to strongly restrict the northern range limit. Under contrasting climatic conditions in winter (between the Pacific and the Sea of Japan sides in eastern Japan) the geographic ranges of the two beech species are differentiated because of a difference in tolerance to heavy snowfall. A range shift model that assumes a migration lag along a horizontal direction because of dispersal limitation can explain the observed distribution patterns of these beech species in relation to climatic change in the Quaternary.     

Geographic variation in genetic and demographic performance: new insights from an old biogeographical paradigm

The ‘centre–periphery hypothesis’ (CPH) is a long‐standing postulate in ecology that states that genetic variation and demographic performance of a species decrease from the centre to the edge of its geographic range. This hypothesis is based on an assumed concordance between geographical peripherality and ecological marginality such that environmental conditions become harsher towards the limits of a species range. In this way, the CPH sets the stage for understanding the causes of distribution limits. To date, no study has examined conjointly the consistency of these postulates. In an extensive literature review we discuss the birth and development of the CPH and provide an assessment of the CPH by reviewing 248 empirical studies in the context of three main themes. First, a decrease in species occurrence towards their range limits was observed in 81% of studies, while only 51% demonstrated reduced abundance of individuals. A decline in genetic variation, increased differentiation among populations and higher rates of inbreeding were demonstrated by roughly one in two studies (47, 45 and 48%, respectively). However, demographic rates, size and population performance less often followed CPH expectations (20–30% of studies). We highlight the impact of important methodological, taxonomic, and biogeographical biases on such validation rates. Second, we found that geographic and ecological marginality gradients are not systematically concordant, which casts doubt on the reliability of a main assumption of the CPH. Finally, we attempt to disentangle the relative contribution of geographical, ecological and historical processes on the spatial distribution of genetic and demographic parameters. While ecological marginality gradients explain variation in species' demographic performance better than geographic gradients, contemporary and historical factors may contribute interactively to spatial patterns of genetic variation. We thereby propose a framework that integrates species' ecological niche characteristics together with current and past range structure to investigate spatial patterns of genetic and demographic variation across species ranges.     

Experimental biogeography: the role of environmental gradients in high geographic diversity in Cape Proteaceae

One of the fundamental dimensions of biodiversity is the rate of species turnover across geographic distance. The Cape Floristic Region of South Africa has exceptionally high geographic species turnover, much of which is associated with groups of closely related species with mostly or completely non-overlapping distributions. A basic unresolved question about biodiversity in this global hotspot is the relative importance of ecological gradients in generating and maintaining high geographic turnover in the region. We used reciprocal transplant experiments to test the extent to which abiotic environmental factors may limit the distributions of a group of closely related species in the genus Protea (Proteaceae), and thus elevate species turnover in this diverse, iconic family. We tested whether these species have a “home site advantage” in demographic rates (germination, growth, mortality), and also parameterized stage-structured demographic models for the species. Two of the three native species were predicted to have a demographic advantage at their home sites. The models also predicted, however, that species could maintain positive population growth rates at sites beyond their current distribution limits. Thus the experiment suggests that abiotic limitation under current environmental conditions does not fully explain the observed distribution limits or resulting biogeographic pattern. One potentially important mechanism is dispersal limitation, which is consistent with estimates based on genetic data and mechanistic dispersal models, though other mechanisms including competition may also play a role. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The evolution of species' distributions: reciprocal transplants across the elevation ranges of Mimulus cardinalis and M. lewisii

Every species occupies a limited geographic area, but it remains unclear why traits that limit distribution do not evolve to allow range expansion. Hypotheses for the evolutionary stability of geographic ranges assume that species are maladapted at the range boundary and unfit beyond the current range, but this assumption has rarely been tested. To examine how fitness varies across species' ranges, we reciprocally transplanted two species of monkeyflowers, Mimulus cardinalis and M. lewisii, within and beyond their present elevation ranges. We used individuals of known parentage from populations collected across the elevation ranges of both species to examine whether populations are adapted to position within the range. For both species we found the greatest average fitness at elevations central within the range, reduced fitness at the range margin, and zero or near-zero fitness when transplanted beyond their present elevation range limits. However, the underlying causes of fitness variation differed between the species. At high elevations beyond its range, M. cardinalis displayed reduced growth and fecundity, whereas at low elevations M. lewisii experienced high mortality. Weak differences in performance were observed among populations within each species and these were not related to elevation of origin. Low fitness of both species at their range margin and weak differentiation among populations within each species suggest that adaptation to the environment at and beyond the range margin is hindered, illustrating that range margins provide an interesting system in which to study limits to adaptation.     

Life on the edge: carnivore body size variation is all over the place

Evolutionary biologists have long been fascinated by both the ways in which species respond to ecological conditions at the edges of their geographic ranges and the way that species'' body sizes evolve across their ranges. Surprisingly, though, the relationship between these two phenomena is rarely studied. Here, we examine whether carnivore body size changes from the interior of their geographic range towards the range edges. We find that within species, body size often varies strongly with distance from the range edge. However, there is no general tendency across species for size to be either larger or smaller towards the edge. There is some evidence that the smallest guild members increase in size towards their range edges, but results for the largest guild members are equivocal. Whether individuals vary in relation to the distance from the range edges often depends on the way edge and interior are defined. Neither geographic range size nor absolute body size influences the tendency of size to vary with distance from the range edge. Therefore, we suggest that the frequent significant association between body size and the position of individuals along the edge-core continuum reflects the prevalence of geographic size variation and that the distance to range edge per se does not influence size evolution in a consistent way.     

Cactophilic Drosophila in South America: A Model for Evolutionary Studies

The Drosophila buzzatii cluster is composed of seven cactophilic species and their known geographical distribution encompasses the open vegetation diagonal, which includes the morphoclimatic Domains of the Caatinga, Chaco and Cerrado, which are situated between the Amazon and the Atlantic forests. Besides these areas, these cactophilic species are also found in a narrow strip along the Atlantic coast from northeastern Brazil to the southern tip of the country. The hypothesis of vicariant events, defining the core areas of each species, is proposed to explain the historical diversification for the cluster. The intraspecific analysis for the cluster shows a population structure with gene flow restricted by distance, range expansion with secondary contact resulting in introgression and simpatry, especially in the limits of the species distribution, polytypic populations and assortative mating in inter population experiments. There is a variation related to these events that depends on the species and geographic origin of the population analyzed. These events are, hypothetically, described as the results of expansion and retraction of the population ranges, as a consequence of their association with cacti, which theoretically follow the expansion and retraction of dry areas during the paleoclimatic oscillations in South America, as that promoted by the glacial cycles of the Quaternary. The Drosophila buzzatii cluster is divided into two groups. The first one is composed of D. buzzatii, a species that has a broad geographic distribution and no significant differentiation between its populations. The second is the Drosophila serido sibling set, which encompasses the others species and is characterized by a significant potential for differentiation.     

The performance of the global protected area system in capturing vertebrate geographic ranges

Given the heavy reliance placed on and investment in protected areas for biological conservation, there has been much debate as to how effective these are in representing biodiversity features within their boundaries. The majority of studies addressing this issue have been conducted on a regional or national basis, precluding a broad picture of patterns of representation at the species level. We present a global assessment of the representation of the terrestrial geographic ranges of complete taxonomic groups: all known extant amphibians, birds and mammals (20,736 species) within the current global system of protected areas. We conclude that it is necessary substantially to improve the levels of coverage of the geographic ranges of the majority of species, even the widespread ones. This is particularly true for rare species, which might be assumed to be foci for protected area systems. To improve on the low levels of coverage of vertebrate ranges attained by the existing areas, key regions should be targeted, but heavy reliance will also have to be placed on approaches to sustaining populations in the wider, unprotected landscape.     

Physiological Limits along an Elevational Gradient in a Radiation of Montane Ground Beetles

A central challenge in ecology and biogeography is to determine the extent to which physiological constraints govern the geographic ranges of species along environmental gradients. This study tests the hypothesis that temperature and desiccation tolerance are associated with the elevational ranges of 12 ground beetle species (genus Nebria) occurring on Mt. Rainier, Washington, U.S.A. Species from higher elevations did not have greater cold tolerance limits than lower-elevation species (all species ranged from -3.5 to -4.1°C), despite a steep decline in minimum temperature with elevation. Although heat tolerance limits varied among species (from 32.0 to 37.0°C), this variation was not generally associated with the relative elevational range of a species. Temperature gradients and acute thermal tolerance do not support the hypothesis that physiological constraints drive species turnover with elevation. Measurements of intraspecific variation in thermal tolerance limits were not significant for individuals taken at different elevations on Mt. Rainier, or from other mountains in Washington and Oregon. Desiccation resistance was also not associated with a species’ elevational distribution. Our combined results contrast with previously-detected latitudinal gradients in acute physiological limits among insects and suggest that other processes such as chronic thermal stress or biotic interactions might be more important in constraining elevational distributions in this system.     

Thermal sensitivity of cold climate lizards and the importance of distributional ranges

One of the fundamental goals in macroecology is to understand the relationship among species’ geographic ranges, ecophysiology, and climate; however, the mechanisms underlying the distributional geographic patterns observed remain unknown for most organisms. In the case of ectotherms this is particularly important because the knowledge of these interactions may provide a robust framework for predicting the potential consequences of climate change in these organisms. Here we studied the relationship of thermal sensitivity and thermal tolerance in Patagonian lizards and their geographic ranges, proposing that species with wider distributions have broader plasticity and thermal tolerance. We predicted that lizard thermal physiology is related to the thermal characteristics of the environment. We also explored the presence of trade-offs of some thermal traits and evaluated the potential effects of a predicted scenario of climate change for these species. We examined sixteen species of Liolaemini lizards from Patagonia representing species with different geographic range sizes. We obtained thermal tolerance data and performance curves for each species in laboratory trials. We found evidence supporting the idea that higher physiological plasticity allows species to achieve broader distribution ranges compared to species with restricted distributions. We also found a trade-off between broad levels of plasticity and higher optimum temperatures of performance. Finally, results from contrasting performance curves against the highest environmental temperatures that lizards may face in a future scenario (year 2080) suggest that the activity of species occurring at high latitudes may be unaffected by predicted climatic changes.     

Geographic range size is predicted by plant mating system

Species' geographic ranges vary enormously, and even closest relatives may differ in range size by several orders of magnitude. With data from hundreds of species spanning 20 genera in 15 families, we show that plant species that autonomously reproduce via self‐pollination consistently have larger geographic ranges than their close relatives that generally require two parents for reproduction. Further analyses strongly implicate autonomous self‐fertilisation in causing this relationship, as it is not driven by traits such as polyploidy or annual life history whose evolution is sometimes correlated with selfing. Furthermore, we find that selfers occur at higher maximum latitudes and that disparity in range size between selfers and outcrossers increases with time since their evolutionary divergence. Together, these results show that autonomous reproduction—a critical biological trait that eliminates mate limitation and thus potentially increases the probability of establishment—increases range size.     

The geographic range size and vulnerability to extinction of angiosperm epiphytes in the Atlantic Forest of Brazil

Aim

Angiosperm epiphytes have long been reported to have larger geographic ranges than terrestrial species, despite evidence of their outstanding diversity and endemism. This apparent contradiction calls for further investigation of epiphytes' poorly understood range size patterns. Here, we address the question of whether epiphytes have larger geographic ranges and different vulnerability to extinction than terrestrial species.

Location

The Atlantic Forest of Brazil, a global centre of tropical epiphyte diversity with relatively well-known flora, where we can estimate the geographic ranges of a large number of species with reasonable confidence.

Time period

Occurrence records from the 17th century to the year 2021.

Major taxa studied

Flowering plants (angiosperms).

Methods

We downloaded, processed and cleaned all occurrence records for the angiosperm species native to the Atlantic Forest of Brazil available in the speciesLink network and the Global Biodiversity Information Facility. We estimated the extent of occurrence and area of occupancy of 12,679 native flowering plants, including 1251 epiphytic species. We compared the geographic ranges of epiphytes and other life forms at broad (e.g. Angiosperms, Monocots) and more restricted taxonomic scales (e.g. individual families), assuming species are independent entities and also when accounting for species phylogenetic dependence.

Results

We found that epiphytes have among the smallest geographic ranges of flowering plants. We found no consistent evidence that epiphytism leads to differences in geographic ranges between close relatives. However, both epiphytes and non-epiphytes in epiphyte-rich lineages have small ranges and likely a high vulnerability to extinction.

Main Conclusions

Our findings contrast with the long-held hypothesis that epiphytes have larger geographic ranges than terrestrial species. Epiphytes and their close relatives share many diversification mechanisms and ecological adaptations (‘epiphyte-like traits’), which probably explain why both sets of species have small range sizes and high vulnerability to extinction.     

Calymperes (Musci: Calymperaceae): World ranges,implications for patterns of historical dispersion and speciation,and comments on phylogeny

Refinements in the taxonomy of the genus Calymperes (Musci: Calymperaceae) allow increasingly accurate estimates of the total number of species worldwide and new assessments of world ranges of the species. Instead of the 272 correct names of Calymperes listed in Volume 1 of Index Muscorum, the world total of “good” species now appears to be in the range of 30 to 40, and contrary to the high degree of narrow endemism implied from the compilation of Index Muscorum, many species of Calymperes have broad ranges in the paleo- or neotropics. A few species, including C. afzelii, C. erosum, C. palisotii, C. lonchophyllum, and C. tenerum, are pantropical. World ranges of species of Calymperes are mapped and implications for historical patterns of dispersal and speciation are discussed. Suites of endemic species and species groups in the neotropics differ from those in the paleotropics, demonstrating different ancestries, a long period of geographic separation, and independent evolution in two fundamentally different environments. It is suggested that Calymperes shares close phylogenetic relationship with limbate Syrrhopodon.     

Environmental marginality and geographic range limits: a case study with Arabidopsis lyrata ssp. lyrata

Understanding the factors that govern the distribution of species is a central goal of evolutionary ecology. It is commonly assumed that geographic range limits reflect ecological niche limits and that species experience increasingly marginal conditions towards the edge of their ranges. Using spatial data and ecological niche models we tested these hypotheses in Arabidopsis lyrata. Specifically, we asked whether range limits coincide with predicted niche limits in this system and whether the suitability of sites declines towards the edge of the species’ range in North America. We further explored patterns of environmental change towards the edge of the range and asked whether genome‐wide patterns of genetic diversity decline with increasing peripherality and environmental marginality. Our results suggest that latitudinal range limits coincide with niche limits. Populations experienced increasingly marginal environments towards these limits – though patterns of environmental change were more complex than most theoretical models for range limits assume. Genomic diversity declined towards the edge of the species’ range and with increasing distance from the estimated centre of the species’ niche in environmental space, but not with the suitability of sites based on niche model predictions. Thus while latitudinal range limits in this system are broadly associated with niche limits, the link between environmental conditions and genetic diversity (and thus the adaptive potential of populations) is less clear.     

Species Boundaries and Host Range of Tortoise Mites (Uropodoidea) Phoretic on Bark Beetles (Scolytinae), Using Morphometric and Molecular Markers

Understanding the ecology and evolutionary history of symbionts and their hosts requires accurate taxonomic knowledge, including clear species boundaries and phylogenies. Tortoise mites (Mesostigmata: Uropodoidea) are among the most diverse arthropod associates of bark beetles (Curculionidae: Scolytinae), but their taxonomy and host associations are largely unstudied. We tested the hypotheses that (1) morphologically defined species are supported by molecular data, and that (2) bark beetle uropodoids with a broad host range comprise cryptic species. To do so, we assessed the species boundaries of uropodoid mites collected from 51 host species, across 11 countries and 103 sites, using morphometric data as well as partial cytochrome oxidase I (COI) and nuclear large subunit ribosomal DNA (28S). Overall, morphologically defined species were confirmed by molecular datasets, with a few exceptions. Twenty-nine of the 36 uropodoid species (Trichouropoda, Nenteria and Uroobovella) collected in this study had narrow host ranges, while seven species had putative broad host ranges. In all but one species, U. orri, our data supported the existence of these host generalists, which contrasts with the typical finding that widespread generalists are actually complexes of cryptic specialists.