Maladaptation beyond a geographic range limit driven by antagonistic and mutualistic biotic interactions across an abiotic gradient

Species’ geographic range limits often result from maladaptation to the novel environments beyond the range margin. However, we rarely know which aspects of the n‐dimensional environment are driving this maladaptation. Especially of interest is the influence of abiotic versus biotic factors in delimiting species’ distributions. We conducted a 2‐year reciprocal transplant experiment involving manipulations of the biotic environment to explore how spatiotemporal gradients in precipitation, fatal mammalian herbivory, and pollination affected lifetime fitness within and beyond the range of the California annual plant, Clarkia xantiana ssp. xantiana. In the first, drier year of the experiment, fitness outside the range edge was limited mainly by low precipitation, and there was some evidence for local adaptation within the range. In the second, wetter year, we did not observe abiotic limitations to plant fitness outside the range; instead biotic interactions, especially herbivory, limited fitness outside the range. Together, protection from herbivory and supplementation of pollen resulted in three‐ to sevenfold increases in lifetime fitness outside the range margin in the abiotically benign year. Overall, our work demonstrates the importance of biotic interactions, particularly as they interact with the abiotic environment, in determining fitness beyond geographic range boundaries.     

Climate warming is associated with smaller body size and shorter lifespans in moose near their southern range limit

Despite the importance of body size for individual fitness, population dynamics and community dynamics, the influence of climate change on growth and body size is inadequately understood, particularly for long‐lived vertebrates. Although temporal trends in body size have been documented, it remains unclear whether these changes represent the adverse impact of climate change (environmental stress constraining phenotypes) or its mitigation (via phenotypic plasticity or evolution). Concerns have also been raised about whether climate change is indeed the causal agent of these phenotypic shifts, given the length of time‐series analysed and that studies often do not evaluate – and thereby sufficiently rule out – other potential causes. Here, we evaluate evidence for climate‐related changes in adult body size (indexed by skull size) over a 4–decade period for a population of moose (Alces alces) near the southern limit of their range whilst also considering changes in density, predation, and human activities. In particular, we document: (i) a trend of increasing winter temperatures and concurrent decline in skull size (decline of 19% for males and 13% for females) and (ii) evidence of a negative relationship between skull size and winter temperatures during the first year of life. These patterns could be plausibly interpreted as an adaptive phenotypic response to climate warming given that latitudinal/temperature clines are often accepted as evidence of adaptation to local climate. However, we also observed: (iii) that moose with smaller skulls had shorter lifespans, (iv) a reduction in lifespan over the 4‐decade study period, and (v) a negative relationship between lifespan and winter temperatures during the first year of life. Those observations indicate that this phenotypic change is not an adaptive response to climate change. However, this decline in lifespan was not accompanied by an obvious change in population dynamics, suggesting that climate change may affect population dynamics and life‐histories differently.     

Measures of geographic range size: the effects of sample size

A number of methods have been used for quantifying the sizes of the geographic ranges of species. The consequences of different levels of sampling (the proportion of actual spatial occurrences) are explored for eight of these, using data on the occurrences of butterfly species on a 10 × 10 km grid across Britain. For all methods, the percentage error of estimation (PEE) decreases with the number of 10 × 10 km squares which a species occupies, most rapidly for extent measures, and more rapidly for area measures than for measures of numbers of units occupied. The rate of decline in PEE itself falls as sampling effort increases. At a given sampling level, rank correlations between range sizes measured by different methods are generally high, but there is no consistent change in the magnitude of these correlations as the level of sampling increases. The composition of the set of species with the smallest range sizes changes with the level of sampling.     

Global patterns of geographic range size in birds

Large-scale patterns of spatial variation in species geographic range size are central to many fundamental questions in macroecology and conservation biology. However, the global nature of these patterns has remained contentious, since previous studies have been geographically restricted and/or based on small taxonomic groups. Here, using a database on the breeding distributions of birds, we report the first (to our knowledge) global maps of variation in species range sizes for an entire taxonomic class. We show that range area does not follow a simple latitudinal pattern. Instead, the smallest range areas are attained on islands, in mountainous areas, and largely in the southern hemisphere. In contrast, bird species richness peaks around the equator, and towards higher latitudes. Despite these profoundly different latitudinal patterns, spatially explicit models reveal a weak tendency for areas with high species richness to house species with significantly smaller median range area. Taken together, these results show that for birds many spatial patterns in range size described in geographically restricted analyses do not reflect global rules. It remains to be discovered whether global patterns in geographic range size are best interpreted in terms of geographical variation in species assemblage packing, or in the rates of speciation, extinction, and dispersal that ultimately underlie biodiversity.     

The relationship between dispersal ability and geographic range size

There are a variety of proposed evolutionary and ecological explanations for why some species have more extensive geographical ranges than others. One of the most common explanations is variation in species' dispersal ability. However, the purported relationship between dispersal distance and range size has been subjected to few theoretical investigations, and empirical tests reach conflicting conclusions. We attempt to reconcile the equivocal results of previous studies by reviewing and synthesizing quantitative dispersal data, examining the relationship between average dispersal ability and range size for different spatial scales, regions and taxonomic groups. We use extensive data from marine taxa whose average dispersal varies by seven orders of magnitude. Our results suggest dispersal is not a general determinant of range size, but can play an important role in some circumstances. We also review the mechanistic theories proposed to explain a positive relationship between range size and dispersal and explore their underlying rationales and supporting or refuting evidence. Despite numerous studies assuming a priori that dispersal influences range size, this is the first comprehensive conceptual evaluation of these ideas. Overall, our results indicate that although dispersal can be an important process moderating species' distributions, increased attention should be paid to other processes responsible for range size variation.     

Climate tolerance and interspecific variation in geographic range size

The fact that climate influences the continental-scale distributions of species is one of the central tenets of ecology and biogeography. Equally elemental is that species exhibit enormous variation in geographic range size, with most occupying comparatively small areas. The degree to which climate can account for this variation remains unclear. Here, I test three alternative climate-based hypotheses for variation in range size using a large sample of tree and shrub species native to North America north of Mexico. I show that the lowest value of January average daily minimum temperature encompassed by a species' geographic range (T(MIN)), representing the 'climate extremes hypothesis', explains almost 80% of the variation in range size. Hypotheses based on seasonality and climate optima find substantially less support. The relationship between range size and T(MIN) does not change across the breadth of latitudes examined, and is general for conifers and hardwoods, and growth form (tree versus shrub). Differential freezing resistance gains support as the mechanism underlying interspecific variation in range size: using 35 species for which data were available, both T(MIN) and range size are shown to be strongly correlated with measures of freezing resistance.     

Interactions between soil habitat and geographic range location affect plant fitness

Populations are often found on different habitats at different geographic locations. This habitat shift may be due to biased dispersal, physiological tolerances or biotic interactions. To explore how fitness of the native plant Chamaecrista fasciculata depends on habitat within, at and beyond its range edge, we planted seeds from five populations in two soil substrates at these geographic locations. We found that with reduced competition, lifetime fitness was always greater or equivalent in one habitat type, loam soils, though early-season survival was greater on sand soils. At the range edge, natural populations are typically found on sand soil habitats, which are also less competitive environments. Early-season survival and fitness differed among source populations, and when transplanted beyond the range edge, range edge populations had greater fitness than interior populations. Our results indicate that even when the optimal soil substrate for a species does not change with geographic range location, the realized niche of a species may be restricted to sub-optimal habitats at the range edge because of the combined effects of differences in abiotic and biotic effects (e.g. competitors) between substrates.     

Evolutionary patterns in the geographic range size of Atlantic Forest plants

Species’ geographic range size is arguably the single most important predictor of vulnerability to extinction and a key metric in ecology. Despite this, patterns of specific variation in range size and their underlying reasons are still poorly understood. For example, hypotheses on how evolutionary history affects range size have scarcely been tested. To address these questions, we focused on Brazil's Atlantic Forest flora, one of the most species-rich in the world, relatively well-known and highly threatened. We investigated whether and how lineages’ diversification rate, number of species and age are associated with species’ geographic range size. We estimated the extent of occurrence and area of occupancy of each of 13 283 plant species native to the Atlantic Forest region based on over 500 000 unique records. We used phylogenetic least squares and logistic regressions to analyze how the predictors affect the geographic range size. On average, the higher the diversification rate and number of species in the lineage, the smaller the species range size and the higher the proportion of species with vulnerably small range size. Lineage age showed no clear effect on average range size. The results support our expectations that dynamics of diversification and taxonomic richness considerably affect the species range size. Finally, this work reveals poorly known patterns of range size variation and some of the mechanisms driving variation in range size and vulnerability to extinction.     

物种分布区范围地理格局的Rapoport法则

Rapoport法则最初是关于物种分布的纬度位置与纬度分布宽度的关系的假说, 被认为是宏观生态学的重要假说之一。Rapoport法则提出以后引发了广泛的实证检验和理论探讨, 并扩展到海拔和海洋深度梯度的分析。对Rapoport法则的检验发展了多种算法, 包括基于中域效应模型的比较分析法, 提出了气候变异性等多种机理解释, 并建立了物种分布区范围格局与物种丰富度格局、分布边界限制等重要宏观生态学现象之间的联系。关于该法则的普适性仍存在明显的意见分歧, 但分析和检验方法被认为是影响结果的重要因素, 方法的改进将是这一领域今后研究的关键。本文主要从Rapoport法则的概念、检验方法、机理性解释, 关于其普适性的争论及其与物种丰富度格局的关系等方面, 综述这一领域的研究进展。     

Thermal tolerance and preference of exploited turbinid snails near their range limit in a global warming hotspot

Predicted global climate change has prompted numerous studies of thermal tolerances of marine species. The upper thermal tolerance is unknown for most marine species, but will determine their vulnerability to ocean warming. Gastropods in the family Turbinidae are widely harvested for human consumption. To investigate the responses of turbinid snails to future conditions we determined critical thermal maxima (CTMax) and preferred temperatures of Turbo militaris and Lunella undulata from the tropical-temperate overlap region of northern New South Wales, on the Australian east coast. CTMax were determined at two warming rates: 1 °C/30 min and 1 °C/12 h. The number of snails that lost attachment to the tank wall was recorded at each temperature increment. At the faster rate, T. militaris had a significantly higher CTMax (34.0 °C) than L. undulata (32.2 °C). At the slower rate the mean of both species was lower and there was no significant difference between them (29.4 °C for T. militaris and 29.6 °C for L. undulata). This is consistent with differences in thermal inertia possibly allowing animals to tolerate short periods at higher temperatures than is possible during longer exposure times, but other mechanisms are not discounted. The thermoregulatory behaviour of the turban snails was determined in a horizontal thermal gradient. Both species actively sought out particular temperatures along the gradient, suggesting that behavioural responses may be important in ameliorating short-term temperature changes. The preferred temperatures of both species were higher at night (24.0 °C and 26.0 °C) than during the day (22.0 °C and 23.9 °C). As the snails approached their preferred temperature, net hourly displacement decreased. Preferred temperatures were within the average seasonal seawater temperature range in this region. However, with future predicted water temperature trends, the species could experience increased periods of thermal stress, possibly exceeding CTMax and potentially leading to range contractions.     

Colonization,abundance, and geographic range size of gravestone lichens

Macroecological studies often find that species with large geographic range sizes are also locally abundant. Superior colonization ability of species with large ranges is a possible/plausible explanation for this pattern, yet direct measures of colonization ability are difficult, and thus the relationship between colonization ability and range size is rarely investigated directly. Using a data set of gravestone lichens spanning more than 300 years, we investigated relationships among colonization ability, abundance, and geographic range size. Pairwise correlations were used to compare colonization ability and local abundance with area of occupancy (a measure of range size) and spore size within England, Scotland, and Wales on two different types of gravestones. Indices of colonization ability and abundance were positively correlated with area of occupancy. Colonization ability was significantly positively correlated with local abundance, but it was not at all related to propagule size. When lichen species were grouped categorically by colonization ability, the strongest area-occupancy relationships were observed within the subset of species that were the best colonizers. Significant differences among genera were found in spore size but not for other variables. Lichen species that occupy the largest geographic area were the best colonizers: they were the first species present on newly erected stones. These results complement the more commonly observed macroecological pattern that widespread species are also locally abundant.     

Null models of geographic range size evolution reaffirm its heritability

Most models of allopatric speciation predict that the two daughter species will have range sizes different from each other's and potentially from that of their common ancestor. However, I find that this difference is less than that expected under a variety of null models of range evolution. Sister species' range values may therefore become more similar in the time following speciation. Greater-than-expected similarity (symmetry) has also been treated as a form of range size heritability. I therefore compare the results of this symmetry approach to a test for phylogenetic signal, using the range sizes of North American birds. I find that range size is heritable under both tests. I suggest that null models for range size heritability should be informed by an explicit model of evolution. Comparative methods may give erroneous results if they fail to take the unusual form of inheritance of range size into account.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Interspecific patterns of species richness, geographic range size, and body size among New World venomous snakes

Many higher taxa exhibit latitudinal gradients in species richness, geographic range size, and body size. However, these variables are often interdependent, such that examinations of univariate or bivariate patterns alone may be misleading. Therefore, I examined latitudinal gradients in, and relationships between, species richness, geographic range size, and body size among 144 species of New World venomous snakes [families Elapidae (coral snakes) and Viperidae (pitvipers)]. Both lineages are monophyletic, collectively span 99° of latitude, and are extremely variable in body size and geographic range sizes. Coral snakes exhibit highest species richness near the equator, while pitviper species richness peaks in Central America. Species – range size distributions were strongly right-skewed for both families. There was little support for Bergmann's rule or Rapoport's rule for snakes of either family, as neither body size nor range size increased significantly with latitude. However, range area and median range latitude were positively correlated above 15° N, indicating a possible "Rapoport effect" at high northern latitudes. Geographic range size was positively associated with body size. Available continental area strongly influenced range size. Comparative (phylogenetically-based) analyses revealed that shared history is a poor predictor of range size variation within clades. Among vipers, trends in geographic range sizes may have been structured more by historical biogeography than by macroecological biotic factors.     

The significance of geographic range size for spatial diversity patterns in Neotropical palms

We examined the effect of range size in commonly applied macroecological analyses using continental distribution data for all 550 Neotropical palm species (Arecaceae) at varying grain sizes from 0.5° to 5°. First, we evaluated the relative contribution of range-restricted and widespread species on the patterns of species richness and endemism. Second, we analysed the impact of range size on the predictive value of commonly used predictor variables. Species sequences were produced arranging species according to their range size in ascending, descending, and random order. Correlations between the cumulative species richness patterns of these sequences and environmental predictors were performed in order to analyse the effect of range size. Despite the high proportion of rare species, patterns of species richness were found to be dominated by a minority of widespread species (∼20%) which contained 80% of the spatial information. Climatic factors related to energy and water availability and productivity accounted for much of the spatial variation of species richness of widespread species. In contrast, species richness of range-restricted species was to a larger extent determined by topographical complexity. However, this effect was much more difficult to detect due to a dominant influence of widespread species. Although the strength of different environmental predictors changed with spatial scale, the general patterns and trends proved to be relatively stabile at the examined grain sizes. Our results highlight the difficulties to approximate causal explanations for the occurrence of a majority of species and to distinguish between contemporary climatic factors and history.