Correlated evolution between climate and suites of traits along a fast–slow continuum in the radiation of Protea

Evolutionary radiations are responsible for much of Earth's diversity, yet the causes of these radiations are often elusive. Determining the relative roles of adaptation and geographic isolation in diversification is vital to understanding the causes of any radiation, and whether a radiation may be labeled as “adaptive” or not. Across many groups of plants, trait–climate relationships suggest that traits are an important indicator of how plants adapt to different climates. In particular, analyses of plant functional traits in global databases suggest that there is an “economics spectrum” along which combinations of functional traits covary along a fast–slow continuum. We examine evolutionary associations among traits and between trait and climate variables on a strongly supported phylogeny in the iconic plant genus Protea to identify correlated evolution of functional traits and the climatic‐niches that species occupy. Results indicate that trait diversification in Protea has climate associations along two axes of variation: correlated evolution of plant size with temperature and leaf investment with rainfall. Evidence suggests that traits and climatic‐niches evolve in similar ways, although some of these associations are inconsistent with global patterns on a broader phylogenetic scale. When combined with previous experimental work suggesting that trait–climate associations are adaptive in Protea, the results presented here suggest that trait diversification in this radiation is adaptive.     

What explains patterns of species richness? The relative importance of climatic‐niche evolution,morphological evolution,and ecological limits in salamanders

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic‐niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species‐rich family of salamanders. Earlier studies have suggested that climatic‐niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with “ecological limits” on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic‐niche evolution. Using phylogenetic multiple regression, we show that rates of climatic‐niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic‐niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns.     

Developmental life history is associated with variation in rates of climatic niche evolution in a salamander adaptive radiation*

Rates of climatic niche evolution vary widely across the tree of life and are strongly associated with rates of diversification among clades. However, why the climatic niche evolves more rapidly in some clades than others remains unclear. Variation in life history traits often plays a key role in determining the environmental conditions under which species can survive, and therefore, could impact the rate at which lineages can expand in available climatic niche space. Here, we explore the relationships among life-history variation, climatic niche breadth, and rates of climatic niche evolution. We reconstruct a phylogeny for the genus Desmognathus, an adaptive radiation of salamanders distributed across eastern North America, based on nuclear and mitochondrial genes. Using this phylogeny, we estimate rates of climatic niche evolution for species with long, short, and no aquatic larval stage. Rates of climatic niche evolution are unrelated to the mean climatic niche breadth of species with different life histories. Instead, we find that the evolution of a short larval period promotes greater exploration of climatic space, leading to increased rates of climatic niche evolution across species having this trait. We propose that morphological and physiological differences associated with variation in larval stage length underlie the heterogeneous ability of lineages to explore climatic niche space. Rapid rates of climatic niche evolution among species with short larval periods were an important dimension of the clade's adaptive radiation and likely contributed to the rapid rate of lineage accumulation following the evolution of an aquatic life history in this clade. Our results show how variation in a key life-history trait can constrain or promote divergence of the climatic niche, leading to variation in rates of climatic niche evolution among species.     

Correlated evolution of thermal niches and functional physiology in tropical freshwater fishes

The role of ecology in phenotypic and species diversification is widely documented. Nonetheless, numerous nonadaptive processes can shape realized niches and phenotypic variation in natural populations, complicating inferences about adaptive evolution at macroevolutionary scales. We tested for evolved differences in thermal tolerances and their association with the realized thermal niche (including metrics describing diurnal and seasonal patterns of temperature extremes and variability) across a genus of tropical freshwater fishes reared in a standardized environment. There was limited evolution along the thermal niche axis associated with variation in maximum temperature and in upper thermal limits. In contrast, there was considerable diversification along the first major axis of the thermal niche associated with minimum temperatures and in lower thermal limits. Across our adaptive landscape analyses, 70% of species exhibited evidence of divergence in thermal niches. Most importantly, the first two major axes of thermal niche variation were significantly correlated with variation in lower thermal limits. Our results indicate adaptation to divergent thermal niches and adaptive evolution of related functional traits, and highlight the importance of divergence in lower thermal limits for the evolution of tropical biodiversity.     

Highlighting convergent evolution in morphological traits in response to climatic gradient in African tropical tree species: The case of genus Guibourtia Benn.

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Phylogeny and biogeography of the American live oaks (Quercus subsection Virentes): a genomic and population genetics approach

The nature and timing of evolution of niche differentiation among closely related species remains an important question in ecology and evolution. The American live oak clade, Virentes, which spans the unglaciated temperate and tropical regions of North America and Mesoamerica, provides an instructive system in which to examine speciation and niche evolution. We generated a fossil‐calibrated phylogeny of Virentes using RADseq data to estimate divergence times and used nuclear microsatellites, chloroplast sequences and an intron region of nitrate reductase (NIA‐i3) to examine genetic diversity within species, rates of gene flow among species and ancestral population size of disjunct sister species. Transitions in functional and morphological traits associated with ecological and climatic niche axes were examined across the phylogeny. We found the Virentes to be monophyletic with three subclades, including a southwest clade, a southeastern US clade and a Central American/Cuban clade. Despite high leaf morphological variation within species and transpecific chloroplast haplotypes, RADseq and nuclear SSR data showed genetic coherence of species. We estimated a crown date for Virentes of 11 Ma and implicated the formation of the Sea of Cortés in a speciation event ~5 Ma. Tree height at maturity, associated with fire tolerance, differs among the sympatric species, while freezing tolerance appears to have diverged repeatedly across the tropical–temperate divide. Sympatric species thus show evidence of ecological niche differentiation but share climatic niches, while allopatric and parapatric species conserve ecological niches, but diverge in climatic niches. The mode of speciation and/or degree of co‐occurrence may thus influence which niche axis plants diverge along.     

Expanding the understanding of local community assembly in adaptive radiations

Communities are thought to be assembled by two types of filters: by the environment relating to the fundamental niche and by biotic interactions relating to the realized niche. Both filters include parameters related to functional traits and their variation along environmental gradients. Here, we infer the general importance of environmental filtering of a functional trait determining local community assembly within insular adaptive radiations on the example of Caribbean Anolis lizards. We constructed maps for the probability of presence of Anolis ecomorphs (ecology‐morphology‐behavior specialists) on the Greater Antilles and overlaid these to estimate ecomorph community completeness (ECC) over the landscape. We then tested for differences in environmental parameter spaces among islands for real and cross‐fitted ECC values to see whether the underlying assembly filters are deterministic (i.e., similar among islands). We then compared information‐theoretic models of climatic and landscape parameters among Greater Antillean islands and inferred whether body mass as functional trait determines ECC. We found areas with high ECC to be strongly correlated with environmental filters, partly related to elevation. The environmental parameters influencing high ECC differed among islands. With the exception of the Jamaican twig ecomorph (which we suspect to be misclassified), smaller ecomorphs were more restricted to higher elevations than larger ones which might reflect filtering on the basis of differential physiological restrictions of ecomorphs. Our results in Anolis show that local community assembly within adaptive island radiations of animals can be determined by environmental filtering of functional traits, independently from species composition and realized environmental niche space.     

Morphological innovation, diversification and invasion of a new adaptive zone

How ecological opportunity relates to diversification is a central question in evolutionary biology. However, there are few empirical examples of how ecological opportunity and morphological innovation open new adaptive zones, and promote diversification. We analyse data on diet, skull morphology and bite performance, and relate these traits to diversification rates throughout the evolutionary history of an ecologically diverse family of mammals (Chiroptera: Phyllostomidae). We found a significant increase in diversification rate driven by increased speciation at the most recent common ancestor of the predominantly frugivorous subfamily Stenodermatinae. The evolution of diet was associated with skull morphology, and morphology was tightly coupled with biting performance, linking phenotype to new niches through performance. Following the increase in speciation rate, the rate of morphological evolution slowed, while the rate of evolution in diet increased. This pattern suggests that morphology stabilized, and niches within the new adaptive zone of frugivory were filled rapidly, after the evolution of a new cranial phenotype that resulted in a certain level of mechanical efficiency. The tree-wide speciation rate increased non linearly with a more frugivorous diet, and was highest at measures of skull morphology associated with morphological extremes, including the most derived Stenodermatines. These results show that a novel stenodermatine skull phenotype played a central role in the evolution of frugivory and increasing speciation within phyllostomids.     

Pleistocene glacial cycles drove lineage diversification and fusion in the Yosemite toad (Anaxyrus canorus)

Species endemic to alpine environments can evolve via steep ecological selection gradients between lowland and upland environments. Additionally, many alpine environments have faced repeated glacial episodes over the past two million years, fracturing these endemics into isolated populations. In this “glacial pulse” model of alpine diversification, cycles of allopatry and ecologically divergent glacial refugia play a role in generating biodiversity, including novel admixed (“fused”) lineages. We tested for patterns of glacial pulse lineage diversification in the Yosemite toad (Anaxyrus [Bufo] canorus), an alpine endemic tied to glacially influenced meadow environments. Using double‐digest RADseq on populations densely sampled from a portion of the species range, we identified nine distinct lineages with divergence times ranging from 18 to 724 thousand years ago (ka), coinciding with multiple Sierra Nevada glacial events. Three lineages have admixed origins, and demographic models suggest these fused lineages have persisted throughout past glacial cycles. Directionality indices supported the hypothesis that some lineages recolonized Yosemite from east of the ice sheet, whereas other lineages remained in western refugia. Finally, refugial niche reconstructions suggest that low‐ and high‐elevation lineages have convergently adapted to similar climatic niches. Our results suggest glacial cycles and refugia may be important crucibles of adaptive diversity across deep evolutionary time.     

Climatic Niche Dynamics and Its Role in the Insular Endemism of <Emphasis Type="Italic">Anolis</Emphasis> Lizards

Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.     

Tests of the contribution of acclimation to geographic variation in water loss rates of the West Indian lizard <Emphasis Type="Italic">Anolis cristatellus</Emphasis>

Phenotypic plasticity can contribute to the process of adaptive radiation by facilitating population persistence in novel environments. West Indian Anolis lizards provide a classic example of an adaptive radiation, in which divergence has occurred along two primary ecological axes: structural microhabitat and climate. Adaptive plasticity in limb morphology is hypothesized to have facilitated divergence along the structural niche axis in Anolis, but very little work has explored plasticity in physiological traits. Here, we experimentally ask whether Puerto Rican Anolis cristatellus from mesic and xeric habitats differ in desiccation rates, and whether these lizards exhibit an acclimation response to changes in relative humidity. We first present microclimatic data collected at lizard perch sites that demonstrate that abiotic conditions experienced by lizards differ between mesic and xeric habitat types. In Experiment 1, we measured desiccation rates of lizards from both habitats maintained under identical laboratory conditions. This experiment demonstrated that desiccation rates differ between populations; xeric lizards lose water more slowly than mesic lizards. In Experiment 2, lizards from each habitat were either maintained under the conditions of Experiment 1, or under extremely low relative humidity. Desiccation rates did not differ between lizards from the same habitat maintained under different treatments and xeric lizards maintained lower desiccation rates than mesic lizards within each treatment. Our results demonstrate that A. cristatellus does not exhibit an acclimation response to abrupt changes of hydric conditions, and suggest that tropical Anolis lizards might be unable to exhibit physiological plasticity in desiccation rates in response to varying climatic conditions.     

ECOLOGICAL OPPORTUNITY AND THE RATE OF MORPHOLOGICAL EVOLUTION IN THE DIVERSIFICATION OF GREATER ANTILLEAN ANOLES

The pace of phenotypic diversification during adaptive radiation should decrease as ecological opportunity declines. We test this prediction using phylogenetic comparative analyses of a wide range of morphological traits in Greater Antillean Anolis lizards. We find that the rate of diversification along two important axes of Anolis radiation—body size and limb dimensions—decreased as opportunity declined, with opportunity quantified either as time elapsed in the radiation or as the diversity of competing anole lineages inferred to have been present on an island at different times in the past. Most previous studies of the ecological opportunity hypothesis have focused on the rate of species diversification; our results provide a complementary perspective, indicating that the rate of phenotypic diversification declines with decreasing opportunity in an adaptive radiation.     

Hierarchy in adaptive radiation: A case study using the Carnivora (Mammalia)

Simpson's “early burst” model of adaptive radiation was intended to explain the early proliferation of morphological and functional variation in diversifying clades. Yet, despite much empirical testing, questions remain regarding its frequency across the tree of life. Here, we evaluate the support for an early burst model of adaptive radiation in 14 ecomorphological traits plus body mass for the extant mammalian order Carnivora and its constituent families. We find strong support for early bursts of dental evolution, suggesting a classic Simpsonian adaptive radiation along dietary resource axes. However, the signal of this early burst is not consistently recovered in analyses at the family level, where support for a variety of different models emerges. Furthermore, we find no evidence for early burst–like dynamics in size–related traits, and Bayesian analyses of evolutionary correlations corroborate a decoupling of size and dental evolution, driven in part by dietary specialization. Our results are consistent with the perspective that trait diversification unfolds hierarchically, with early bursts restricted to traits associated with higher level niches, such as macrohabitat use or dietary strategy, and thus with the origins of higher taxa. The lack of support for early burst adaptive radiation in previous phylogenetic studies may be a consequence of focusing on low‐level niche traits (i.e., those associated with microhabitat use) in clades at shallow phylogenetic levels. A richer understanding of early burst adaptive radiation will require a renewed focus on functional traits and their evolution over higher level clades.     

Evolution of the climatic niche in scaly tree ferns (Cyatheaceae,Polypodiopsida)

Although climatic niche conservatism has been assumed by a large number of studies focused on climatic niche evolution, there are examples of climatic niche diversification and adaptation to changing climates. In this article, we reconstruct a climatic niche of scaly tree ferns (Cyatheaceae) using a rigorous analytical procedure which combines climatic niche modelling with reconstruction of continuous characters given a phylogenetic hypothesis. To estimate the limits to climatic niches of species, we used climate envelope modelling and ordination. Ancestral climatic niches of species were reconstructed by maximum likelihood and least‐squares analyses. We observed a trend towards niche conservatism with occasional events of niche transformations in scaly tree ferns. We discuss the implications of our study with respect to the potential and limitations for applications of niche modelling to evolutionary studies. We suggest that future studies of evolution of climatic niches could be considerably improved by employing approaches enabling reconstruction of continuous response to climatic gradients. Further progress may also be achieved by exploring models of character evolution other than the Brownian motion model. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 1–19.     

ADAPTATION AND CONSTRAINT IN THE EVOLUTION OF SPECIALIZATION OF BAHAMIAN ANOLIS LIZARDS

Interspecific interactions affect habitat use and subsequent morphological adaptation in Anolis lizards. We examined populations of two species of Anolis lizards that evolved in the species-rich communities of Cuba and are now widespread in the Bahamas. Because the species occupy islands in the Bahamas that vary in the number of lizard species present and other characteristics, we predicted that directional selection should have led to morphological differentiation. In particular, we expected that populations on one-species islands should have evolved toward a generalist morphology because of the lack of competitors. Divergence in both species has been adaptive; populations that use wider perches have longer legs. Nonetheless, these differences are relatively minor, and none of the populations appears to have differentiated from its ancestral “ecomorph” type toward a more generalized morphology. This stasis mirrors a trend observed in the radiation of Caribbean anoles, which exhibits repeated instances of evolutionary specialization, but few or no cases of reversion to a more generalized condition. The explanation for this directionality of evolution is not obvious but probably involves the tendency of specialized species to continue using and further adapting the niches for which they are specialized even as conditions change.     

MORPHOLOGICAL DIVERSIFICATION AND ADAPTIVE RADIATION: A COMPARISON OF TWO DIVERSE LIZARD CLADES

We compared the morphological diversity (i.e., the amount of morphological space occupied) of two similar clades, the lizard genera Anolis and Sceloporus. These species-rich monophyletic clades are similar in body size, age of origin, and many aspects of their natural history. We examined a number of morphological traits whose variation is likely to represent adaptation to different aspects of the environment, including body size, limb proportions, head dimensions, and tail length. Examination of the position of species in multidimensional space, based on a principal components analysis, indicates that the morphological diversity of Anolis, which we refer to as disparity, is significantly greater than that of Sceloporus. One potential explanation for this pattern is that morphological diversification in Anolis was facilitated by the evolution of subdigital toe-pads, which allow anoles to use the environment in ways not available to Sceloporus. The geographic location of diversification (tropical and subtropical for Anolis, arid for Sceloporus) may also have been important.     

The thermal niche of Neotropical nectar‐feeding bats: Its evolution and application to predict responses to global warming

The thermal niche of a species is one of the main determinants of its ecology and biogeography. In this study, we determined the thermal niche of 23 species of Neotropical nectar‐feeding bats of the subfamily Glossophaginae (Chiroptera, Phyllostomidae). We calculated their thermal niches using temperature data obtained from collection records, by generating a distribution curve of the maximum and minimum temperatures per locality, and using the inflection points of the temperature distributions to estimate the species optimal (STZ) and suboptimal (SRZ) zones of the thermal niche. Additionally, by mapping the values of the STZ and SRZ on a phylogeny of the group, we generated a hypothesis of the evolution of the thermal niches of this clade of nectar‐feeding bats. Finally, we used the characteristics of their thermal niches to predict the responses of these organisms to climate change. We found a large variation in the width and limits of the thermal niches of nectar‐feeding bats. Additionally, while the upper limits of the thermal niches varied little among species, their lower limits differ wildly. The ancestral reconstruction of the thermal niche indicated that this group of Neotropical bats evolved under cooler temperatures. The two clades inside the Glossophaginae differ in the evolution of their thermal niches, with most members of the clade Choeronycterines evolving “colder” thermal niches, while the majority of the species in the clade Glossophagines evolving “warmer” thermal niches. By comparing thermal niches with climate change models, we found that all species could be affected by an increase of 1°C in temperature at the end of this century. This suggests that even nocturnal species could suffer important physiological costs from global warming. Our study highlights the value of scientific collections to obtain ecologically significant physiological data for a large number of species.     

Climatic niche evolution in turtles is characterized by phylogenetic conservatism for both aquatic and terrestrial species

Understanding how the climatic niche of species evolved has been a topic of high interest in current theoretical and applied macroecological studies. However, little is known regarding how species traits might influence climatic niche evolution. Here, we evaluated patterns of climatic niche evolution in turtles (tortoises and freshwater turtles) and whether species habitat (terrestrial or aquatic) influences these patterns. We used phylogenetic, climatic and distribution data for 261 species to estimate their climatic niches. Then, we compared whether niche overlap between sister species was higher than between random species pairs and evaluated whether niche optima and rates varied between aquatic and terrestrial species. Sister species had higher values of niche overlap than random species pairs, suggesting phylogenetic climatic niche conservatism in turtles. The climatic niche evolution of the group followed an Ornstein–Uhlenbeck model with different optimum values for aquatic and terrestrial species, but we did not find consistent evidence of differences in their rates of climatic niche evolution. We conclude that phylogenetic climatic niche conservatism occurs among turtle species. Furthermore, terrestrial and aquatic species occupy different climatic niches but these seem to have evolved at similar evolutionary rates, reinforcing the importance of habitat in understanding species climatic niches and their evolution.     

The role of pollinators in floral diversification in a clade of generalist flowers

Pollinator‐mediated evolutionary divergence has seldom been explored in generalist clades because it is assumed that pollinators in those clades exert weak and conflicting selection. We investigate whether pollinators shape floral diversification in a pollination generalist plant genus, Erysimum. Species from this genus have flowers that appeal to broad assemblages of pollinators. Nevertheless, we recently reported that it is possible to sort plant species into pollination niches varying in the quantitative composition of pollinators. We test here whether floral characters of Erysimum have evolved as a consequence of shifts among pollination niches. For this, we quantified the evolutionary lability of the floral traits and their phylogenetic association with pollination niches. As with pollination niches, Erysimum floral traits show weak phylogenetic signal. Moreover, floral shape and color are phylogenetically associated with pollination niche. In particular, plants belonging to a pollination niche dominated by long‐tongued large bees have lilac corollas with parallel petals. Further analyses suggest, however, that changes in color preceded changes in pollination niche. Pollinators seem to have driven the evolution of corolla shape, whereas the association between pollination niche and corolla color has probably arisen by lilac‐flowered Erysimum moving toward certain pollination niches for other adaptive reasons.     

Ecological and phenotypic divergence in Iberian shrews (Soricidae)

There are examples of coexisting species with similar morphology and ecology, in apparent contradiction to competition theory. Shrews (Soricidae) are a paradigmatic example of this because members of this group exhibit a conserved body form, relatively low variability in lifestyle, and, in many cases, a sympatric distribution. Here, we combined geometric morphometrics and ecological niche modeling to test whether diversification of soricid species inhabiting the Iberian Peninsula has been driven by niche divergence or, conversely, whether niche conservatism has played a paramount role in this process. We also examined whether pairwise morphological distances increase as the degree of niche overlap between species becomes greater, as would be expected if interspecific competition promotes morphological differentiation. Our results showed that water shrews (Neomys), white‐toothed shrews (Crocidurinae), and red‐toothed shrews (Soricinae) are clearly differentiated in terms of both skull shape and mandible shape. However, we found a lack of phylogenetic signal in most morphological traits, indicating that closely related species are not more similar than expected by random chance. Notably, water shrews show a more “triangular” or sharp skull than white‐toothed and red‐toothed shrews, probably as an adaptation to their semiaquatic lifestyle. In agreement with the phenotypic data, climatic traits (mean annual temperature and annual precipitation) were highly labile and sister taxa showed extensive differentiation in their realized niche space. Finally, we found that phenotypic distances between species tend to increase as the degree of niche overlap increases, suggesting that interspecific competition is an important factor in determining the level of morphological resemblance among relatives. Overall, our results indicate that the existence of limited morphological disparity in a given group does not necessarily imply the existence of a niche conservatism signature.