Niche position and niche breadth effects on population abundances: A case study of New World Warblers (Parulidae).

Species abundance patterns are influenced by a myriad of factors, including habitat availability and ecological niche characteristics. However, the evidence concerning the specific impact factors such as niche position and niche breadth on mean and maximum abundances in vertebrates at a broad geographical scale remains inconclusive. In this study, we investigated the influence of niche position and breadth on the abundance of 47 species of birds belonging to the Parulidae family, commonly known as New World Warblers. We obtained data on abundance and presence records spanning the reproductive distribution of these species and employed the outlying mean index analysis to calculate niche position and niche breadth. We assessed the relationship between abundance metrics and niche descriptors using phylogenetic regressions to account for the non-independence resulting from phylogenetic ancestry. Initially, we developed individual models for each predictor and subsequently formulated a multi-predictor model encompassing niche position, niche breadth, and their interaction. Our findings revealed a negative relationship between niche position and both mean and maximum abundance, while niche breadth exhibited a positive relationship with these niche characteristics. Notably, the results of the multi-predictor models indicated that niche position exerted the most substantial influence on both mean and maximum abundance. Additionally, the interaction between niche position and niche breadth had the most positive and significant contribution to mean population abundance. This study underscores the need for future research in other vertebrates to delve into the mechanisms underlying these patterns. Such endeavors will not only enhance our understanding of ecological dynamics but also equip us with predictive capabilities to anticipate population responses to environmental changes effectively.

High Investment in Reproduction Is Associated with Reduced Life Span in Dogs.

AbstractProminent differences in aging among and within species present an evolutionary puzzle. The theories proposed to explain evolutionary differences in aging are based on the axiom that selection maximizes fitness, not necessarily life span. This implies trade-offs between investment in self-maintenance and investment in reproduction, where high investments in growth and current reproduction are associated with short life spans. Fast growth and large adult size are related to shorter life spans in the domestic dog, a bourgeoning model in aging research; however, whether reproduction influences life span in this system remains unknown. Here we test the relationship between reproduction and differences in life span among dog breeds, simultaneously controlling for shared ancestry and recent gene flow. We found that shared ancestry explains a higher proportion of the among-breed variation in life history traits, in comparison with recent gene flow. Our results also show that reproductive investment negatively impacts life span, and more strongly so in large breeds, an effect that is not merely a correlated response of adult size. These results suggest that basic life history trade-offs are apparent in a domestic animal whose diversity is the result of artificial selection and that among-breed differences in life span are due to a combination of size and reproduction.

Diversification dynamics in Caribbean rain frogs (Eleutherodactylus) are uncoupled from the anuran community and consistent with adaptive radiation.

Adaptive radiation is proposed to play a key role in generating differences in species richness among lineages and geographical regions. Due to the importance of ecological divergence in adaptive radiation, species richness is predicted to be influenced by equilibrium diversity dynamics, although the concept continues to generate much debate. An additional important question is whether radiating clades have intrinsic biological characteristics that make them particularly prone to diversify. We tackle these questions by analysing (i) the temporal patterns of diversification of Caribbean Eleutherodactylus frogs, and (ii) assembly of the complete native anuran community of the Caribbean archipelago (197 species), testing for the presence of equilibrium dynamics and whether diversification patterns of Eleutherodactylus differ from those of the rest of the Caribbean anurans. Diversification rates follow the predicted pattern of rapid diversification early in the radiation which gradually decreases towards the present. Eleutherodactylus diversification is significantly faster than that of the Caribbean anuran community, and although equilibrium dynamics influence richness of all Caribbean anurans, Eleutherodactylus shows higher carrying capacity. Our results indicate that ecological opportunity per se is not sufficient for adaptive radiation and that diverse lineages present intrinsic characteristics that enable them to make the most of available opportunity.

The Phylogenetic Limits to Diversity-Dependent Diversification.

While the theory of micro-evolution by natural selection assigns a crucial role to competition, its role in macroevolution is less clear. Phylogenetic evidence for a decelerating accumulation of lineages suggests a feedback of lineage diversity on diversification. However, does this feedback only occur between close relatives, or do distant relatives also influence each other's diversification? In other words: are there phylogenetic limits to this diversity-dependence? Islands form ideal systems to answer these questions because their boundedness facilitates an overview of all potential competitors. The DAISIE (Dynamic Assembly of Island biota through Speciation Immigration and Extinction) framework allows for testing the presence of diversity-dependence on islands given phylogenetic data on colonization and branching times. The current inference models in DAISIE assume that this diversity-dependence only applies within a colonizing clade, i.e., all mainland species can colonize and diversify independently from one another. We term this clade-specific (CS) diversity-dependence. Here we introduce a new DAISIE model that assumes that diversity-dependence applies to all island species of a taxonomic group regardless of their mainland ancestry, i.e., diversity-dependence applies both to species within the same clade and between different clades established by different mainland species. We call this island-wide (IW) diversity-dependence. We present a method to compute a likelihood for this model given phylogenetic data on colonization and branching events and use likelihood ratio bootstrapping to compare it to the likelihood of the CS model in order to overcome biases known for standard model selection. We apply it to the diversification of Eleutherodactylus frogs on Hispaniola. Across the Greater Antilles archipelago, this radiation shows repeated patterns of diversification in ecotypes that are similar across clades. This could be suggestive of overlapping niche space and hence between-clade interactions, i.e., IW diversity-dependence. But it could also be suggestive of only within-clade interactions because between-clade interactions would have blocked the same ecotype from re-appearing. We find that the CS model fits the data much better than the IW model, indicating that different colonizations while resulting in similar ecotypes, are sufficiently distinct to avoid interacting strongly. We argue that non-overlapping distributions between clades (both spatially and in terms of ecotypes) cannot be used as evidence of CS diversity-dependence, because this pattern may be a consequence of IW diversity-dependence. By contrast, by using phylogenetic data rather than distributional data our method does allow for inferring the phylogenetic limits to diversity-dependent diversification. We discuss possibilities for future extensions and applications of our modelling approach. [Adaptive radiation; birth-death model; Caribbean; diversity-dependence; Eleutherodactylus; island biogeography.].

Influence of microhabitat, fecundity, and parental care on the evolution of sexual size dimorphism in Caribbean Eleutherodactylus frogs.

Rensch's rule suggests that sexual size dimorphism (SSD) increases with species size when males are the larger sex, whereas it decreases when females are the larger sex. However, the process responsible for this pattern remains obscure. SSD can result from sexual selection, such as intrasexual competition for access to mates, or from natural selection, due to resource partitioning or fecundity selection. We studied SSD in Caribbean Eleutherodactylus frogs using phylogenetic comparative methods to investigate the influence of microhabitat, fecundity, and parental care. Our results show that in Caribbean Eleutherodactylus females tend to be larger and, contrary to Rensch's rule, dimorphism increases with species size. SSD was not related to microhabitat use. However, SSD was positively correlated with fecundity, mediated by a greater increase in female size. SSD was also influenced by parental care, suggesting that male care promotes larger male size and reduces the female bias in SSD. As suggested for other anurans, female-biased SSD in Caribbean Eleutherodactylus results from fecundity selection, although the magnitude is countered by increased male size in species with paternal care. Our results highlight the importance of considering various selective forces that may act in concert to influence the evolution of SSD.

Sex roles in birds: Phylogenetic analyses of the influence of climate, life histories and social environment.

Sex roles describe sex differences in courtship, mate competition, social pair-bonds and parental care. A key challenge is to identify associations among the components and the drivers of sex roles. Here, we investigate sex roles using data from over 1800 bird species. We found extensive variation and lability in proxies of sex roles, indicating remarkably independent evolution among sex role components. Climate and life history showed weak associations with sex roles. However, adult sex ratio is associated with sexual dimorphism, mating system and parental care, suggesting that social environment is central to explaining variation in sex roles among birds. Our results suggest that sex differences in reproductive behaviour are the result of diverse and idiosyncratic responses to selection. Further understanding of sex roles requires studies at the population level to test how local responses to ecology, life histories and mating opportunities drive processes that shape sex role variation among higher taxa.

The relative effects of pace of life-history and habitat characteristics on the evolution of sexual ornaments: A comparative assessment.

Selection may favor greater investment into sexual ornaments when opportunities for future reproduction are limited, for example, under high adult mortality. However, predation, a key driver of mortality, typically selects against elaborate sexual ornaments. Here, we examine the evolution of sexual ornaments in killifishes, which have marked contrasts in life-history strategy among species and inhabit environments that differ in accessibility to aquatic predators. We first assessed if the size of sexual ornaments (unpaired fins) influenced swimming performance. Second, we investigated whether the evolution of larger ornamental fins is driven primarily by the pace of life-history (investment into current vs. future reproduction) or habitat type (a proxy for predation risk). We found that larger fins negatively affected swimming performance. Further, males from species inhabiting ephemeral habitats, with lower predation risk, had larger fins and greater sexual dimorphism in fin size, compared to males from more accessible permanent habitats. We show that enlarged ornamental fins, which impair locomotion, evolve more frequently in environments that are less accessible to predators, without clear associations to life-history strategy. Our results provide a rare link between the evolution of sexual ornaments, effects on locomotion performance, and natural selection on ornament size potentially through habitat differences in predation risk.

A comment on The adaptive value of gluttony: predators mediate the life history trade-offs of satiation threshold by Pruitt & Krauel (2010).

Inspection of the data that accompany Pruitt and Krauel's study of individual variation in satiation threshold and a comparison of these data with the Materials and Methods and Results sections of the paper have revealed a number of issues that cast doubts on the reliability of the data and any results based on these data. In particular, we show that, following our analyses, the data are unlikely to have been obtained using the study design outlined in the publication and that statistical analyses of these data provide results that differ in important ways from those reported. These findings illustrate the importance of making raw data and analysis code available for the rigour and reproducibility of the scientific literature.

Fast life-histories are associated with larger brain size in killifishes.

The high energetic demands associated with the vertebrate brain are proposed to result in a trade-off between the pace of life-history and relative brain size. However, because both life-history and brain size also have a strong relationship with body size, any associations between the pace of life-history and relative brain size may be confounded by coevolution with body size. Studies on systems where contrasts in the pace of life-history occur without concordant contrasts in body size could therefore add to our understanding of the potential coevolution between relative brain size and life-history. Using one such system - 21 species of killifish - we employed a common garden design across two ontogenetic stages to investigate the association between relative brain size and the pace of life-history. Contrary to predictions, we found that relative brain size was larger in adult fast-living killifishes, compared to slow-living species. Although we found no differences in relative brain size between juvenile killifishes. Our results suggest that fast- and slow-living killifishes do not exhibit the predicted trade-off between brain size and life-history. Instead, fast and slow-living killifishes could differ in the ontogenetic timing of somatic versus neural growth or inhabit environments that differ considerably in cognitive demands.

Environmental heterogeneity shapes physiological traits in tropical direct-developing frogs.

Tropical ectotherm species tend to have narrower physiological limits than species from temperate areas. As a consequence, tropical species are considered highly vulnerable to climate change since minor temperature increases can push them beyond their physiological thermal tolerance. Differences in physiological tolerances can also be seen at finer evolutionary scales, such as among populations of ectotherm species along elevation gradients, highlighting the physiological sensitivity of such organisms.Here, we analyze the influence of elevation and bioclimatic domains, defined by temperature and precipitation, on thermal sensitivities of a terrestrial direct-developing frog (Craugastor loki) in a tropical gradient. We address the following questions: (a) Does preferred temperature vary with elevation and among bioclimatic domains? (b) Do thermal tolerance limits, that is, critical thermal maximum and critical thermal minimum vary with elevation and bioclimatic domains? and (c) Are populations from high elevations more vulnerable to climate warming?We found that along an elevation gradient body temperature decreases as environmental temperature increases. The preferred temperature tends to moderately increase with elevation within the sampled bioclimatic domains. Our results indicate that the ideal thermal landscape for this species is located at midelevations, where the thermal accuracy (db ) and thermal quality of the environment (de ) are suitable. The critical thermal maximum is variable across elevations and among the bioclimatic domains, decreasing as elevation increases. Conversely, the critical thermal minimum is not as variable as the critical thermal maximum.Populations from the lowlands may be more vulnerable to future increases in temperature. We highlight that the critical thermal maximum is related to high temperatures exhibited across the elevation gradient and within each bioclimatic domain; therefore, it is a response to high environmental temperatures.

The role of brain size on mammalian population densities.

The local abundance or population density of different organisms often varies widely. Understanding what determines this variation is an important, but not yet fully resolved question in ecology. Differences in population density are partly driven by variation in body size and diet among organisms. Here we propose that the size of an organism' brain could be an additional, overlooked, driver of mammalian population densities. We explore two possible contrasting mechanisms by which brain size, measured by its mass, could affect population density. First, because of the energetic demands of larger brains and their influence on life history, we predict mammals with larger relative brain masses would occur at lower population densities. Alternatively, larger brains are generally associated with a greater ability to exploit new resources, which would provide a competitive advantage leading to higher population densities among large-brained mammals. We tested these predictions using phylogenetic path analysis, modelling hypothesized direct and indirect relationships between diet, body mass, brain mass and population density for 656 non-volant terrestrial mammalian species. We analysed all data together and separately for marsupials and the four taxonomic orders with most species in the dataset (Carnivora, Cetartiodactyla, Primates, Rodentia). For all species combined, a single model was supported showing lower population density associated with larger brains, larger bodies and more specialized diets. The negative effect of brain mass was also supported for separate analyses in Primates and Carnivora. In other groups (Rodentia, Cetartiodactyla and marsupials) the relationship was less clear: supported models included a direct link from brain mass to population density but 95% confidence intervals of the path coefficients overlapped zero. Results support our hypothesis that brain mass can explain variation in species' average population density, with large-brained species having greater area requirements, although the relationship may vary across taxonomic groups. Future research is needed to clarify whether the role of brain mass on population density varies as a function of environmental (e.g. environmental stability) and biotic conditions (e.g. level of competition).

Long life evolves in large-brained bird lineages.

The brain is an energetically costly organ that consumes a disproportionate amount of resources. Species with larger brains relative to their body size have slower life histories, with reduced output per reproductive event and delayed development times that can be offset by increasing behavioral flexibility. The "cognitive buffer" hypothesis maintains that large brain size decreases extrinsic mortality due to greater behavioral flexibility, leading to a longer lifespan. Alternatively, slow life histories, and long lifespan can be a pre-adaptation for the evolution of larger brains. Here, we use phylogenetic path analysis to contrast different evolutionary scenarios and disentangle direct and indirect relationships between brain size, body size, life history, and longevity across 339 altricial and precocial bird species. Our results support both a direct causal link between brain size and lifespan, and an indirect effect via other life history traits. These results indicate that large brain size engenders longer life, as proposed by the "cognitive buffer" hypothesis.

Macroevolutionary evidence suggests trait-dependent coevolution between behavior and life-history.

Species with fast life-histories typically prioritize current over future reproductive events, compared to species with slow life-histories. These species therefore require greater energetic input into reproduction, and also likely have less time to realize their reproductive potential. Hence, behaviors that increase access to both resources and mating opportunities, at a cost of increased mortality risk, could coevolve with the pace of life-history. However, whether this prediction holds across species, remains untested under standardized conditions. Here, we test how risky behaviors, which facilitate access to resources and mating opportunities (i.e., activity, boldness, and aggression), along with metabolic rate, coevolve with the pace of life-history across 20 species of killifish that present remarkable divergences in the pace of life-history. We found a positive association between the pace of life-history and aggression, but interestingly not with other behavioral traits or metabolic rate. Aggression is linked to interference competition, and in killifishes is often employed to secure mates, while activity and boldness are more relevant for exploiting energetic resources. Our results suggest that the trade-off between current and future reproduction plays a more prominent role in shaping mating behavior, while behaviors related to energy acquisition may be influenced by ecological factors.

Sexually selected sexual selection: Can evolutionary retribution explain female ornamental colour?

By preferring mates with increasingly costly ornaments or courtship displays, females cause an escalation of male reproductive costs. Such increased costs should promote male selectivity based on fecundity-linked female attributes, leading to female ornamentation in species with traditional sex roles. Consequently, female ornamentation should evolve more frequently in taxa where male reproduction is costly than in comparable taxa where it is cheaper. We assessed the prevalence of female ornamental colouration in two clades of viviparous cyprinodontid fish: the Goodeinae, where stringent female choice imposes male mating costs, and the Poeciliinae, whose males can circumvent female mate choice. We found that although in the Poeciliinae female ornamental colour is a correlated, but paler version of male coloration, females of the Goodeinae often display vivid ornamental colours that are distinct from those of males. Thus, male and female ornaments are not (phylo)genetically correlated in the Goodeinae. Furthermore, phylogenetic signal on male and female colour is clearly detectable in the Poeciliinae, but absent in the Goodeinae, suggesting that ornamental colour of males and females in the latter may be the consequence of selection. Given that enforceable female choice has promoted male ornaments, we propose that evolutionary retribution has promoted distinct female ornaments in the Goodeinae.

Ecomorphological convergence in Eleutherodactylus frogs: a case of replicate radiations in the Caribbean.

Replicate radiations, the repeated multiplication of species associated with ecological divergence, have attracted much attention and generated as much debate. Due to the few well-studied cases, it remains unclear whether replicate radiations are an exceptional result of evolution or a relatively common example of the power of adaptation by natural selection. We examined the case of Eleutherodactylus frogs, which radiated in the Caribbean islands resulting in more than 160 species that occupy very diverse habitats. A time-calibrated phylogeny revealed that these frogs independently diversified on all larger islands producing species that occupy a broad range of microhabitats in different islands. Using phylogenetic comparative methods, we found an association between morphological traits and particular microhabitats, and for most microhabitats detected significant morphological convergence. Our results indicate Caribbean Eleutherodactylus are a novel example of replicate radiations, and highlight the predictability of evolutionary processes, as similar ecological opportunities can lead to similar outcomes.

Neotropical frogs and mating songs: The evolution of advertisement calls in glassfrogs.

Anurans emit advertisement calls with the purpose of attracting mates and repelling conspecific competitors. The evolution of call traits is expected to be associated with the evolution of anatomical and behavioural traits due to the physics of call emission and transmission. The evolution of vocalizations might imply trade-offs with other energetically costly behaviours, such as parental care. Here, we investigated the association between body size, calling site, parental care and call properties (call duration, number of notes, peak frequency, frequency bandwidth and call structure) of the advertisement calls of glassfrogs (Centrolenidae)-a family of Neotropical, leaf-dwelling anurans-using phylogenetic comparative methods. We also explored the tempo and mode of evolution of these traits and compared them with those of three morphological traits associated with body size, locomotion and feeding. We generated and compiled acoustic data for 72 glassfrog species (46% of total species richness), including representatives of all genera. We found that almost all acoustic traits have significant, but generally modest, phylogenetic signal. Peak frequency of calls is significantly associated with body size, whereas call structure is significantly associated with calling site and paternal care. Thus, the evolution of body size, calling site and paternal care could constrain call evolution. The estimated disparity of acoustic traits was larger than that of morphological traits and the peak in disparity of acoustic traits generally occurred later in the evolution of glassfrogs, indicating a historically recent outset of the acoustic divergence in this clade.

One strategy does not fit all: determinants of urban adaptation in mammals.

Urbanisation exposes wildlife to new challenging conditions and environmental pressures. Some mammalian species have adapted to these novel environments, but it remains unclear which characteristics allow them to persist. To address this question, we identified 190 mammals regularly recorded in urban settlements worldwide, and used phylogenetic path analysis to test hypotheses regarding which behavioural, ecological and life history traits favour adaptation to urban environments for different mammalian groups. Our results show that all urban mammals produce larger litters; whereas other traits such as body size, behavioural plasticity and diet diversity were important for some but not all taxonomic groups. This variation highlights the idiosyncrasies of the urban adaptation process and likely reflects the diversity of ecological niches and roles mammals can play. Our study contributes towards a better understanding of mammal association to humans, which will ultimately allow the design of wildlife-friendly urban environments and contribute to mitigate human-wildlife conflicts.

Time-limited environments affect the evolution of egg-body size allometry.

Initial offspring size is a fundamental component of absolute growth rate, where large offspring will reach a given adult body size faster than smaller offspring. Yet, our knowledge regarding the coevolution between offspring and adult size is limited. In time-constrained environments, organisms need to reproduce at a high rate and reach a reproductive size quickly. To rapidly attain a large adult body size, we hypothesize that, in seasonal habitats, large species are bound to having a large initial size, and consequently, the evolution of egg size will be tightly matched to that of body size, compared to less time-limited systems. We tested this hypothesis in killifishes, and found a significantly steeper allometric relationship between egg and body sizes in annual, compared to nonannual species. We also found higher rates of evolution of egg and body size in annual compared to nonannual species. Our results suggest that time-constrained environments impose strong selection on rapidly reaching a species-specific body size, and reproduce at a high rate, which in turn imposes constraints on the evolution of egg sizes. In combination, these distinct selection pressures result in different relationships between egg and body size among species in time-constrained versus permanent habitats.

Cryptic within cryptic: genetics, morphometrics, and bioacoustics delimitate a new species of Eleutherodactylus (Anura: Eleutherodactylidae) from Eastern Cuba.

We studied the variation in genetics, bioacustics, and morphology in Eleutherodactylus glamyrus, a regionally endemic frog species restricted to high elevations in the Sierra Maestra Massif, Western Cuba that was originally described as a cryptic species hidden under the name E. auriculatus. Genetic analysis of mtDNA sequences of the 16S and cob genes identify two allopatric and strongly supported mitochondrial clades (phylogroups) which also showed no haplotype sharing in the nuclear Rag-1 gene. Bioacustic, and morphological comparisons concordantly identify these two phylogroups as independent evolutionary lineages. Therefore, we herein restrict the name Eleutherodactylus glamyrus Estrada and Hedges to populations represented in our analyses as the western phylogroup (Cordillera del Turquino to Pico La Bayamesa) and consider specimens from the eastern phylogroup (Sierra del Cobre) to represent a new species described and named as Eleutherodactylus cattus. Our results add to the growing list of Eleutherodactylus species endemic to Cuba and highlight the importance of combining different sources of evidence for obtaining robust assessments of species limits in amphibians.