The evolutionary ecology of interactive synchronism: The illusion of the optimal phenotype

In this article, we discuss some ecological-evolutionary strategies that allow synchronization of organisms, resources, and conditions. Survival and reproduction require synchronization of life cycles of organisms with favourable environmental and ecological features and conditions. This interactive synchronization can occur directly, through pairwise or diffuse co-evolution, or indirectly, for example, as a result of actions of ecosystem engineers and facilitator species. Observations of specific interactions, especially those which have coevolved, may give the false impression that evolution results in optimal genotypes or phenotypes. However, some phenotypes may arise under evolutionary constraints, such as simultaneous evolution of multiple traits, lack of a chain of fit transitional forms leading to an optimal phenotype, or by limits inherent in the process of selection, set by the number of selective deaths and by interference between linked variants. Although there are no optimal phenotypes, optimization models applied to particular species may be useful for a better understanding of the nature of adaptations. The evolution of adaptive strategies results in variable life histories. These strategies can minimize adverse impacts on the fitness of extreme or severe environmental conditions on survival and reproduction, and may include reproductive strategies such as semelparity and iteroparity, or morphological, physiological, or behavioural traits such as diapause, seasonal polyphenism, migration, or bet-hedging. However, natural selection cannot indefinitely maintain intra-population variation, and lack of variation can ultimately extinguish populations

Hierarchical effects on body size evolution and the macroecology of South American rainforest mammals

The relationship between geographic range size and body size analyzed at the continental scale for many species has been described recently by an envelope region defined in bivariate space and limited by ecological and physical constraints. These constraints can be explained by stochastic extinction related to minimum population density and energetic theories for an optimal body size. However, since data are obtained for different species in a large taxonomic group, it is possible that these variables may be strongly affected by phylogenetic autocorrelation. In this paper, we analyzed data on geographic range size and body size for 87 species of Neotropical rainforest mammals from South America, searching for phylogenetic effects changing the shape of constraint space. We used a nested ANOVA design to evaluate and remove large scale hierarchical (phylogenetic) trends in the two variables. Body size was more affected by phylogenetic effects than geographic range size. After removing these effects, we noted that the two macroecological variables analyzed support the shape of constraint space previously observed in North American species. The most important aspect of our analyses is that a model for optimal body size related to life-history variations provides an useful theoretical framework to understand how processes of species dynamics can lead to the observed macroecological patterns

A review of techniques for spatial modeling in geographical, conservation and landscape genetics

Most evolutionary processes occur in a spatial context and several spatial analysis techniques have been employed in an exploratory context. However, the existence of autocorrelation can also perturb significance tests when data is analyzed using standard correlation and regression techniques on modeling genetic data as a function of explanatory variables. In this case, more complex models incorporating the effects of autocorrelation must be used. Here we review those models and compared their relative performances in a simple simulation, in which spatial patterns in allele frequencies were generated by a balance between random variation within populations and spatially-structured gene flow. Notwithstanding the somewhat idiosyncratic behavior of the techniques evaluated, it is clear that spatial autocorrelation affects Type I errors and that standard linear regression does not provide minimum variance estimators. Due to its flexibility, we stress that principal coordinate of neighbor matrices (PCNM) and related eigenvector mapping techniques seem to be the best approaches to spatial regression. In general, we hope that our review of commonly used spatial regression techniques in biology and ecology may aid population geneticists towards providing better explanations for population structures dealing with more complex regression problems throughout geographic space

Exploring patterns in macroecological traits using sequential phylogenetic eigenvector regression

Diferentes métodos han sido propuestos para evaluar el grado de señal filogenética (autocorrelación) en los rasgos macroecológicos. Estos métodos son útiles para desarrollar maneras alternativas de evitar el problema de la falta de independencia entre especies y, actualmente, han demostrado ser importantes para inferir la velocidad de evolución de un rasgo al compararlo con modelos evolutivos alternativos, tales como el movimiento Browniano o el proceso Ornstein-Uhlenbeck (OU). Recientemente, desarrollamos un método llamado Curva de Señal-Representación Filogenética (PSR, por sus siglas en inglés), una expansión de la regresión de autovectores filogenéticos (PVR, por sus siglas en inglés) propuesto en 1998, que consiste en estimar diferentes coeficientes de determinación mediante la regresión de un rasgo de interés contra los autovectores extraídos de una matriz de distancias filogenéticas. El primer modelo únicamente utiliza los primeros de estos autovectores como variable explicativa, el segundo modelo utiliza tanto los primeros como los segundos autovectores y así sucesivamente. Posteriormente, los coeficientes de determinación resultantes son graficados contra los autovalores acumulados y la forma de esta curva es relacionada con modelos evolutivos determinando la variación del rasgo (i.e. un patrón lineal es esperado bajo movimiento Browniano). Aquí, aplicamos la curva PSR para estudiar los patrones de variación interespecífica en la talla corporal y tamaño de área de distribución de los carnívoros a nivel mundial y comparamos dichas curvas con aquellas simuladas bajo diferentes modelos de evolución. Nuestros resultados apoyan inequívocamente nuestras expectativas basadas en estudios previos acerca de la talla corporal presentando una fuerte señal filogenética, aproximada por un patrón OU con baja restricción, mientras el tamaño de área de distribución es más variable y mejor ajustado por un modelo nulo sugiriendo la ausencia de señal filogenética.A number of methods have been proposed to estimate the level of phylogenetic signal (autocorrelation) in macroecological traits. These methods are useful to devise alternative ways to circumvent the problem of lack of independence among species and, recently, they have also proved valuable to infer how fast a trait has evolved in comparison with alternative evolutionary models, such as a Brownian motion or Ornstein-Uhlenbeck process. Recently, we developed a method called phylogenetic signal-representation (PSR) curve, an expansion of the phylogenetic eigenvector regression (PVR) proposed in 1998, which consists in estimating different coefficients of determination by regressing a trait of interest on the eigenvectors extracted from a phylogenetic distance matrix. The first model uses only the first of these eigenvectors as an explanatory variable; the second model uses both the first and the second and so on. After, the resultant coefficients of determination are plotted against the cumulative eigenvalues, and the shape of this curve is related to evolutionary models driving trait variation (i.e., a linear pattern is expected under Brownian evolution). Here, we used the PSR curve to study patterns of interspecific variation in Carnivora body size and geographical range size, and compared them with simulated curves under distinct evolutionary processes. Our results unequivocally support our expectations based on previous studies that body size has a strong phylogenetic signal, approximated by an OU pattern with low restraining force, whereas geographic range size is more labile and better fits the null expectations (i.e., absence of phylogenetic signal)

Community phylogenetics at the biogeographical scale: cold tolerance, niche conservatism and the structure of North American forests

Aim The fossil record has led to a historical explanation for forest diversity gradients within the cool parts of the Northern Hemisphere, founded on a limited ability of woody angiosperm clades to adapt to mid-Tertiary cooling. We tested four predictions of how this should be manifested in the phylogenetic structure of 91,340 communities: (1) forests to the north should comprise species from younger clades (families) than forests to the south; (2) average cold tolerance at a local site should be associated with the mean family age (MFA) of species; (3) minimum temperature should account for MFA better than alternative environmental variables; and (4) traits associated with survival in cold climates should evolve under a niche conservatism constraint. Location The contiguous United States. Methods We extracted angiosperms from the US Forest Service's Forest Inventory and Analysis database. MFA was calculated by assigning age of the family to which each species belongs and averaging across the species in each community. We developed a phylogeny to identify phylogenetic signal in five traits: realized cold tolerance, seed size, seed dispersal mode, leaf phenology and height. Phylogenetic signal representation curves and phylogenetic generalized least squares were used to compare patterns of trait evolution against Brownian motion. Eleven predictors structured at broad or local scales were generated to explore relationships between environment and MFA using random forest and general linear models. Results Consistent with predictions, (1) southern communities comprise angiosperm species from older families than northern communities, (2) cold tolerance is the trait most strongly associated with local MFA, (3) minimum temperature in the coldest month is the environmental variable that best describes MFA, broad-scale variables being much stronger correlates than local-scale variables, and (4) the phylogenetic structures of cold tolerance and at least one other trait associated with survivorship in cold climates indicate niche conservatism. Main conclusions Tropical niche conservatism in the face of long-term climate change, probably initiated in the Late Cretaceous associated with the rise of the Rocky Mountains, is a strong driver of the phylogenetic structure of the angiosperm component of forest communities across the USA. However, local deterministic and/or stochastic processes account for perhaps a quarter of the variation in the MFA of local communities

A dark scenario for Cerrado plant species: effects of future climate, land-use, and protected areas ineffectiveness

Aim: The anthropogenic climate change and land-use change are considered two of the main factors that are altering biodiversity at the global scale. An evaluation that combined both factors can be relevant to detect which species could be the most vulnerable and reveal the regions of highest stability or susceptibility to biodiversity.We aimed to (i) assess the effect of the climate change and land-use on the distribution of the Cerrado plant species for different countries where it occurs, (ii) evaluate the efficiency of the current protected areas (PAs) network to safeguards species under different greenhouse gas (GHG) emissions and land-use and (iii) estimate the vulnerability of species caused by protection efficiency and habitat loss.Location: Bolivia, Brazil, and ParaguayMethods: We modeled the distribution of 1,553 plant species of Cerrado and evaluated species range loss caused by present and future land-use and two GHG for 2050 and 2080. We assessed the species vulnerability combining the representativeness of a species´ distribution within conservation units and the loss of species range outside PAs.Results: We found that climate change and land-use will cause great damage to Cerrado flora by 2050 and 2080, even under optimistic conditions. Unfortunately, the greatest intensity and extent of land-use will have to overcome on the regions where the greatest richness will be harbored. The conservation of the species will be seriously affected since the PAs network is not as efficient in safeguarding them under current or future conditions. Main conclusions: The low level of protection together with the losses caused by the advance of the agricultural frontier will lead to most species being highly vulnerable. Due to the climate and land-use, effects showed different interactions in each country, conservation strategies should be implemented at transboundary and national levels.Fil: Velazco, Santiago José Elías. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú | Universidad Nacional de Misiones. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú; Argentina. Universidade Federal do Paraná; BrasilFil: Villalobos, Fabricio. Instituto de Ecología; México. Universidade Federal de Goiás; BrasilFil: Galvao, Franklin. Universidade Federal do Paraná; BrasilFil: de Marco Junior, Paulo. Universidade Federal de Goiás; Brasi

Community phylogenetics at the biogeographical scale: cold tolerance, niche conservatism and the structure of North American forests

Aim The fossil record has led to a historical explanation for forest diversity gradients within the cool parts of the Northern Hemisphere, founded on a limited ability of woody angiosperm clades to adapt to mid-Tertiary cooling. We tested four predictions of how this should be manifested in the phylogenetic structure of 91,340 communities: (1) forests to the north should comprise species from younger clades (families) than forests to the south; (2) average cold tolerance at a local site should be associated with the mean family age (MFA) of species; (3) minimum temperature should account for MFA better than alternative environmental variables; and (4) traits associated with survival in cold climates should evolve under a niche conservatism constraint. Location The contiguous United States. Methods We extracted angiosperms from the US Forest Service's Forest Inventory and Analysis database. MFA was calculated by assigning age of the family to which each species belongs and averaging across the species in each community. We developed a phylogeny to identify phylogenetic signal in five traits: realized cold tolerance, seed size, seed dispersal mode, leaf phenology and height. Phylogenetic signal representation curves and phylogenetic generalized least squares were used to compare patterns of trait evolution against Brownian motion. Eleven predictors structured at broad or local scales were generated to explore relationships between environment and MFA using random forest and general linear models. Results Consistent with predictions, (1) southern communities comprise angiosperm species from older families than northern communities, (2) cold tolerance is the trait most strongly associated with local MFA, (3) minimum temperature in the coldest month is the environmental variable that best describes MFA, broad-scale variables being much stronger correlates than local-scale variables, and (4) the phylogenetic structures of cold tolerance and at least one other trait associated with survivorship in cold climates indicate niche conservatism. Main conclusions Tropical niche conservatism in the face of long-term climate change, probably initiated in the Late Cretaceous associated with the rise of the Rocky Mountains, is a strong driver of the phylogenetic structure of the angiosperm component of forest communities across the USA. However, local deterministic and/or stochastic processes account for perhaps a quarter of the variation in the MFA of local communities

Modeling the ecology and evolution of biodiversity: Biogeographical cradles, museums, and graves

Individual processes shaping geographical patterns of biodiversity are increasingly understood, but their complex interactions on broad spatial and temporal scales remain beyond the reach of analytical models and traditional experiments. To meet this challenge, we built a spatially explicit, mechanistic simulation model implementing adaptation, range shifts, fragmentation, speciation, dispersal, competition, and extinction, driven by modeled climates of the past 800,000 years in South America. Experimental topographic smoothing confirmed the impact of climate heterogeneity on diversification. The simulations identified regions and episodes of speciation (cradles), persistence (museums), and extinction (graves). Although the simulations had no target pattern and were not parameterized with empirical data, emerging richness maps closely resembled contemporary maps for major taxa, confirming powerful roles for evolution and diversification driven by topography and climate

Ecological and evolutionary factors in the morphological diversification of South American spiny rats

Understanding the processes underlying morphological diversification is a central goal in ecology and evolutionary biology and requires the integration of information about phylogenetic divergence and ecological niche diversity. In the present study, we use geometric morphometrics and comparative methods to investigate morphological diversification in Neotropical spiny rats of the family Echimyidae. Morphological diversification is studied as shape variation in the skull, comprising a structure composed of four distinct units: vault, base, orognathofacial complex, and mandible. We demonstrate association among patterns of variation in shape in different cranial units, levels of phylogenetic divergence, and ecological niche diversification. At the lower level of phylogenetic divergence, there is significant and positive concordance between patterns of phylogenetic divergence and cranial shape variation in all cranial units. This concordance may be attributable to the phylogenetic and shape distances being calculated between species that occupy the same niche. At higher phylogenetic levels of divergence and with ecological niche diversity, there is significant concordance between shape variation in all four cranial units and the ecological niches. In particular, the orognathofacial complex revealed the most significant association between shape variation and ecological niche diversity. This association may be explained by the great functional importance of the orognathofacial complex.Facultad de Ciencias Naturales y Muse