**6. Conclusions**

#### **6.1 Future directions**

In conclusion, to test the robustness of the theoretical foundations of evolutionary and ecological genetics, three main categories of questions should be answered, using a multidisciplinary approach consisting of: (A) experimental population genetics, (B) collection and analysis of empirical data, and (C) computational population genetics combined with ecological information as for example life history characteristics of the study organism.

#### **6.1.1 Experimental population genetics**

Numerous experiments should be conducted including different model organisms. Many of these experiments should be based on innovative methods accounting for the experimental

always the case as for example in constant environments genetic variability in a quantitative character creates a segregational load each generation due to stabilizing selection against individuals that deviate from the optimum phenotype (Lande & Shannon, 1996). Consider a presumably ordinary situation where natural selection acting on quantitative characters favours intermediate phenotypes. In an intermediate-optimum model, the genetic variability may be either beneficial or detrimental, depending on the pattern of environmental change (the frequency, the amplitude and the degree of autocorrelation of

The genetic consequences of CIEC can be subdivided in two main categories, namely

In small populations, random genetic processes (genetic drift) lead to loss of genetic variability, which may depress the evolutionary potential and thus the ability to respond to changing environments (Pertoldi et al., 2006a). It is also anticipated that populations only persist if the rate of adaptive evolution at least matches the rate of environmental change since the evolutionary response of quantitative traits to selection necessitates the presence of genetic variability (Burger and Lynch, 1995). In fact, this is the case even in the presence significant capacity to respond plastically, including adaptations in behaviour, physiology, morphology, growth, life history and demography. The rate of loss of genetic variability in populations is associated to a reduction of NE. Reduction of NE due to amplified population fluctuations, reduce the evolutionary potential, by reducing the additive genetic variance

the environmental oscillations) (Lande & Shannon, 1996, Björklund et al., 2011).

a) and the heritability (h2) of the traits, which in turn is inversely related to σ<sup>2</sup>

therefore the importance of plasticity is quite evident (Pertoldi et al., 2007b).

In large populations, the regime of alternating selective pressures has the potential to increase the average population fitness, selecting for genes implicated in the expression of plasticity. Various modelling approaches have shown that to optimize fitness, phenotypic plasticity evolves by trading the adaptation to acquire resources against the costs of maintaining the potential for plasticity (Ernande & Dieckman, 2004). Plastic responses include changes in behavior, physiology, morphology, growth, life history and demography, and can be expressed either within the lifespan of an individual or across generations (Pertoldi et al., 2005; Røgilds et al., 2005). Two ways of adapting to environmental changes are therefore possible, by evolutionary or by plastic responses, including maternal transmission (trans-generational plasticity). Hence, the survival of populations relies on genetic variation and/or phenotypic plasticity. Populations with small NE and/or little genetic variability have mainly the option of adapting in a plastic way,

In conclusion, to test the robustness of the theoretical foundations of evolutionary and ecological genetics, three main categories of questions should be answered, using a multidisciplinary approach consisting of: (A) experimental population genetics, (B) collection and analysis of empirical data, and (C) computational population genetics combined with

Numerous experiments should be conducted including different model organisms. Many of these experiments should be based on innovative methods accounting for the experimental

ecological information as for example life history characteristics of the study organism.

e.

consequences in small populations and consequences in large populations:

(σ<sup>2</sup>

**6. Conclusions** 

**6.1 Future directions** 

**6.1.1 Experimental population genetics** 

errors due to unpredictable environmental components (for applications see Kristensen et al., 2004). Clonally reproducing strains should be used to study the extent of adaptive phenotypic plasticity, and maternal effects, including the effect of parental ageing. The use of clonal strains will allow us to exclude the genetic components and their interactions with the environment. Therefore, unbiased estimates of genetic and environmental canalization, plasticity, developmental homeostasis and σ<sup>2</sup> e, will be obtained. A more correct interpretation of the interplay between these parameters will provide important contributions to: 1) The evolutionary importance of phenotypic plasticity, maternal effects, environmental and genetic stressors. 2) The consequences of outbreeding on population fitness and phenotypic plasticity, and 3) The selective effects of fluctuating selective regimes on plasticity genes.

In order to investigate in which way environments fluctuating with different intervals can affect the mean population average fitness, and to quantify the costs and benefits of genetic variability in fluctuating environments, sexually reproducing strains ought to be utilised, creating fluctuating temperature environments and making truncated selection experiments in which the extreme phenotypes at the two tails of the phenotypic distribution are selected away. Important information could in this way be obtained about the extent of the environmental information that will be transmitted to the offspring, and to what extent it can enlarge the plastic response of a trait when selecting for plasticity genes.

#### **6.1.2 Collection and analysis of empirical data**

Molecular and quantitative genetics studies should be conducted on several species with different ecological characteristics and with different demographic history, such as recent and ancient population decline or expansion. Changes in population size and range are frequent consequences of CIEC, and examples include habitat fragmentation and rapid colonization or recolonization processes. Extensive collections of several species provide the opportunity to analyse large numbers of samples on a temporal scale and directly document changes in genetic diversity. The results of these analyses will improve our understanding of the historical dimension of population change, and provide important data for the interpretation of genetic diversity studies in an ecological and evolutionary context. The possibility of amplifying ancient DNA from old museum specimen (Pertoldi et al., 2005b), should also be used. Furthermore, a phylogeographic approach should be carried out. The innovative aspect of this approach consists of the fact that different molecular and quantitative genetics techniques should be employed simultaneously.

To document the range of genetically based morphological variation within and among populations, a comparison of the degree of quantitative genetics distance (QST) with neutral genetic distance (FST) should be made (Mckay & Latta, 2002). Comparisons of morphometrical (for example, size and shape) and life-history variability (for example, longevity and fecundity) of populations and their crosses with molecular variability (using microsatellites) could present important information about the influences of environmental and genetic components in a non-genetic-equilibrium situation. Furthermore, it will provide important information about the extent to which crosses between different strains affect the various components of σ<sup>2</sup> p (plasticity, developmental homeostasis, canalization and σ2e).

The combination of ecological models of the distribution of the species investigated with both mitochondrial DNA (mtDNA) data and synthetic genetic maps constructed from

Possible Evolutionary Response to Global Change – Evolutionary Rescue? 99

molecular data in combination with quantitative traits and environmental data, and **4)** Unravelling the distribution of variation at functional vs. non-coding sequences in natural

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multivariate analysis of microsatellites and morphometric data will allow us to discuss hypothesized historical biogeographic scenarios. By directly dating and quantifying changes in genetic diversity, these investigations will allow examination of postulated causes of population decline, including habitat loss and temperature increase.

The genetic data obtained from the investigations mentioned above provides the information on postglacial history as well as on current demographic threats (fragmentation, relict populations, marginal populations; levels of inbreeding). The results of this approach will provide important information about: 1) The genetic consequences of population fragmentation and rapid recolonization caused by climate change. 2) The extent of the genetic diversity of modern populations compared to that in the late Pleistocenic environment, and 3) Pattern of species recolonization in Europe and their response to environmental change after the last glaciation.

#### **6.1.3 Computational population genetics**

Stochastic simulation tools, based on a quantitative infinitesimal model, where the size of NE can be varied, should be developed. In the simulation models, each phenotype should be considered to be the sum of independent genetic components (σ 2a, dominance and epistasis) and σ<sup>2</sup> e, and the σ<sup>2</sup> p in a population should be described as the sum of independent variances for each of these effects. Several questions could be answered:


Given that the information obtained from the computational approach can be combined with empirical data, obtained from approaches 6.1.1 and 6.1.2 the model will be a powerful tool for understanding complex dynamics and to make predictions concerning the possible effects of CIEC and their interactions with other factors.

#### **6.2 Expected yield of the multidisciplinary approach**

The establishment of such an approach which integrate experimental, theoretical and applied ecological and evolutionary genetics, will create synergistic effects and contribute to the understanding of the consequences of CIEC and the questions addressed will provide important contributions to general ecology and conservation genetics as there is a requirement for detailed studies on how variation at the level of genes translates through developmental and physiological processes, into phenotypic variation for ecologically important traits. Further scientific progress will be achieved by merging and complementing recent efforts in evolutionary and ecological genetics by: 1) Collecting informative genetic and environmental data sets in natural populations and from preserved specimens, 2) Merging taxonomic, ecological and genetic databases, 3) Using molecular data in combination with quantitative traits and environmental data, and **4)** Unravelling the distribution of variation at functional vs. non-coding sequences in natural populations.

#### **7. References**

98 International Perspectives on Global Environmental Change

multivariate analysis of microsatellites and morphometric data will allow us to discuss hypothesized historical biogeographic scenarios. By directly dating and quantifying changes in genetic diversity, these investigations will allow examination of postulated causes of

The genetic data obtained from the investigations mentioned above provides the information on postglacial history as well as on current demographic threats (fragmentation, relict populations, marginal populations; levels of inbreeding). The results of this approach will provide important information about: 1) The genetic consequences of population fragmentation and rapid recolonization caused by climate change. 2) The extent of the genetic diversity of modern populations compared to that in the late Pleistocenic environment, and 3) Pattern of species recolonization in Europe and their response to

Stochastic simulation tools, based on a quantitative infinitesimal model, where the size of NE can be varied, should be developed. In the simulation models, each phenotype should be considered to be the sum of independent genetic components (σ 2a, dominance and epistasis)

1. Understanding how different environmental scenarios can affect both genetic and

2. Understanding how much difference in life history between ecologically similar species can cause substantial differences in NE and σ2a, and to what degree fluctuations in vital

3. Quantifying the interactions of each particular life history parameter with other factors

4. Quantifying the effects and the interactions that factors such as NE, inbreeding, gametic phase disequilibrium, plasticity, and developmental homeostasis have on the speed at

Given that the information obtained from the computational approach can be combined with empirical data, obtained from approaches 6.1.1 and 6.1.2 the model will be a powerful tool for understanding complex dynamics and to make predictions concerning the possible

The establishment of such an approach which integrate experimental, theoretical and applied ecological and evolutionary genetics, will create synergistic effects and contribute to the understanding of the consequences of CIEC and the questions addressed will provide important contributions to general ecology and conservation genetics as there is a requirement for detailed studies on how variation at the level of genes translates through developmental and physiological processes, into phenotypic variation for ecologically important traits. Further scientific progress will be achieved by merging and complementing recent efforts in evolutionary and ecological genetics by: 1) Collecting informative genetic and environmental data sets in natural populations and from preserved specimens, 2) Merging taxonomic, ecological and genetic databases, 3) Using

p in a population should be described as the sum of independent

population decline, including habitat loss and temperature increase.

variances for each of these effects. Several questions could be answered:

rate parameters induced by environmental change can alter NE,

which a population can react to a selective pressure.

effects of CIEC and their interactions with other factors.

**6.2 Expected yield of the multidisciplinary approach** 

environmental change after the last glaciation.

**6.1.3 Computational population genetics** 

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**Historical Environmental Change** 


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**6** 

*1,3Spain 2,4Argentina* 

**How Did Past Environmental Change** 

*1Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, CSIC, Madrid,* 

An important aspect of evolutionary biology is to know the faunal diversity changes through time. There is a general agreement that the knowledge of evolutionary patterns through time from an ecological perspective provides information that is not available from ecological studies on extant faunas (Vrba, 1985, 1995a; Behrensmeyer et al*.*, 1992; Jablonski, 2005). Iberian Cenozoic basins provide an exceptional record of fossil mammals and continental environments, giving a great opportunity to evaluate the ecological and evolutionary responses of mammalian communities to climatic changes through the last millions years (Azanza et al., 1999, 2000; Hernández Fernández et al., 2007; Van Dam et al., 2006). This knowledge is essential for linking the dynamics of biotic change from ecological to evolutionary time scales and for understanding the processes that transform ecosystems

Previous works have proposed different explanations for biodiversity changes through time, emphasizing the influence of both physic and biotic factors (e.g., Van Valen, 1973; Janis, 1989; Stucky, 1990; Vrba, 1995a, 1995b, 2000; Prothero, 1999, 2004; Alroy, 2000; Alroy et al*.*, 2000; Barnosky, 2001, 2005; Vrba & DeGusta, 2004). The response of mammals throughout the late Cenozoic has been often reflected by migrations or variation of their area of distribution, related to the vegetation cover and latitudinal displacement of biomes. Patterns of change in the home range size (HR, the size of the minimum area that can sustain the individual's energetic requirements) through time can provide important insights into the ecological and evolutionary responses of mammalian communities to new environmental conditions. In a lesser degree, mammals can also respond evolving into new species. These events could modify the structure of mammalian communities, triggering new internal

**1. Introduction** 

over geologic times (Badgley et al., 2008).

*4Departamento de Ciencias de la Tierra, Universidad de Zaragoza, Zaragoza,* 

**Affect Carnivore Diversity and** 

**Home-Range-Size in Spain?** 

Esperanza Cerdeño3 and Beatriz Azanza4

*2INCUAPA, Departamento de Arqueología, F.C.S., Universidad Nacional del Centro, Olavarría, 3Departamento de Paleontología, IANIGLA,* 

*Centro Científico Tecnológico CONICET, Mendoza,* 

María T. Alberdi1, José L. Prado2,
