Phylogenetic comparative analysis of life-history variation among populations of the lizard Sceloporus undulatus: an example and prognosis

Over the past 15 years, phylogenetic comparative methods (PCMs) have become standard in the study of life-history evolution. To date, most studies have focused on variation among species or higher taxonomic levels, generally revealing the presence of significant phylogenetic effects as well as residual variation potentially attributable to adaptive evolution. Recently, population-level phylogenetic hypotheses have become available for many species, making it possible to apply PCMs directly to the level at which experiments are typically used to test adaptive hypotheses. In this study, we present the results of PCMs applied to life-history variation among populations of the widespread and well-studied lizard Sceloporus undulatus. Using S. undulatus (which may represent four closely related species) as an example, we explore the benefits of using PCMs at the population level, as well as consider the importance of several thorny methodological problems including but not limited to nonindependence of populations, lack of sufficient variation in traits, and the typically small sample sizes dictated by the difficulty of collecting detailed demographic data. We show that phylogenetic effects on life-history variation among populations of S. undulatus appear to be unimportant, and that several classic trade-offs expected by theory and revealed by many interspecific comparisons are absent. Our results suggest that PCMs applied to variation in life-history traits below the species level may be of limited value, but more studies like ours are needed to draw a general conclusion. Finally, we discuss several outstanding problems that face studies seeking to apply PCMs below the species level.     

Allozyme variation in 16 natural populations of Faidherbia albida (Del.) A. Chev

The objectives of this study were (1) to investigate the genetic variability; and (2) to determine the phylogenetic relationship in the leguminous tree Faidherbia albida through out its range of distribution in East, West and Southern Africa. A total of 16 populations were subjected to enzyme electrophoresis and 6 enzyme systems (Adh, Mdh, G6pdh, Idh, 6Pgdh, and alpha-Est) encoded by 12 loci were scored. A total of fifty one different alleles were detected, with an average of 2.5 alleles per locus. Forty three percent of the loci were polymorphic at a 95% criterion. The average expected heterozygosity (gene diversity index H(e)) was 0.141. All provenances showed significant deviation from Hardy-Weinberg expectation. The UPGMA cluster analysis, based on Modified Rogers distance revealed close similarities between Eastern and Southern African provenances, except Debre zeit from Ethiopia, which was closest to West African populations than to the East African populations. Also, Bignona from West Africa was peculiarly closer to East African provenances. Differentiation of West African populations from Eastern and Southern African provenances was quite evident, emphasising the clinical pattern of variation in these two major geographical races of F. albida in Africa. Wright's F-statistics showed an overall significant deficit of heterozygotes, a common feature in mixed mating, entomophilous, widespread species such as F. albida. The dendrogram analysis showed wide separation among the three Ethiopian provenances indicating a high level of genetic differentiation and diversity among them.     

A novel primer set for multilocus phylogenetic inference in East African cichlid fishes

The cichlid fishes in the East African Great Lakes are a prime model system for the study of adaptive radiation. Therefore, the availability of an elaborate phylogenetic framework is an important prerequisite. Previous phylogenetic hypotheses on East African cichlids are mainly based on mitochondrial and/or fragment‐based markers, and, to date, no taxon‐rich phylogeny exists that is based on multilocus DNA sequence data. Here, we present the design of an extensive new primer set (24 nuclear makers) for East African cichlids that will be used for multilocus phylogenetic analyses in the future. The primers are designed to work for both Sanger sequencing and next‐generation sequencing with the 454 technology. As a proof of principle, we validate these primers in a phylogenetically representative set of 16 cichlid species from Lake Tanganyika and main river systems in the area and provide a basic evaluation of the markers with respect to marker length and diversity indices.     

Population structure in African Drosophila melanogaster revealed by microsatellite analysis

Tropical sub-Saharan regions are considered to be the geographical origin of Drosophila melanogaster. Starting from there, the species colonized the rest of the world after the last glaciation about 10 000 years ago. Consistent with this demographic scenario, African populations have been shown to harbour higher levels of microsatellite and sequence variation than cosmopolitan populations. Nevertheless, limited information is available on the genetic structure of African populations. We used X chromosomal microsatellite variation to study the population structure of D. melanogaster populations using 13 sampling sites in North, West and East Africa. These populations were compared to six European and one North American population. Significant population structure was found among African D. melanogaster populations. Using a Bayesian method for inferring population structure we detected two distinct groups of populations among African D. melanogaster. Interestingly, the comparison to cosmopolitan D. melanogaster populations indicated that one of the divergent African groups is closely related to cosmopolitan flies. Low, but significant levels of differentiation were observed for sub-Saharan D. melanogaster populations from West and East Africa.     

Molecular evidence for species level divergence in African Nile Crocodiles Crocodylus niloticus (Laurenti, 1786)

Relationships of the newly discovered dwarf crocodiles from Mauritania were inferred from mitochondrial 12S sequences. Specimens from 13 different Crocodylus niloticus populations (from East Africa, West Africa and Madagascar) were compared. Additional representatives of the genus Crocodylus (one from Africa and one from Australia), the African genus Osteolaemus and the South American alligatorid Paleosuchus palpebrosus (as outgroup) were included in the analysis. Maximum-likelihood and Bayesian analyses yielded relationships that were strikingly different from currently prevailing phylogenetic hypotheses. Both analyses consistently revealed two groups, one consisting of the monophyletic West- and Central African populations and the other of a paraphyletic group containing the East African and Madagascan populations. High genetic divergence between those groups indicates separation on the species level. Furthermore ‘C’ cataphractus is clearly shown not to be a member of the genus Crocodylus. The resulting nomenclatural changes are discussed. To cite this article: A. Schmitz et al., C. R. Palevol 2 (2003).     

Genetic diversity of donkey populations from the putative centers of domestication

Donkey domestication drastically changed ancient transport systems in Africa and Asia, enabling overland circulation of people and goods and influencing the organization of early cities and pastoral societies. Genetic studies based on mtDNA have pointed to the African wild ass as the most probable ancestor of the domestic donkey, but questions regarding its center of origin remain unanswered. Endeavoring to pinpoint the geographical origin of domestic donkey, we assessed levels and patterns of genetic diversity at 15 microsatellite loci from eight populations, representing its three hypothesized centers of origin: northeast Africa, the Near East and the Arabian Peninsula. Additionally, we compared the donkey genotypes with those from their wild relative, the African wild ass (Equus africanus somaliensis) to visualize patterns of differentiation among wild and domestic individuals. Obtained results revealed limited variation in levels of unbiased expected heterozygosity across populations in studied geographic regions (ranging from 0.637 in northeast Africa to 0.679 in the Near East). Both allelic richness (Ar) and private allelic richness presented considerably higher values in northeast Africa and in the Arabian Peninsula. By looking at variation at the country level, for each region, we were able to identify Sudan and Yemen as the countries possessing higher allelic richness and, cumulatively, Yemen also presented higher values for private allelic richness. Our results support previously proposed northeast Africa as a putative center of origin, but the high levels of unique diversity in Yemen opens the possibility of considering this region as yet another center of origin for this species.     

GM haplotype diversity of 82 populations over the world suggests a centrifugal model of human migrations

This study investigates the GM genetic relationships of 82 human populations, among which 10 represent original data, within and among the main broad geographic areas of the world. Different approaches are used: multidimensional scaling analysis and test for isolation by distance, to assess the correlation between genetic variation and spatial distributions; analysis of variance, to investigate the genetic structure at different hierarchical levels of population subdivision; genetic similarity map (geographic map distorted by available genetic information), to identify regions of high and low genetic variation; and minimal spanning network, to point out possible migration routes across continental areas. The results show that the GM polymorphism is characterized by one of the highest amounts of genetic variation observed so far among populations of different continents (Fct=0.3915, P < 0.0001). GM diversity can be explained by a model of isolation by distance (IBD) at most continental levels, with a particularly significant fit to IBD for the Middle East and Europe. Five peripheral regions of the world (Europe, west and south sub-Saharan Africa, Southeast Asia, and America) exhibit a low level of genetic diversity both within and among populations. By contrast, East and North African, Southwest Asian, and Northeast Asian populations are highly diverse and interconnected genetically by large genetic distances. Therefore, the observed GM variation can be explained by a "centrifugal model" of modern humans peopling history, involving ancient dispersals across a large intercontinental area spanning from East Africa to Northeast Asia, followed by recent migrations in peripheral geographic regions.     

Plumage-based phylogenetic analyses of the Merops bee-eaters

I review previous systematic work on the family Meropidae and present phylogenetic hypotheses derived from my analyses of colour, pattern and shape variation in 30 plumage regions among species and subspecies in this family. Consistent patterns are seen across shallow portions of the trees. Uncertainty remains concerning the placement of several deep branches within this group's phylogeny. In particular, the phylogenetic placement of Meropogon forsteni and Merops breweri , M. ornatus , M. hirundineus and M. boehmi remains uncertain. The biogeographical patterns in the resultant trees are similar with either a Southeast Asian or African origin for the family, with most of the early diversification occurring in Africa, and with multiple independent subsequent invasions of non-African areas.     

Reanalysis of oviposition and development rates in the Phytoseiidae using a phylogenetic autoregressive method

Life history patterns are usually identified by comparisons of extant species. Because of inferences regarding phylogenetic constraints, comparative data are often not statistically independent. In order to remove phylogenetic patterns embedded in life history data completely, we adopted a phylogenetic autoregressive method to reanalyse a data set of the ovipositional and developmental rates of 45 Phytoseiidae species. We first calculated the phylogenetic correlation in relation to different taxonomic levels using Moran's I statistics. Significant and positive phylogenetic correlations were found at the subgenus and subfamily levels. This indicates that some variation in both of these life-history traits could be accounted for by phylogeny. Phylogenetic associations, therefore, were removed by a phylogenetic autoregressive method. Using corrected data from this method, the specific components of the ovipositional rate are positively correlated with the specific components of th e developmental rate. The method that we have used obtains the same conclusion as others but differs from the phylogenetic effect in the way that it influences the relationship between comparative data. Because of no data reduction in the phylogenetic autoregressive method, the specific components are more useful than the mean values derived from the higher taxonomic nodes for testing ecological and evolutionary hypotheses about life history patterns. © Rapid Science Ltd. 1998     

Autoregressive models for estimating phylogenetic and environmental effects: accounting for within-species variations

Comparative methods are used to investigate the attributes of present species or higher taxa. Difficulties arise from the phylogenetic heritage: taxa are not independent and neglecting phylogenetic inertia can lead to inaccurate results. Within-species variations in life-history traits are also not negligible, but most comparative methods are not designed to take them into account. Taxa are generally described by a single value for each trait. We have developed a new model which permits the incorporation of both the phylogenetic relationships among populations and within-species variations. This is an extension of classical autoregressive models. This family of models was used to study the effect of fishing on six demographic traits measured on 77 populations of teleost fishes.     

Evolutionary history of the parrotfishes: biogeography,ecomorphology, and comparative diversity

The family Scaridae comprises about 90 species of herbivorous coral reef, rock reef, and seagrass fishes. Parrotfishes are important agents of marine bioerosion who rework the substrate with their beaklike oral jaws. Many scarid populations are characterized by complex social systems including highly differentiated sexual stages, territoriality, and the defense of harems. Here, we test a hypothesis of relationships among parrotfish genera derived from nearly 2 kb of nuclear and mitochondrial DNA sequence. The DNA tree is different than a phylogeny based on comparative morphology and leads to important reinterpretations of scarid evolution. The molecular data suggest a split among seagrass and coral reef associated genera with nearly 80% of all species in the coral reef clade. Our phylogenetic results imply an East Tethyan origin of the family and the recurrent evolution of excavating and scraping feeding modes. It is likely that ecomorphological differences played a significant role in the initial divergence of major scarid lineages, but that variation in color and breeding behavior has triggered subsequent diversification. We present a two-phase model of parrotfish evolution to explain patterns of comparative diversity. Finally, we discuss the application of this model to other adaptively radiating clades.     

Testing evolutionary hypotheses about human biological adaptation using cross-cultural comparison

Physiological data from a range of human populations living in different environments can provide valuable information for testing evolutionary hypotheses about human adaptation. By taking into account the effects of population history, phylogenetic comparative methods can help us determine whether variation results from selection due to particular environmental variables. These selective forces could even be due to cultural traits-which means that gene-culture co-evolution may be occurring. In this paper, we outline two examples of the use of these approaches to test adaptive hypotheses that explain global variation in two physiological traits: the first is lactose digestion capacity in adults, and the second is population sex-ratio at birth. We show that lower than average sex ratio at birth is associated with high fertility, and argue that global variation in sex ratio at birth has evolved as a response to the high physiological costs of producing boys in high fertility populations.     

Morphological variation of major human populations based on nonmetric dental traits

The patterns of inter- and intra-regional variation among 12 major geographical groups from around the world were investigated based on 15 nonmetric dental traits. The R-matrix method was applied using a pooled within-group variance-covariance matrix estimated with the maximum likelihood method (tetrachoric correlation matrix) and the threshold value for each trait estimated by univariate probit analysis. Using average heritability rates that range from 0.40 to 1.00, the inter-regional variation represented by Fst falls between 7.19% and 16.23% of the total variance. This range of variation is compatible with those obtained by genetic, craniometric, and odontometric data. Subsaharan Africans show the largest intra-regional diversity among the groups compared. The degree of intra-regional variation shows, moreover, rough clinalities from subsaharan Africa to peripheral regions. The relationship between regional variation and geographic distance from subsaharan Africa supports serial bottlenecks and the founder effect of ancient populations originating in Africa. The variation of East/Northeast Asians is relatively large, suggesting a complex population history such as possible earlier divergence and multiple migrations from outside sources. The present findings are in agreement with both the recent African model for the origin of anatomically modern humans and the current scenario for human migration history suggested by genetic analyses.     

Cultural macroevolution and the transmission of traits

Cultural traits are distributed across human societies in a patterned way. Study of the mechanisms whereby cultural traits persist and change over time is key to understanding human cultural diversity. For more than a century, a central question has engaged anthropologists interested in the study of cultural trait variation: What is the source of cultural variation? More precisely, are cultural traits transmitted primarily from ancestral to descendant populations (vertical transmission) or between contemporary, typically neighboring populations (horizontal transmission), or do they emerge as independent innovations? While debates among unilineal evolutionists and diffusionists have long since faded, there is still much uncertainty about how traits are transmitted at this macroevolutionary level, as well as about the implications of these transmission patterns for testing hypotheses regarding the adaptive function of particular cultural traits across human populations.     

Comparison of the Mutation Spectrum of Hypervariable Segment 1 for Phylogeographical Groups of Human Mitochondrial DNA

The nucleotide sequence variation of hypervariable segment 1 (HVS1) was analyzed for mtDNAs of 88 phylogeographical clusters characteristic of African, West Eurasian, or East Eurasian populations. A significant difference in the distribution of mutations was revealed for the mitochondrial gene pools of the regional human populations. The HVS1 positions were identified whose instability is explained by strand displacement during mtDNA replication. Strand displacement was assumed to be a major mechanism of context-dependent mutagenesis associated with the regional differentiation of human populations.     

Modèle de mélange et analyse comparative des traits démographiques des poissons: Mixture models and comparative analysis of life history traits of Teleost fishes

The comparative approach in biology is used to investigate the attributes of actual species. Differences between them are used to formulate adaptive hypotheses of life history, morphology, demography, etc. Difficulties arise from the phylogenetic heritage: species are not independent and neglecting this fact can lead to inaccurate results. Fishing can have important effects but usually they are mixed with other variations such as environmental ones. Single species studies cannot succeed completely in analysing the effect of fishing. To disentangle them from orher fluctuations, we are led to the use of a comparative approach. Thus, we need to account for phylogenetic links between populations. In this paper we propose a new family of models, which allows us to split life history traits into parts.     

Global analysis of regional differences in craniometric diversity and population substructure.

Estimates of genetic diversity in major geographic regions are frequently made by pooling all individuals into regional aggregates. This method can potentially bias results if there are differences in population substructure within regions, since increased variation among local populations could inflate regional diversity. A preferred method of estimating regional diversity is to compute the mean diversity within local populations. Both methods are applied to a global sample of craniometric data consisting of 57 measurements taken on 1734 crania from 18 local populations in six geographic regions: sub-Saharan Africa, Europe, East Asia, Australasia, Polynesia, and the Americas. Each region is represented by three local populations. Both methods for estimating regional diversity show sub-Saharan Africa to have the highest levels of phenotypic variation, consistent with many genetic studies. Polynesia and the Americas both show high levels of regional diversity when regional aggregates are used, but the lowest mean local population diversity. Regional estimates of F(ST) made using quantitative genetic methods show that both Polynesia and the Americas also have the highest levels of differentiation among local populations, which inflates regional diversity. Regional differences in F(ST) are directly related to the geographic dispersion of samples within each region; higher F(ST) values occur when the local populations are geographically dispersed. These results show that geographic sampling can affect results, and suggest caution in making inferences regarding regional diversity when population substructure is ignored.     

Ecological and Phylogenetic Effects on Song Sharing in Songbirds

As a consequence of vocal learning, the songs of male songbirds often vary geographically. We examined three proximate hypotheses that predict variation in the incidence of local song sharing among species: the annual cycle, breeding season duration, and epiphenomenon hypotheses. To test these hypotheses, we compiled data from the literature on local song sharing, annual cycle, breeding location isophane (an index of breeding season duration), repertoire size, and phylogenetic relationships of 65 populations in the family Fringillidae. Mapping characters on the phylogenetic tree revealed that local song sharing, migratory status, and breeding location isophane were randomly distributed on the phylogeny. Breeding isophane and migratory status were themselves correlated and had significant effects on the probability of sharing local songs. As predicted by the annual cycle and breeding season duration hypotheses, males in sedentary populations and populations breeding at low isophanes were more likely to share local songs. In subsets of the data, the annual cycle hypothesis received more support than did the breeding season duration hypothesis. Neither hypothesis explained more than 17% of the variation in the incidence of song sharing, and we suggest that detailed information on song function, male dispersal, and constraints on the timing of song learning will be needed to better predict the occurrence of local song sharing.     

Evolutionary history of the honey bee Apis mellifera inferred from mitochondrial DNA analysis

Variability of mitochondrial DNA (mtDNA) of the honey bee Apis mellifera L. has been investigated by restriction and sequence analyses on a sample of 68 colonies from ten different subspecies. The 19 mtDNA types detected are clustered in three major phylogenetic lineages. These clades correspond well to three groups of populations with distinct geographical distributions: branch A for African subspecies (intermissa, monticola, scutellata, andansonii and capensis), branch C for North Mediterranean subspecies (caucasica, carnica and ligustica) and branch M for the West European populations (mellifera subspecies). These results partially confirm previous hypotheses based on morphometrical and allozymic studies, the main difference concerning North African populations, now assigned to branch A instead of branch M. The pattern of spatial structuring suggests the Middle East as the centre of dispersion of the species, in accordance with the geographic areas of the other species of the same genus. Based on a conservative 2% divergence rate per Myr, the separation of the three branches has been dated at about 1 Myr BP.     

Indirect evidence for the genetic determination of short stature in African Pygmies

Central African Pygmy populations are known to be the shortest human populations worldwide. Many evolutionary hypotheses have been proposed to explain this short stature: adaptation to food limitations, climate, forest density, or high mortality rates. However, such hypotheses are difficult to test given the lack of long-term surveys and demographic data. Whether the short stature observed nowadays in African Pygmy populations as compared to their Non-Pygmy neighbors is determined by genetic factors remains widely unknown. Here, we study a uniquely large new anthropometrical dataset comprising more than 1,000 individuals from 10 Central African Pygmy and neighboring Non-Pygmy populations, categorized as such based on cultural criteria rather than height. We show that climate, or forest density may not play a major role in the difference in adult stature between existing Pygmies and Non-Pygmies, without ruling out the hypothesis that such factors played an important evolutionary role in the past. Furthermore, we analyzed the relationship between stature and neutral genetic variation in a subset of 213 individuals and found that the Pygmy individuals' stature was significantly positively correlated with levels of genetic similarity with the Non-Pygmy gene-pool for both men and women. Overall, we show that a Pygmy individual exhibiting a high level of genetic admixture with the neighboring Non-Pygmies is likely to be taller. These results show for the first time that the major morphological difference in stature found between Central African Pygmy and Non-Pygmy populations is likely determined by genetic factors.