Population Genomic Analysis Reveals a Rich Speciation and Demographic History of Orang-utans (Pongo pygmaeus and Pongo abelii)

To gain insights into evolutionary forces that have shaped the history of Bornean and Sumatran populations of orang-utans, we compare patterns of variation across more than 11 million single nucleotide polymorphisms found by previous mitochondrial and autosomal genome sequencing of 10 wild-caught orang-utans. Our analysis of the mitochondrial data yields a far more ancient split time between the two populations (∼3.4 million years ago) than estimates based on autosomal data (0.4 million years ago), suggesting a complex speciation process with moderate levels of primarily male migration. We find that the distribution of selection coefficients consistent with the observed frequency spectrum of autosomal non-synonymous polymorphisms in orang-utans is similar to the distribution in humans. Our analysis indicates that 35% of genes have evolved under detectable negative selection. Overall, our findings suggest that purifying natural selection, genetic drift, and a complex demographic history are the dominant drivers of genome evolution for the two orang-utan populations.     

The active evolutionary lives of echinoderm larvae

Echinoderms represent a researchable subset of a dynamic larval evolutionary cosmos. Evolution of echinoderm larvae has taken place over widely varying time scales from the origins of larvae of living classes in the early Palaeozoic, approximately 500 million years ago, to recent, rapid and large-scale changes that have occurred within living genera within a span of less than a million years to a few million years. It is these recent evolutionary events that offer a window into processes of larval evolution operating at a micro-evolutionary level of evolution of discrete developmental mechanisms. We review the evolution of the diverse larval forms of living echinoderms to outline the origins of echinoderm larval forms, their diversity among living echinoderms, molecular clocks and rates of larval evolution, and finally current studies on the roles of developmental regulatory mechanisms in the rapid and radical evolutionary changes observed between closely related congeneric species.     

Archaeology and Human Evolution

Archaeology is the study of human behavior through material culture, the things we rely on for survival. Behavioral change was likely a driving factor in the evolution of our species, and archaeology therefore plays a central role in understanding human origins from the beginning of the known archaeological record some 2.5 million years ago. From its origins to subsequent diversification, the material record of human behavioral innovation provides an essential learning tool for understanding human behavioral diversity and also serves as a gateway to critical thinking in education.     

The Evolution of Human Origins

Because humans are the product of our evolutionary past, learning how we evolved is fundamental to all anthropological investigations. We now realize that reconstructing why unique human attributes evolved requires an understanding of our starting point, but this is a relatively recent perspective. One hundred years ago, the question of human origins was identical to that of hominin origins. Accepting Australopithecus into human ancestry, coupled with the modern synthesis of evolution, led anthropologists to consider humans as products of natural selection. They realized that increased intelligence did not initially distinguish our lineage, and that early hominins were apelike in many ways. Australopithecus brought bipedalityr and brain expansion came with Homo . Because the human mind and behavior are products of evolution, we must reconstruct the selective pressures that shaped our lineage in order to understand ourselves today. Paleoanthropology, as with all anthropology, is becoming ever more question oriented, drawing on many areas of inquiry. [Keywords: human origins, human evolution, history, data, theory]     

De l’origine des anthropoïdes à l’émergence de la famille humaine

The human lineage has a very ancient origin, as most of the mammals. Its oldest representatives, anthropoid primates, have been described from Asia some 45 million years ago. During this long evolutionary story, two critical stages have appeared as especially important, their beginning in Asia and the emergence of hominids in Africa, some seven million years ago. These two stages are discussed hereby with new data relative to their Asian origins and their dispersal into Africa between 45 and 40 million years ago. Following this dispersal event, these primates evolved in Africa and gave rise to the early hominids. These appeared around seven million years ago and have three distinct representatives. Among them, Toumaï appears as the oldest and the closest to our ancestry, a point that is evidenced here.     

The evolution of the thrombospondin gene family

Summary Thrombospondin-1 is an adhesive glycoprotein that is involved in cellular attachment, spreading, migration, and proliferation. To date, four genes have been identified that encode for the members of the thrombospondin gene family. These four genes are homologous to each other in the EGF-like (type 2) repeats, the calcium-binding (type 3) motifs, and the COOH-terminal. The latter has been reported to be a cell-binding domain in thrombospondin-1. Phylogenetic trees have been constructed from the multisequence alignment of thrombospondin sequences from human, mouse, chicken, and frog. Two different algorithms generate comparable results in terms of the topology and the branch lengths. The analysis indicates that an early form of the thrombospondin gene duplicated about 925 million years ago. The gene duplication that produced the thrombospondin-1 and -2 branches of the family is predicted to have occurred 583 million years ago, whereas the gene duplication that produced the thrombospondin-3 and -4 branches of the family is predicted to have occurred 644 million years ago. These results indicate that the members of the thrombospondin gene family have existed throughout the evolution of the animal kingdom and thus probably participate in functions that are common to most of its members.     

Tests of pleistocene speciation in montane grasshoppers (genus Melanoplus) from the sky islands of western North America

There has a been a resurgence of debate on whether the Pleistocene glaciations inhibited speciation. This study tests a model of Pleistocene speciation, estimating the phylogenetic relationships and divergence times of 10 species of montane grasshoppers, genus Melanoplus, using 1300 bp of the mitochondrial gene cytochrome oxidase I (COI). Based on average pairwise distances (corrected for multiple substitutions using Kimura's two-parameter model), all species appear to have originated within the Pleistocene. Sequence divergences between species are less than 4%, corresponding to divergence times less than 1.7 million years ago. Branching patterns among the species suggest that speciation was associated with more than one glacial-interglacial cycle. A likelihood-ratio test rejected a model of simultaneous species origins, the predicted branching pattern if species arose from the fragmentation of a widespread ancestor. These grasshoppers live in an area that was previously glaciated and, as inhabitants of the northern Rocky Mountain sky islands, underwent latitudinal and probably altitudinal shifts in distribution in response to climatic fluctuations. Given the repeated distributional shifts and range overlap of the taxa, there most likely has been ample opportunity for population mixing. However, despite periodic glacial cycles, with more than 10 major glaciations over the past million years and climatic fluctuations over as short a time scale as 10(3) to 10(4) years, the dynamic history of the Pleistocene did not preclude speciation. Although relationships among some taxa remain unresolved, these grasshopper species, even with their recent origins, exhibit genetic coherence and monophyletic or paraphyletic gene trees. The frequency of glacial cycles suggests that the speciation process must have been extremely rapid. These species of grasshoppers are morphologically very similar, differing primarily in the shape of the male genitalia. These characters are posited to be under sexual selection, may play an important role in reproductive isolation, and are known to diverge rapidly. This suggests the rapidity of evolution of reproductive isolation may determine whether species divergences occurred during the Pleistocene glaciations.     

The importance of prebiotic chemistry in the RNA world

In vitro selection experiments have clearly demonstrated that RNA can perform many of the functions necessary to support an RNA world. Moreover, it appears that novel functions could have readily evolved from existing functional RNA molecules. Therefore, diverse molecular ecosystems could potentially have arisen from an initial, small population of functional replicators. These findings suggest that the sequences of living systems may have been determined in part by chance occurrences at origins. Any extrapolations linking sequences (as opposed to functions) obtained in the laboratory to what may have occurred ca. 4 billion years ago are tenuous at best. Thus, perhaps the best way to understand origins is not by examining relatively unconstrained sequence information, but by examining the inherent constraints imposed by prebiotic chemistry.     

Changes in twelve homoeologous genomic regions in soybean following three rounds of polyploidy

With the advent of high-throughput sequencing, the availability of genomic sequence for comparative genomics is increasing exponentially. Numerous completed plant genome sequences enable characterization of patterns of the retention and evolution of genes within gene families due to multiple polyploidy events, gene loss and fractionation, and differential evolutionary pressures over time and across different gene families. In this report, we trace the changes that have occurred in 12 surviving homoeologous genomic regions from three rounds of polyploidy that contributed to the current Glycine max genome: a genome triplication before the origin of the rosids (~130 to 240 million years ago), a genome duplication early in the legumes (~58 million years ago), and a duplication in the Glycine lineage (~13 million years ago). Patterns of gene retention following the genome triplication event generally support predictions of the Gene Balance Hypothesis. Finally, we find that genes in networks with a high level of connectivity are more strongly conserved than those with low connectivity and that the enrichment of these highly connected genes in the 12 highly conserved homoeologous segments may in part explain their retention over more than 100 million years and repeated polyploidy events.     

Lemuriform origins as viewed from the fossil record

Fossils relevant to lemuriform origins are reviewed. Omanodon seems very close to the other early tooth-combed lemuriforms Karanisia, Wadilemur and Saharagalago, whereas Bugtilemur is rejected from the Lemuriformes. The Djebelemurinae, including Djebelemur and 'Anchomomys' milleri, are considered as stem lemuriforms preceding tooth comb differentiation; they are shown to be very distinct from European adapiforms. With tooth-combed lemuriforms present in Africa around 40 million years ago, and stem lemuriforms without tooth combs present on the same continent around 50-48 million years ago, a reasonable scenario can be proposed: tooth comb differentiation and lemuriform dispersal to Madagascar between 52-40 million years ago. The possible significance of Plesiopithecus for daubentoniid origins is raised. A critique of molecular dates is presented in the light of the fossil record. Azibiids are possibly early African prosimians. The timing of the dispersal of primates to Africa and the problem of strepsirhine origins are briefly examined.     

Evolution: red algal genome affirms a common origin of all plastids

Photosynthetic organelles (plastids) come in many forms and were originally thought to have multiple origins. The complete genome of the thermophilic red alga Cyanidioschizon merolae provides further evidence that all plastids derive from a single endosymbiotic event more than 600 million years ago.     

Molecular characterisation and evolution of the hemocyanin from the European spiny lobster,<Emphasis Type="Italic"> Palinurus elephas</Emphasis>

The hemocyanin of the European spiny lobster Palinurus elephas (synonym: Palinurus vulgaris) is a hexamer composed by four closely related but distinct subunits. We have obtained the full cDNA sequences of all four subunits, which cover 2275-2298 bp and encode for native polypeptides of 656 and 657 amino acids. The P. elephas hemocyanin subunits belong to the alpha-type of crustacean hemocyanins, whereas beta- and gamma-subunits are absent in this species. An unusual high ratio of non-synonymous versus synonymous nucleotide substitutions was observed, suggesting positive selection among subunits. Assuming a constant evolution rate, the P. elephas hemocyanin subunits emerged from a single hemocyanin gene around 25 million years ago. The alpha-type hemocyanins of P. elephas and the American spiny lobster Panulirus interruptus split around 100 million years ago. This is about five times older than the assumed divergence time of the species and suggests that the genera may have split with the formation of the Atlantic Ocean. The application of the Bayesian method for phylogenetic inference allows for the first time a solid reconstruction of the evolution of the decapod hemocyanins, showing that the beta-subunit types diverged first and that the crustacean pseudo-hemocyanins are associated with the gamma-type subunits.     

A comparative analysis of the Darwin-Wallace papers and the development of the concept of natural selection

The classical theory of descent with modification by means of natural selection had no mother, but did have two English fathers, Charles Darwin (1809–1882) and Alfred Russel Wallace (1823–1913). In 1858,the Linnean Society of London published two contributions of these naturalists and acknowledged both authors as the proponents of a novel hypothesis on the driving force of organismic evolution. In the present report the most important sections of the Darwin-Wallace papers are summarized. This close reading of both publications reveals six striking differences in emphasis: Darwin and Wallace did not propose identical ideas. The species definitions of both authors are described and the further development of the concept of natural selection in wild populations is reviewed. It is shown that the contributions of A.R. Wallace, who died 90 years ago, are more significant than usually acknowledged. I conclude that natural selection's lesser known co-discoverer should be regarded as one of the most important pioneers of evolutionary biology, whose original contributions are underestimated by most contemporary scientists.     

Do Women Prefer More Complex Music around Ovulation?

The evolutionary origins of music are much debated. One theory holds that the ability to produce complex musical sounds might reflect qualities that are relevant in mate choice contexts and hence, that music is functionally analogous to the sexually-selected acoustic displays of some animals. If so, women may be expected to show heightened preferences for more complex music when they are most fertile. Here, we used computer-generated musical pieces and ovulation predictor kits to test this hypothesis. Our results indicate that women prefer more complex music in general; however, we found no evidence that their preference for more complex music increased around ovulation. Consequently, our findings are not consistent with the hypothesis that a heightened preference/bias in women for more complex music around ovulation could have played a role in the evolution of music. We go on to suggest future studies that could further investigate whether sexual selection played a role in the evolution of this universal aspect of human culture.     

Phylogenetic perspectives on the origins of nodulation

Recent refinements to the phylogeny of rosid angiosperms support the conclusion that nodulation has evolved several times in the so-called N(2)-fixing clade (NFC), and provide dates for these origins. The hypothesized predisposition that enabled the evolution of nodulation occurred approximately 100 million years ago (MYA), was retained in the various lineages that radiated rapidly shortly thereafter, and was functional in its non-nodulation role for at least an additional 30 million years in each nodulating lineage. Legumes radiated rapidly shortly after their origin approximately 60 MYA, and nodulation most likely evolved several times during this radiation. The major lineages of papilionoid legumes diverged close to the time of origin of nodulation, accounting for the diversity of nodule biology in the group. Nodulation symbioses exemplify the concept of "deep homology," sharing various homologous components across nonhomologous origins of nodulation, largely due to recruitment from existing functions, notably the older arbuscular mycorrhizal symbiosis. Although polyploidy may have played a role in the origin of papilionoid legume nodules, it did not do so in other legumes, nor did the prerosid whole-genome triplication lead directly to the predisposition of nodulation.     

Did sex chromosome turnover promote divergence of the major mammal groups?

Comparative mapping and sequencing show that turnover of sex determining genes and chromosomes, and sex chromosome rearrangements, accompany speciation in many vertebrates. Here I review the evidence and propose that the evolution of therian mammals was precipitated by evolution of the male‐determining SRY gene, defining a novel XY sex chromosome pair, and interposing a reproductive barrier with the ancestral population of synapsid reptiles 190 million years ago (MYA). Divergence was reinforced by multiple translocations in monotreme sex chromosomes, the first of which supplied a novel sex determining gene. A sex chromosome‐autosome fusion may have separated eutherians (placental mammals) from marsupials 160 MYA. Another burst of sex chromosome change and speciation is occurring in rodents, precipitated by the degradation of the Y. And although primates have a more stable Y chromosome, it may be just a matter of time before the same fate overtakes our own lineage. Also watch the video abstract .     

Speciation, diversity, and Mode 1 technologies: the impact of variability selection

Over geological timescales, organisms encounter periodic shifts in selective conditions driven by environmental change. The variability selection hypothesis suggests that increases in environmental fluctuation have led to the evolution of complex, flexible behaviours able to respond to novel and unpredictable adaptive settings. This hypothesis is tested via the framework of a single locus genetic model in which an invading ‘versatilist’ allele competes with two opposed specialists in a selection regime driven by a fluctuating environment, modelled initially as a sine wave and subsequently as an empirical climate curve covering the past 5 million years. Results demonstrate that generalist alleles achieve fixation in the sine wave environment, whilst versatilist alleles do so in the empirical environment, even at a range of very low fitness advantages over the basic generalist template. Variability selection is found to be a particularly strong force between approximately 2.5 and 1.2 Ma (millions of years ago). These results are discussed in relation to the spread of Oldowan lithics and the patterns of speciation and extinction documented in the hominin fossil record. It is suggested that the flexibility required for survival in a variable climatic regime may have been a stimulus to the development of the first stone tool technologies, whilst the ecological opportunities provided by heightened variability may have been a factor in prompting the hominin adaptive radiation evidenced during this period.     

Clock gene evolution and functional divergence

In considering the impact of the earth's changing geophysical conditions during the history of life, it is surprising to learn that the earth's rotational period may have been as short as 4 h, as recently as 1900 million years ago (or 1.9 billion years ago). The implications of such figures for the origin and evolution of clocks are considerable, and the authors speculate on how this short rotational period might have influenced the development of the "protoclock" in early microorganisms, such as the Cyanobacteria, during the geological periodsin which they arose and flourished. They then discuss the subsequent duplication of clock genes that took place around and after the Cambrian period, 543 million years ago, and its consequences. They compare the relative divergences of the canonical clock genes, which reveal the Per family to be the most rapidly evolving. In addition, the authors use a statistical test to predict which residues within the PER and CRY families may have undergone functional specialization.     

Three novel mammalian toll-like receptors: gene structure, expression, and evolution

We describe three novel genes, encoding members of the Toll-like receptor (Tlr) family (TLR7, TLR8, and TLR9). These Tlr family members, unlike others reported to date, were identified within a genomic database. TLR7 and TLR8 each have three exons, two of which have coding function, and lie in close proximity to one another at Xp22, alongside a pseudogene. The remaining gene (TLR9) resides at 3p21.3 (in linkage with the MyD88 gene), and is expressed in at least two splice forms, one of which is monoexonic and one of which is biexonic, the latter encoding a protein with 57 additional amino acids at the N-terminus. The novel Tlrs comprise a cluster as nearest phylogenetic neighbors. Combining all sequence data related to Toll-like receptors, we have drawn several inferences concerning the phylogeny of vertebrate and invertebrate Tlrs. According to our best estimates, mammalian TLRs 1 and 6 diverged from a common mammalian ancestral gene 95 million years ago. TLR4, which encodes the endotoxin sensor in present-day mammals, emerged as a distinct entity 180 million years ago. TLRs 3 and 5 diverged from a common ancestral gene approximately 150 million years ago, as did Tlr7 and Tlr8. Very likely, fewer Tlrs existed during early vertebrate evolution: at most three or four were transmitted with the primordial vertebrate line. Phylogenetic data that we have adduced in the course of this work also suggest the existence of a Drosophila equivalent of MyD88, and indicate that the plasma membrane protein SIGIRR is close functional relative of MyD88 in mammals. Finally, a single present-day representative of the Toll-like proteins in Drosophila has striking cytoplasmic domain homology to mammalian Tlrs within the cluster that embraces TLRs 1, 2, 4, and 6. This would suggest that an ancestral (pre-vertebrate) Tlr may have adopted a pro-inflammatory function 500 million years ago.     

Evolution, systematics, and phylogeography of pleistocene horses in the new world: a molecular perspective

The rich fossil record of horses has made them a classic example of evolutionary processes. However, while the overall picture of equid evolution is well known, the details are surprisingly poorly understood, especially for the later Pliocene and Pleistocene, c. 3 million to 0.01 million years (Ma) ago, and nowhere more so than in the Americas. There is no consensus on the number of equid species or even the number of lineages that existed in these continents. Likewise, the origin of the endemic South American genus Hippidion is unresolved, as is the phylogenetic position of the "stilt-legged" horses of North America. Using ancient DNA sequences, we show that, in contrast to current models based on morphology and a recent genetic study, Hippidion was phylogenetically close to the caballine (true) horses, with origins considerably more recent than the currently accepted date of c. 10 Ma. Furthermore, we show that stilt-legged horses, commonly regarded as Old World migrants related to the hemionid asses of Asia, were in fact an endemic North American lineage. Finally, our data suggest that there were fewer horse species in late Pleistocene North America than have been named on morphological grounds. Both caballine and stilt-legged lineages may each have comprised a single, wide-ranging species.