Pan-Africanism vs. single-origin of Homo sapiens: Putting the debate in the light of evolutionary biology.

The scenario of Homo sapiens origin/s within Africa has become increasingly complex, with a pan-African perspective currently challenging the long-established single-origin hypothesis. In this paper, we review the lines of evidence employed in support of each model, highlighting inferential limitations and possible terminological misunderstandings. We argue that the metapopulation scenario envisaged by pan-African proponents well describes a mosaic diversification among late Middle Pleistocene groups. However, this does not rule out a major contribution that emerged from a single population where crucial derived features-notably, a globular braincase-appeared as the result of a punctuated, cladogenetic event. Thus, we suggest that a synthesis is possible and propose a scenario that, in our view, better reconciles with consolidated expectations in evolutionary theory. These indicate cladogenesis in allopatry as an ordinary pattern for the origin of a new species, particularly during phases of marked climatic and environmental instability.

The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal.

A combination of evidence, based on genetic, fossil and archaeological findings, indicates that Homo sapiens spread out of Africa between ~70-60 thousand years ago (kya). However, it appears that once outside of Africa, human populations did not expand across all of Eurasia until ~45 kya. The geographic whereabouts of these early settlers in the timeframe between ~70-60 to 45 kya has been difficult to reconcile. Here we combine genetic evidence and palaeoecological models to infer the geographic location that acted as the Hub for our species during the early phases of colonisation of Eurasia. Leveraging on available genomic evidence we show that populations from the Persian Plateau carry an ancestry component that closely matches the population that settled the Hub outside Africa. With the paleoclimatic data available to date, we built ecological models showing that the Persian Plateau was suitable for human occupation and that it could sustain a larger population compared to other West Asian regions, strengthening this claim.

Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa.

The population dynamics that followed the Out of Africa (OoA) expansion and the whereabouts of the early migrants before the differentiation that ultimately led to the formation of Oceanian, West and East Eurasian macropopulations have long been debated. Shedding light on these events may, in turn, provide clues to better understand the cultural evolution in Eurasia between 50 and 35 ka. Here, we analyze Eurasian Paleolithic DNA evidence to provide a comprehensive population model and validate it in light of available material culture. Leveraging on our integrated approach we propose the existence of a Eurasian population Hub, where Homo sapiens lived between the OoA and the broader colonization of Eurasia, which was characterized by multiple events of expansion and local extinction. A major population wave out of Hub, of which Ust'Ishim, Bacho Kiro, and Tianyuan are unadmixed representatives, is broadly associated with Initial Upper Paleolithic lithics and populated West and East Eurasia before or around 45 ka, before getting largely extinct in Europe. In this light, we suggest a parsimonious placement of Oase1 as an individual related to Bacho Kiro who experienced additional Neanderthal introgression. Another expansion, started before 38 ka, is broadly associated with Upper Paleolithic industries and repopulated Europe with sporadic admixtures with the previous wave (GoyetQ116-1) and more systematic ones, whereas moving through Siberia (Yana, Mal'ta). Before these events, we also confirm Zlatý Kun as the most basal human lineage sequenced to date OoA, potentially representing an earlier wave of expansion out of the Hub.

Mosaic evolution in hominin phylogeny: meanings, implications, and explanations.

In paleoanthropological literature, the use of the term "mosaic" (mosaic evolution, mosaic trait, mosaic species, and so on) is becoming more and more frequent. In order to promote a clarification of the use of the concept in literature, we propose here a classification in three different meanings of the notion of mosaic in human evolution: 1) morphological (inter-specific and intra-specific) instability in a certain phase of a branched phylogeny; 2) multiple trajectories and versions of the same adaptive trait in a branched phylogeny; 3) the trait itself as a complex mosaic of sub-traits with different phylogenetic stories (as is the case in language). We argue that the relevance of such mosaic patterns needs a macro-evolutionary interpretation, which takes into consideration the interaction between general selective pressures (promoting different versions of the same adaptation) and a cladogenetic approach in which speciation played a crucial role, due to ecological instability, habitat fragmentation, and geographical dispersals in human evolution.

Mirror neurons in the tree of life: mosaic evolution, plasticity and exaptation of sensorimotor matching responses.

Considering the properties of mirror neurons (MNs) in terms of development and phylogeny, we offer a novel, unifying, and testable account of their evolution according to the available data and try to unify apparently discordant research, including the plasticity of MNs during development, their adaptive value and their phylogenetic relationships and continuity. We hypothesize that the MN system reflects a set of interrelated traits, each with an independent natural history due to unique selective pressures, and propose that there are at least three evolutionarily significant trends that gave raise to three subtypes: hand visuomotor, mouth visuomotor, and audio-vocal. Specifically, we put forward a mosaic evolution hypothesis, which posits that different types of MNs may have evolved at different rates within and among species. This evolutionary hypothesis represents an alternative to both adaptationist and associative models. Finally, the review offers a strong heuristic potential in predicting the circumstances under which specific variations and properties of MNs are expected. Such predictive value is critical to test new hypotheses about MN activity and its plastic changes, depending on the species, the neuroanatomical substrates, and the ecological niche.

Multi-level human evolution: ecological patterns in hominin phylogeny.

Evolution is a process that occurs at many different levels, from genes to ecosystems. Genetic variations and ecological pressures are hence two sides of the same coin; but due both to fragmentary evidence and to the influence of a gene-centered and gradualistic approach to evolutionary phenomena, the field of paleoanthropology has been slow to take the role of macro-evolutionary patterns (i.e. ecological and biogeographical at large scale) seriously. However, several very recent findings in paleoanthropology stress both climate instability and ecological disturbance as key factors affecting the highly branching hominin phylogeny, from the earliest hominins to the appearance of cognitively modern humans. Allopatric speciation due to geographic displacement, turnover-pulses of species, adaptive radiation, mosaic evolution of traits in several coeval species, bursts of behavioral innovation, serial dispersals out of Africa, are just some of the macro-evolutionary patterns emerging from the field. The multilevel approach to evolution proposed by paleontologist Niles Eldredge is adopted here as interpretative tool, and has yielded a larger picture of human evolution that integrates different levels of evolutionary change, from local adaptations in limited ecological niches to dispersal phenotypes able to colonize an unprecedented range of ecosystems. Changes in global climate and Earth's surface most greatly affected human evolution. Precisely because it is cognitively hard for us to appreciate the long-term common destiny we share with the whole biosphere, it is particularly valuable to highlight the accumulating evidence that human evolution has been deeply affected by global ecological changes that transformed our African continent of origin.

Individuals and groups in evolution: Darwinian pluralism and the multilevel selection debate.

Outlined here is an updated review of the long-standing 'kin selection vs group selection' debate. Group selection is a highly contentious concept, scientifically and philosophically. In 2012, Dawkins' attack against Wilson's latest book about eusociality concentrated all the attention on group selection and its mutual exclusivity with respect to inclusive fitness theory. Both opponents seem to be wrong, facing the general consensus in the field, which favours a pluralistic approach. Historically, despite some misunderstandings in current literature, such a perspective is clearly rooted in Darwin's writings, which suggested a plurality of levels of selection and a general view that we propose to call 'imperfect selfishness'. Today, the mathematically updated hypothesis of group selection has little to do with earlier versions of 'group selection'. It does not imply ontologically unmanageable notions of 'groups'. We propose here population structure as the main criterion of compatibility between kin selection and group selection. The latter is now evidently a pattern among others within a more general 'multilevel selection' theory. Different explanations and patterns are not mutually exclusive. Such a Darwinian pluralism is not a piece of the past, but a path into the future. A challenge in philosophy of biology will be to figure out the logical structure of this emerging pluralistic theory of evolution in such contentious debates.

Italian natural history museums on the verge of collapse?

The Italian natural history museums are facing a critical situation, due to the progressive loss of scientific relevance, decreasing economic investments, and scarcity of personnel. This is extremely alarming, especially for ensuring the long-term preservation of the precious collections they host. Moreover, a commitment in fieldwork to increase scientific collections and concurrent taxonomic research are rarely considered priorities, while most of the activities are addressed to public events with political payoffs, such as exhibits, didactic meetings, expositions, and talks. This is possibly due to the absence of a national museum that would have better steered research activities and overall concepts for collection management. We here propose that Italian natural history museums collaborate to instate a "metamuseum", by establishing a reciprocal interaction network aimed at sharing budgetary and technical resources, which would assure better coordination of common long-term goals and scientific activities.

Many ways of being human, the Stephen J. Gould's legacy to Palaeo-Anthropology (2002-2012).

As an invertebrate palaeontologist and evolutionary theorist, Stephen J. Gould did not publish any direct experimental results in palaeo-anthropology (with the exception of Pilbeam and Gould, 1974), but he did prepare the stage for many debates within the discipline. We argue here that his scientific legacy in the anthropological fields has a clear and coherent conceptual structure. It is based on four main pillars: (1) the famed deconstruction of the "ladder of progress" as an influential metaphor in human evolution; (2) Punctuated Equilibria and their significance in human macro-evolution viewed as a directionless "bushy tree" of species; (3) the trade-offs between functional and structural factors in evolution and the notion of exaptation; (4) delayed growth, or neoteny, as an evidence in human evolution. These keystones should be considered as consequences of the enduring theoretical legacy of the eminent Harvard evolutionist: the proposal of an extended and revised Darwinism, coherently outlined in the last twenty years of his life (1982-2002) and set out in 2002 in his final work, The Structure of Evolutionary Theory. It is in the light of his "Darwinian pluralism", able to integrate in a new frame the multiplicity of explanatory patterns emerging from different evolutionary fields, that we understand Stephen J. Gould's legacy in palaeo-anthropology today, both in terms of provocative shocks to comfortable visions of human evolution and, above all, in terms of specific scientific predictions about future research.

Exaptation in human evolution: how to test adaptive vs exaptive evolutionary hypotheses.

Palaeontologists, Stephen J. Gould and Elisabeth Vrba, introduced the term "ex-aptation" with the aim of improving and enlarging the scientific language available to researchers studying the evolution of any useful character, instead of calling it an "adaptation" by default, coming up with what Gould named an "extended taxonomy of fitness". With the extension to functional co-optations from non-adaptive structures ("spandrels"), the notion of exaptation expanded and revised the neo-Darwinian concept of "pre-adaptation" (which was misleading, for Gould and Vrba, suggesting foreordination). Exaptation is neither a "saltationist" nor an "anti-Darwinian" concept and, since 1982, has been adopted by many researchers in evolutionary and molecular biology, and particularly in human evolution. Exaptation has also been contested. Objections include the "non-operationality objection".We analyze the possible operationalization of this concept in two recent studies, and identify six directions of empirical research, which are necessary to test "adaptive vs. exaptive" evolutionary hypotheses. We then comment on a comprehensive survey of literature (available online), and on the basis of this we make a quantitative and qualitative evaluation of the adoption of the term among scientists who study human evolution. We discuss the epistemic conditions that may have influenced the adoption and appropriate use of exaptation, and comment on the benefits of an "extended taxonomy of fitness" in present and future studies concerning human evolution.