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Mostly Out of Africa

6 & 7 November 2023 10.00am to 5.30pm
in the Flett Lecture Theatre, Natural History Museum, South Kensington, London SW7 5BD

In honour of Chris Stringer’s Huxley Medal, the RAI is organising a conference on the theme of ‘Mostly Out of Africa’.

‘Out of Africa’ is commonly used to represent the evolutionary origins of Homo sapiens, and dispersal from the African homeland about 60,000 years ago. However, there were no doubt several Out of Africa events in human evolution before this, and probably some ‘In to Africa’ dispersals as well. The theme of this conference is ‘Mostly Out of Africa’, reflecting both the wide range of Chris Stringer’s research interests, and the fact that genetic data have shown that the spread of H. sapiens was accompanied by bouts of interbreeding with Neanderthals and Denisovans. Thus we are mostly, but not entirely, of African origin.


Tickets must be purchased in advance for this event.

Tickets are £50 / £40 for RAI Fellows and Members / £25 Students

Lunch is not included, but there will be tea and coffee in the breaks and a drinks reception at the end of Tuesday 7 November.

To book your place please go to https://www.eventbrite.co.uk/e/mostly-out-of-africa-conference-and-huxley-lecture-tickets-726781863117?aff=oddtdtcreator (if you book a ticket for the conference this automatically includes a ticket for the Huxley Lecture and RAI AGM, you do not need to book separately for this).


Monday 6 November

Registration – 10.00-10.30

Introduction – 10.30

Panel Session 1 – 10.45-12.25

Lunch – 12.25-1.45

Panel Session 2 – 1.45-3.05

Break – 3.05-3.35

Panel Session 3 – 3.35-5.00

General Discussion – 5.00-5.30

Tuesday 7 November

Registration – 10.00-10.30

Panel Session 4 – 10.30-11.15

General Discussion – 11.15-12.00

Lunch – 12.00-1.30

Huxley Lecture by Chris Stringer – 1.30-2.30

RAI AGM – 2.30-4.30

Drinks Reception – 4.30-5.30

Monday 6 November 2023

10.00-10.30 Registration

10:30 Introduction

Session 1 – Entirely out of Africa: Ancient Britain

10:45 Nick Ashton (British Museum)
A re-evaluation of the emergence of modern behaviour from a European perspective

There has been long debate about what constitutes modern behaviour with much of the focus on Africa and the emergence of Homo sapiens. But there has also been recognition by many researchers that facets of modern behaviour can be traced back into the Middle Pleistocene and that a multi-regional approach addresses the question more broadly by contributing evidence from other hominin species. This paper re-evaluates some of the evidence from Europe over the last half million years, drawing on developments in technology, economising behaviour, landscape use, social organisation, and emblematic use of material culture.

11:05 Matthew Pope (University College London)
The Middle to Upper Palaeolithic Transition in northern France and southern Britain: Approaching a challenging record

Two Palaeolithic sites, each flanking the Western Approaches of the English Channel, preserved fossil human material relating to populations which may have lived close in time to the Middle Upper Palaeolithic transition in the region. Kents Cavern in south-western Britain preserved a fragment of human jaw now assigned to Homo sapiens with a date modelled to 44 – 41 kya. 171 miles to the south, at La Cotte de St Brelade, Jersey, 13 human teeth show a mosaic of Homo neanderthalensis and Homo sapiens features and may date to <48 kya. In both cases, the fossil material was recovered during old excavations and, despite recent and ongoing reanalysis of associated archives, it is so far impossible to determine their precise age and cultural associations. More broadly, understanding the nature and timing of replacement of Neanderthal people by Homo sapiens in wider La Manche region is made difficult due to both the lack of sequenced DNA from any of these fossils and the relative absence of well-dated ‘transitional’ or Initial Upper Palaeolithic assemblages. The absence of the Châtelperronian in northern France and Britain alongside the absence of the LRJ, and Early Aurignacian in north-west France, and only poorly dated occurrences of either technology in Southern Britain presents a challenging research environment. In this paper we review the sparse evidential record for the Middle to Upper Palaeolithic transition in the La Manche region. We discuss the implications of the current data gap and how, on both sides of the English Channel, we might develop proactive approaches to researching this critical evolutionary event on the edge of the human range.

11:25 Rob Dinnis (University of Aberdeen)
Reconstructing the early Homo sapiens occupations of Britain: the archaeological potential of Wogan Cavern

Relative to elsewhere in Europe, our understanding of Britain’s earliest Homo sapiens occupations remains extremely limited. In large part this is due to extensive early excavations, which completely or almost completely emptied Britain’s most important archaeological caves. Extant collections from this work are usually of limited research value. To meaningfully improve our understanding will require new intact archaeological deposits, and probably a new site.

Current fieldwork at Wogan Cavern, a large cave underlying Pembroke Castle, is demonstrating its potential as such a site. Test excavations have shown the presence of undisturbed Pleistocene and early Holocene sedimentary deposits in several parts of the cave. Small-scale testing of these has revealed evidence for several periods of early prehistoric activity, including an Early Upper Palaeolithic layer with clear similarity to the historical collection from Paviland Cave, one of Britain’s most important early H. sapiens sites. Although in its early stages, work at Wogan Cavern has already shown it to be one of Britain’s most significant archaeological caves, and that it has the potential to address many outstanding questions about early H. sapiens occupation of northwestern Europe.

11:45 Silvia M. Bello (Natural History Museum)
125 Years of Research at Gough’s Cave (Somerset, UK)

Gough’s Cave (Somerset, UK) is the richest Upper Palaeolithic site in the British Isles in terms of human remains, stone and organic artefacts, and butchered faunal remains. The cave has a long history of investigations that stretches back to the discovery and opening of the cave as a tourist attraction by Richard Cox Gough between 1892 and 1898. The remaining deposits at the front of the cave were almost completely excavated by R.F. Parry over several winter seasons between 1927 and 1932 and A.V. Painter continued work up to 1953. The most recent investigations were directed by the late Roger Jacobi, Andy Currant and Chris Stringer, who undertook exploratory work in 1986-87, 1989-90.

Our knowledge of the recolonization of north-west Europe at the end of the Last Glacial Maximum depends to a large extent on finds from Gough’s Cave. The artefact collection from Gough’s Cave is impressive for a British Late Upper Palaeolithic site with numerous flint tool-types based on blades and bladelets, bone, antler and ivory artefacts. The faunal assemblage suggests the site functioned as a short-lived, multi-activity seasonal camp with a focus on horse and red deer hunting. A particularly intriguing aspect of the human bone assemblage is the evidence of butchery of the cadavers with traces from scalping, intensive cleaning of the bones to remove edible soft tissue, fracturing of bone shafts to extract marrow, and chewing and consumption of spongy bone. Heads were also modified to make skull cups, and a radius is marked with a chevron motif, suggesting that cannibalism was ritual in nature. Recent aDNA analyses of the human remains, the oldest human DNA thus far obtained from Britain or Ireland, have determined that Late Upper Palaeolithic individuals from Gough’s Cave probably traced all their ancestry to Magdalenian-associated individuals closely related to those from sites such as El Mirón Cave, Spain, and Troisième Caverne in Goyet, Belgium.

In this talk, I will review how the contributions of many researchers working on Gough’s Cave over the past 125 have been instrumental in transforming our understanding of the Upper Palaeolithic in Europe.

12:05 William Marsh (Natural History Museum)
Using ancient biomolecules to reveal the social structure and behaviours of the Magdalenian at the Upper Palaeolithic cave site of Gough’s Cave, UK

The human remains excavated from Gough’s Cave represents the largest deposit in Britain associated with the Upper Palaeolithic Magdalenian culture (dated to ~15k calBP). Evidence of anthropic modifications associated with processing of human remains post-mortem and cannibalism is prevalent across the 205 cranial and postcranial remains (MNI=6), with the most remarkable examples including three skull cups manufactured from cranial vaults. Prior biomolecular results have shown one individual to share ancestry with the genetic cluster associated with Magdalenian populations across western Europe, with two separate individuals showing bulk isotope signatures consistent with a terrestrial diet. Here, we expand these biomolecular methods onto a larger set of skeletal remains to further clarify the composition, ancestry, and diet of the group, which when combined with previous archaeological studies develop our understanding of the Magdalenian more generally. Compound specific isotope analyses support an omnivorous diet, with no measurable marine component. Of the six individuals sequenced from the cave, all show a genetic affinity to the GoyetQ2 cluster associated with the Magdalenian culture. Four individuals show a second degree or higher degree of relatedness, with two separate close family groups identified. Both sexes are represented, which when considered alongside the presence of juvenile remains suggests group composition mirroring that seen in modern hunter gatherer groups. A low effective population size is inferred, along with a lack of close kin breeding, compatible with sexual behaviours that avoided close union despite the inferred high level of group relatedness. The skeletal elements for five sequenced individuals show evidence of cannibalism and post-mortem modification, indicative of endocannibalism and a kin-mediated funerary behaviour. Genetic results conform with the predicted group structure of the Magdalenian where small groups containing related individuals maintained broader connections to neighbouring regional groups allowing for resource sharing and the avoidance of consanguineous relationships.

12:25-1:45 Lunch

Session 2 Mostly out of Africa 1: Chronology and Regional Studies

1:45 Rainer Grün (Australian National University and University of Tübingen)
Early Homo sapiens in and out of Africa

It has been established for some 40 years that anatomically modern humans appeared in the Near East some 100,000 years ago at Skhul and Qafzeh. Within Africa, modern humans have been dated to about 300,000 years (Irhoud) and ~260,000 years (Florisbad) and ~ 200,000 years (Omo). Recent studies claimed that the “oldest” Homo sapiens outside Africa appeared in the Levant at around 180,000 years ago (Mislya) and in Europe even earlier, at > 210,000 years ago (Apidima). I will discuss the dating results and the criticism thereof in more detail.

2:05 Katerina Harvati (University of Tübingen)
Modern human origins and dispersals: the view from South-East Europe

South-East Europe is commonly thought to lie directly on the principal migration route through which modern humans dispersed from Africa to Europe through the Near East. Despite its hypothesised central position as an early entry point to the European continent, little evidence exists to support this idea. Indeed it is common for maps of Paleolithic Europe to indicate population dispersals as arrows that either entirely bypass or ‘fly over’ much of the Balkan region. Today I will present evidence from my team’s work in Greece both in the laboratory and in the field, which brings new data to the debate and highlights the regions’ biogeographic importance as both dispersal corridor and glacial refugium for human populations. Results strongly suggest that the paucity of currently available data is due to lack of research, and highlight the potential of the Balkan’s record to shed light on the complex processes underlying human evolution in Europe.

2:25 Huw Groucutt (University of Malta, Max Planck Institutes for Geoanthropology, Chemical Ecology, and Biogeochemistry, and University of Cologne)
Human evolution at the interface of Africa and Eurasia

That all living and recent humans are of purely or overwhelmingly African origin (the Recent African Origin model) is one of the most important scientific developments in recent decades. Understanding how our species then expanded beyond Africa, and mixed with the Eurasian Neanderthals, are major questions which then flow from this model. In this paper I will evaluate recent research in Northeast Africa and Southwest Asia which contribute to the understanding of the origin and spread of our species. In particular, I will explore how archaeological evidence can contribute in areas where very few hominin fossils are known. This is explored in terms of the biogeographical context, such as whether early Homo sapiens dispersals were the result of ‘Africa growing’. Finally, the uncertainty around chronological age estimates for key fossil, archaeological, and palaeoclimatic findings will be evaluated. While much has been learned about the evolution and spread of Homo sapiens, many lacunae and uncertainties remain and it is argued that these need to be recognized and considered so as to most fruitfully aim future research.

2:45 Eleanor Scerri (Max Planck Institute of Geoanthropology, University of Cologne, and University of Malta)
Human Origins: The View from West Africa

Multiple, independent, lines of evidence confirm a structured African origin of our species, Homo sapiens. However, many questions remain regarding the character of early human population structure, and its physical, behavioural, and evolutionary consequences. These gaps in knowledge in part exist because of knowledge gaps in vast swathes of Africa. Recent work across a West African transect is attempting to over come this barrier in a major region of the continent. New data from Senegal, the Ivory Coast, Nigeria, Benin, and Guinea is providing critical new evidence from which to test expectations of a Pan-African model by evaluating the age of the Middle Stone Age, the time-depth of human habitation in rainforests, the signatures of regionally-specific material culture sequences, and the diversity of human-environment interactions. In this paper, I will present the most recent results of this work and consider both how different African regions mediated population structure and what role they played in the origins of our species.

3:05-3:35 Tea & Coffee Break

Session 3 Mostly out of Africa 2: Dispersals, phylogeny and behaviour

3:35 Thomas Higham (University of Vienna)
Tracking the earliest dispersals of Homo sapiens across Eurasia

Obtaining reliable chronologies is key if we are to track human dispersals out of Africa. The latter part of the story of human evolution occurs at just the time that radiocarbon dating reaches its limit of 50-60,000 years. Improvements to the reliability of the method have been necessary to generate dates that are not contaminated, and can be trusted. In this talk I will outline some of the most recent work in this area and in the application of improved methods (including non-radiocarbon ages) from some of the key sites on the path of human dispersal in Eurasia.

3:55 Marta Lahr (University of Cambridge) and Rob Foley (University of Cambridge)
Hominin dispersals: causes, rates and outcomes

The out of Africa model, from its first inception in the 1980s, put an emphasis on dispersals as a mechanism driving human evolution, distribution and diversity. Dispersal has remained at the heart of virtually all subsequent iterations of the model. However, across this time virtually all aspects of hominin dispersals have been debated and challenged – one or many, only out of Africa or both ways, with or without admixture, driven by external climatic factors or human adaptive success. In this paper we will consider the current evidence for hominin dispersals related to recent human evolution, and discuss the drivers of range expansion and the conditions shaping different outcomes. As dispersal models have become increasingly important in biogeography more generally, we also examine how models of hominin diversity fit into a broader framework of evolutionary processes.

4:15 Xijun Ni (Chinese Academy of Science) and Chris Stringer (Natural History Museum)
Homo phylogeny and source-sink of diversification

The genus Homo contains quite diverse species, probably more diverse than many scientists expected. The traditional idea postulated that there had been no speciation event since the origin of Homo erectus, resulting in all later Homo fossils being included in a broad and vaguely defined species of Homo sapiens, but later researchers found that Homo neanderthalensis, Homo heidelbergensis, and other groups are distinct enough to be considered species. More recent discoveries, such as Homo floresiensis, Homo antecessor, Homo luzonensis, Homo naledi, and Homo longi, have provided even stronger evidence that several independent species coexisted with H. sapiens, especially during the Middle Pleistocene. The evolutionary relationships among these different Homo species are critical for understanding the origin of H. sapiens. We developed a phenomic character data matrix containing 632 discrete and metric characters scored for 55 Homo species/populations. We used parsimony analysis and Bayesian tip-dating inference to reconstruct the Homo phylogeny and estimate the divergence time of all Homo clades. Our results suggest that H. sapiens and H. neanderthalensis are monophyletic groups that arose around 770 ka and 737 ka, respectively. Asian H. longi-like Homo, such as Dali, Hualongdong, Jinniushan, and Xiahe form another monophyletic group that diverged from H. sapiens around 949 ka. H. neanderthalensis has traditionally been regarded as the sister group of H. sapiens, but our results showed that this Asian monophyletic group is the sister group of H. sapiens. Asian, European and African H. erectus belong to a paraphyletic group. African and European H. heidelbergensis-like Homo, such as Broken Hill, Saldanha, Bodo, Petralona, Ceprano, and Arago, also belong to a paraphyletic group. Based on the Homo phylogeny, we tested 18 biogeographic models and estimated the number and type of biogeographic events. Our simulation suggests that instead of a unidirectional “out of Africa” model, a multidirectional “shuttle dispersal” model is more likely to explain the complex phylogenetic relationships among African and Eurasian Homo species/populations. The common ancestor of H. longi-clade and H. sapiens dispersed from Africa to Eurasia between 949 ka and 917 ka. The common ancestor of H. sapiens and H. neanderthalensis dispersed from Europe to Africa between 1007 ka and 978 ka. H. sapiens dispersed from Africa to Eurasia between 490 ka and 416 ka. Africa gave more dispersals to other continents, while Asia received more dispersals from other continents. From this perspective, we suggest that Africa is the “source” of evolution and biogeographic dispersal, while Asia is the “sink” of evolution and dispersal of Homo.

4:35 Francesco d’Errico (Université de Bordeaux, Centre for Early Sapiens Behaviour (SapienCE), and University of Bergen)
The archaeology of modern human origin

How can the study of past material culture shed light on the process that led to the emergence of our species and its unique cognition in the animal kingdom? The scenario proposing that our species was the outcome of a punctuated event in Eastern or Southern Africa, 200,000 years ago, carries implications going well beyond paleoanthropology. By giving rise to our species, natural selection would have provided this small modern population with extra advantages: new cognitive faculties and behavioural correlates such a language comparable to ours, and the ability to develop and transmit complex technological innovations and symbolic cultures. Armed with these advantages, modern humans would have rapidly replaced anatomically and cognitively archaic populations, first in Africa and later in Eurasia. In contrast, the scenario that envisions the origin of modern populations in Africa as the result of a gradual and pan-African process lasting for at least 300,000 years implies a gradual and probably asynchronous emergence of modern cognitive abilities across the entirety of this continent. Building on my recent and ongoing research in Africa and Eurasia I will argue that archaeological discoveries made in the last two decades support the latter scenario. Innovations often interpreted as reflecting modern cognition (use of pigments, abstract and figurative representations, ornaments, burials, elaborate techniques of stone knapping, complex osseous technologies) emerge gradually in Africa and appear in certain regions before others. If we add to this picture data supporting systematic genetic exchanges between modern and archaic populations, and the fact that some of these latter populations develop key innovations in parallel with their African counterparts, we reach the inescapable conclusion that Paleolithic human populations considered distinct species in the past must now be regarded as the phenotypic expression of the same evolving species, characterized by strong cognitive plasticity, also in evolution. As a consequence, it becomes essential to focus our attention on the conditions and mechanisms that have triggered cultural innovations at regional scale, and on the factors that have enabled them to be preserved and disseminated.

5:00-5:30 General Discussion


Tuesday 7 November 2023

10:00-10:30 Registration

Session 4 Mostly out of African 3: Genomics

10:30 Pontus Skoglund (The Francis Crick Institute)
Origins of modern human ancestry: genetic approaches to its reconstruction

New finds in the fossil and genomic record have changed our view of the origins of modern human ancestry. I will review our current understanding of how the ancestry of modern humans can be traced into the deep past, focusing on key phases that are surrounded by major questions. Of particular interest is the broadly African origin of modern human diversity between 60 and 300 ka, where much of our understanding is still fragmentary. I will discuss why no specific point in time can currently be identified at which modern human ancestry was confined to a limited birthplace, and approaches in computational genomics and ancient proteomics that could advance our understanding of the origin of our species and its relationship to others.

10:50 Mateja Hajdinjak (Max Planck Institute for Evolutionary Anthropology)
Re-examining the genetic relationships among Neandertal populations

Before their disappearance at around 40,000 years before present (i.e. ~40 ka BP), Neandertals lived throughout Europe, western and central Asia, and the Near East, appearing in the European fossil record at least ~430 ka BP. However, despite genomic data being recovered from more than 10,000 ancient humans to date, the genomic data recovered from Neandertals are still relatively sparse. Thus far, genome-wide data have been retrieved for 31 Neandertals from 16 archaeological sites spanning their long history and across large parts of their geographical range. These data have offered a broad overview of Neandertal populations, indicating the existence of multiple distinct groups over time and space, as well as the occurrences of multiple population turnovers and migrations. Here, I will summarize and re-examine what we currently know about the relationships among different Neandertal populations using their nuclear, Y-chromosomal and mtDNA data, with a specific focus on the Neandertals from Gibraltar, which have for a long time been considered as some of the last Neandertals in Europe. I will also touch upon what the new analyses of the old data can tell us about the relationship of different Neandertal groups to the Neandertals that mixed with the ancestors of all present-day and ancient humans outside of sub-Saharan Africa.

11:10 Matthias Meyer (Max Planck Institute for Evolutionary Anthropology) and the Ancient Environmental DNA Analysis Consortium
Beyond bones and teeth – exploring alternative sources of ancient DNA for investigating the human past

The retrieval of complete and partial genome sequences from the skeletal remains of Neandertals, Denisovans and modern humans has greatly improved our understanding of the history of these groups, their relatedness and interactions. However, many questions about the human past cannot be answered by analysing human skeletal remains alone, which are often rare or absent in archaeological assemblages. In 2017, we reported on the recovery of Neandertal and Denisovan mitochondrial DNA from Pleistocene cave sediments, suggesting that sediment DNA analysis could overcome our dependency on the scarce fossil record from this period. However, this work also raised a number of questions that are important in determining the significance of this newly discovered DNA source for future research: How common is the preservation of ancient human DNA in sediments? What are the temporal limits of preservation of ancient DNA in sediments? What is the source of this DNA? Do individual samples contain DNA from one or more individuals? Can human DNA move between layers? What are the limits of resolution that can be achieved when analysing DNA sequences from sediment?

To address these questions we conducted several large studies at different scales. On a global scale, we screened sediment samples from more than 200 archaeological sites, mainly in Eurasia, to assess the preservation of ancient human and faunal DNA. On a local scale, we reconstructed genetic time-transects from archaeological sites based on hundreds of sediment samples. In Denisova Cave in Russia, for example, we detected the DNA of Neandertals, Denisovans and modern humans in 175 out of 728 samples, allowing the reconstruction of the site’s occupational history with unprecedented resolution. At the Galería de las Estatuas in Spain, we expanded the search of DNA in sediment from mitochondrial to nuclear DNA, which allowed us to detect a turnover in the Neandertal populations that occupied the site. On a microscale, we analyzed DNA from blocks of sediment that had been impregnated in resin for micromorphological analysis, targeting microstructures derived from bone fragments, coprolites and minerals. Taken together, the results of these studies show that ancient human DNA is a very small but frequently detectable component of DNA in Pleistocene cave sediment, that isolation of DNA from single individuals from sediment is possible, and that movement of DNA across archaeological layers is not a common phenomenon.

More recently, we succeeded in isolating ancient human DNA from Palaeolithic tooth pendant, demonstrating that DNA analysis of artefacts made from bones and teeth may represent another avenue towards linking the cultural and genetic records of the past. Current efforts are focused on improving and simplifying the methods for extracting and analysing DNA sequences from sediments and artefacts, with the aim of making DNA analysis an even more widely used tool in prehistoric archaeology.

11:30 General Discussion

12:00-1:30 Lunch

Chris Stringer
(Natural History Museum)
Mostly Out of Africa: how, when and where

During the last 50 years the Recent African Origin model has risen from nowhere to become the dominant explanation for the origins of our species Homo sapiens. However, there are many outstanding issues in resolving the details of how, when and where we evolved. It is not yet clear whether the last common ancestor (LCA) of H. sapiens, Neanderthals, and Denisovans lived in Europe, Asia, or Africa, and when it lived. Estimates using genomic data from H. sapiens, Neanderthals, and Denisovans calibrate the LCA to between about 500,000 and 700,000 years ago. However, some recent stud­ies of dental and cranial variation in fossil hominins place the LCA earlier, between about 800,000 and 1.2  million years ago. Thus identifying potential candidate fossils for the LCA is highly problematic against these uncertainties. Moreover, some genetic models for the deep ancestry of H. sapiens and Neanderthals suggest that concepts of a single LCA in time and space might be illusory.

A further problem is that the fossil human record from Africa for the period between about 200,000 and 1 million years ago is relatively sparse, and yet this timespan was the critical one for the evolution of Homo sapiens. The record we have is generally not well-dated, and limited to small areas of the continent, with no fossils at all from large regions of West and Central Africa. In the past, researchers have proposed a range of different models, from those with a single location of origin (usually East or South Africa, which have the best fossil records for the period) through to pan-African models (sometimes also called African multiregionalism, a term I prefer not to use now because of potential confusion with the refuted global multiregional theory). Pan-African models envisage lineages of basal H. sapiens populations in different regions of Africa that evolved separately at times, but which also sporadically mixed and merged with each other, finally giving rise to the form of Homo sapiens that eventually spread globally.

That global spread is usually assumed to have been initiated about 60,000 years ago, but there is increasing evidence of earlier dispersals from Africa, at least one of which led to an episode of gene flow with a basal Neanderthal lineage. We now know that the main dispersal was also accompanied by bouts of gene flow with the Neanderthal and Denisovan lineages – hence the term Mostly Out of Africa (which I first heard being used by Svante Pääbo) is the most appropriate simple description for the evolutionary origins of our species.

2:30-4:30 RAI AGM

4:30-5:30 Drinks Reception

Location: Flett Lecture Theatre
Natural History Museum
South Kensington