Christophe Amiot
email: christophe.amiot@univ-angers.fr

Weihong Ji
email: weihong@massey.ac.nz

Erle C Ellis

Michael G Anderson
email: m.g.anderson@massey.ac.nz

Temporal and sociocultural effects of human colonisation on native biodiversity: filtering and rates of adaptation

Keywords: Human colonisation, Extinction / Extirpation, Native biodiversity, Evolutionary history, Sociocultural niche construction

ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

MAIN TEXT

Colonisation of habitats by modern human societies, have radically changed local and regional native species richness and the ecological and evolutionary processes of ecosystems [START_REF] Koch | Late quaternary extinctions: State of the debate[END_REF][START_REF] Barnosky | Megafauna biomass tradeoff as a driver of Quaternary and future extinctions[END_REF][START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF], Malhi et al. 2016). As a hypothesis to explain these changes, we propose that the rates and types of changes in native species richness following modern human colonisation are mediated by the temporal dynamics and types of the coloniser's societies.

Specifically, rates of extinction and extirpation following modern human colonisation can be understood as analogous to habitat filtering processes [START_REF] Cornwell | A trait-based test for Habitat filtering: Convex hull volume[END_REF][START_REF] Kraft | Community assembly, coexistence and the environmental filtering metaphor[END_REF]. In this framework, time elapsed since colonisation, the number of times colonised by a different society type and its sociocultural regime at the time of colonisation all play significant roles in shaping the ecological and evolutionary responses to sociocultural filtering processes.

Previous studies have shown a divergence in life history traits through nest predation constraints between regions colonised by humans at different times [START_REF] Martin | Nest predation and avian life-history evolution in Europe versus North America: a possible role of humans?[END_REF].

Rates of change (e.g., alterations of reproductive strategies) may be more rapid in sites more recently colonised by humans [START_REF] Cartwright | Anthropogenic natal environment of effects on life histories in a wild bird population[END_REF]. We propose that these patterns result from two processes; 1) anthropogenic filtering, whereby extinctions of non-adaptive species occur rapidly following colonisation, and 2) adaptation processes occur within species that survive the habitat changes following colonisation. We suggest that these processes can be applied together with well-established biogeographic principles, such as latitude, altitude, habitat size and degree of isolation (Mc Arthur 1972[START_REF] Helmus | Island Biogeography of the Anthropocene[END_REF], to predict current and future patterns of native species richness and life-history traits, especially with regard to the capacity of species to adapt to changing environments and climate. By incorporating the temporal and sociocultural aspects of human colonisation alongside the classic dimensions of biogeography, it may be possible to develop a more comprehensive framework for understanding biodiversity patterns in an increasingly anthropogenic biosphere. Indeed, previous studies using chronometric resolution have highlighted that large-sized fauna extinction were more related to anthropogenic causes than to a climatic role in Australia [START_REF] Rule | The aftermath of megafaunal extinction: Ecosystem transformation in Pleistocene Australia[END_REF], Patagonia [START_REF] Villavicencio | Combination of humans, climate, and vegetation change triggered Late Quaternary megafauna extinction in the Última Esperanza region, southern Patagonia, Chile[END_REF], and in New Zealand (Holdaway et al. 2001, Trewick and[START_REF] Trewick | Vicars, tramps and assembly of the New Zealand avifauna: a review of molecular phylogenetic evidence[END_REF].

Changes in human societies

In this study, we consider the differing effects of human colonisation on species richness by three major types of human societies: hunter-gatherer, agrarian and industrial, each with profoundly different levels of societal complexity, subsistence regimes, resource use, ecological and material inheritances, and ecosystem engineering practices (Ellis 2015, Table 1). Admittedly archaeological, historical and ethnographic evidence has highlighted a strong correlation between these societies' complexities and the land productivity, the resource management, the human population size and its density, and the amount of nonhuman energy used per capita (Ellis et al;2018, Freeman et al. 2020)).

Hunter-gatherer societies, while still present today represent the earliest forms of modern human societies [START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF]. These societies generally depend on mobile to moderately sedentary social foraging strategies for subsistence (i.e. on primary and secondary productivity), which maximise the use of seasonally available local or regional food resources Typical patterns of resource exploitations are first by harvesting pressure (i.e. unsustainable hunting and overharvesting) on the most desirable large-sized fauna, then by broadening hunting (Stiner et al. 1999) and foraging strategies across taxa (niche broadening) as well as on flora with the used of wood for fuel or other parts of the plant for their antibacterial, antifungal, anti-inflammatory, antimicrobial or feeding properties (Esteban et al. 2020) and then later by the intentional use of fire to maintain more productive early-successional ecosystems [START_REF] Hamilton | The complex structure of hunter-gatherer social networks[END_REF][START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF][START_REF] Ellis | Involve social scientist in defining the Anthropocene[END_REF]; Table 1). The practice of this society has acted as a first anthropogenic and unnatural force transforming ecological pattern and process across the biosphere through the harvesting and the predation, and then by the burning practices (Braje and Erlandson 2013, Sullivan et al. 2017, Burger and Fristoe 2018), that drove to first natural failure in the ecosystems functioning. These societies have shown the first major taxonomic turnover on biodiversity through the direct effect of harvesting pressures, causing translocation, extinction, extirpation, range shifts and, through landscape modification and the use of fire, both intentionally and unintentionally [START_REF] Grayson | The archaeological record of human impacts on animal populations[END_REF][START_REF] Barnosky | Assessing the causes of Late Pleistocene extinctions on the continents[END_REF][START_REF] Koch | Late quaternary extinctions: State of the debate[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]. For example in some region of Chile ( Villavicencio et al. 2015) or in Madagascar (Burns et al 2016), the arrival of hunter-gatherer communities have led to strong decline megafauna and some case sone species extinction due to overhunting (see also Andermann et al. 2020) and the burning of human society. These societies generally mediated species introductions, through the intentional introduction of human commensals, species used as hunting aids (e.g., dogs), and the unintentional establishment of species (e.g., rats, mice; see table 1; Weisssbrod et al. 2017). This first translocation of species might also have had an important impact on behavioural ecology and evolutionary biology of endemic taxa, like in insular system (e.g. in New Zealand avifauna, Whitwell et al. 2012). More than, the subsistence and habitat-modifying behaviours of this society, depending on the presence or absence of transition with pre-hominins societies, have mediated varying effects on the natural ecosystem as well as their contribution role in biodiversity extinction. The presence of Pre-hominins societies and the dietary change observed in some of them via the consumption of animal material, through the combination of hunting and passive or active scavenging, has already started pre-anthropogenic filtering by extirpating some wildlife, due to Pre-hominins diet pressure, as observed by Werdelin and Lewis (2013) on eastern Africa carnivore guilds. The pre-hominins-environment interactions through their harvesting strategies could have already mediated ecological changes (e.g. Werdelin and Lewis 2013), which would explain the divergent impact on cascading effect on local ecosystem function from Huntergatherer society in the different ecosystem (e.g. New Zealand vs. North America vs. Europe), leading to different starting points for modern human-environment filtering.

A transition from hunter-gatherer to early agrarian societies produces even greater alterations of native biodiversity (Table 1). Indeed, the cumulative cultural evolution observed into the hunter-gatherer societies through innovations has allowed pushing back some constraints that previously limited the geographic and population expansions of these societies and now favoured the emergence of the agrarian society (Ellis et al. 2018). These culturally inherited and socially learned technologies, practices, and more productive strategies, led to an increase of human density on the order of 100 times higher than the most abundant period of hunter-gatherer society (Ellis et al. 2018, Freeman et al. 2020). With a change to Agrarian societies came a change in their associated impacts, through their intensive agricultural practices that systematically denuded vegetation from landscapes (e.g., grazing by livestock; Brigg et al. 2006) to increase the productivity of specific resources (e.g., horticultural and agricultural practices; [START_REF] Lightfoot | Rethinking the Study of Landscape Management Practices Among Hunter-Gatherers in North America[END_REF]. Agrarian practices ranged from temporary shifting cultivation that , resembled burning by hunter-gatherers, to the continuous use of land by annual cultivation and the use of irrigation, causing permanent habitat loss and fragmentation [START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]. Further impacts of agrarian societies include the introduction of domesticates, diseases, ruderals, and feral species (table 1), as well as additional pressures from larger, denser, and more rapidly growing human populations [START_REF] Grayson | The archaeological record of human impacts on animal populations[END_REF][START_REF] Koch | Late quaternary extinctions: State of the debate[END_REF][START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]. The permanent settlements associated with these societies, has also shifted the hunting pressure to a more localized area, while also increasing the rate of exotic species introductions (e.g., weeds; [START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]; Table 1), ultimately leading to a loss of biodiversity at the local scale. Thus, the change in patterns of hunting pressure has caused a widening set of pressure on local biodiversity such as on the carnivore community to avoid human-carnivore conflict (i.e. competition on wild prey, predation on domestic livestock...; Galetti et al. 2018) or by the selection and protection of 'noble game' species (e.g. deer, wild boar in Europe; Ashby 2002) for recreational hunting (Crees et al. 2019). This new human sociocultural niche construction through its different inheritance (e.g., cultural, material, ecological ...) and its increasing population density has helped the cultural evolution of the human species to outpace the rate of biological evolution, putting humans as the major ecosystems engineer of the transformation causing the terrestrial biosphere [START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF] and significant impacts on global biodiversity (see Sullivan et al. 2017). This new societal evolution represents the second anthropogenic force to transform ecological patterns and processes across the biosphere, through the intensification and the sophistication of ecosystem engineering. This has occurred via domestication and agriculture practices, by shaping new trajectories of environmental change and new ecological disturbances, and so by exerting new evolutionary pressures for biodiversity. These new transformations of ecological patterns and process, through the reduction and fragmentation of natural habitats and the development of a new anthropogenic habitat "cropland", has caused a substantial reduction of native biodiversity, which were often replaced by a fewer number of domesticated species.

Most recently, the industrial revolution and the massive food requirements of growing human populations have led to an increase of agricultural productivity for commercial purposes, by the use of toxic agrichemicals, excess nutrients, mechanisation, loss of remaining habitat fragments, increased drainage of land, intensive grassland management, and the construction of transportation networks and movement of materials and biota across the planet [START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]; Table 1). Modern agricultural techniques have therefore resulted in a further increase of land-use intensity the development of less suitable areas for agriculture [START_REF] Benton | Farmland biodiversity: is habitat heterogeneity the key?[END_REF], and the development of non-agricultural land use (i.e., urbanization; [START_REF] Ramalho | Time for a change: dynamic urban ecology[END_REF]. This new Human sociocultural niche construction sustained the third anthropogenic force that transformed ecological patterns and processes across the biosphere. The new "industrial" phase resulted from the overharvesting of nonhuman energy (e.g. fossil biomass fuels, abiotic resources of energy) to support human populations and their subsistence regimes. This drove the decline in population of previously unaffected forms of biodiversity, both locally and regionally, caused by the creation of novel selective pressures. Furthermore, the sociocultural evolution that occurred during the industrial societies also resulted in new combinations of constraints, most notably via the appearance of new philosophical concepts, such as habitat alterations for the sake of 'improvements' or 'aesthetics' of the landscape. For example, these ideas led to the creation of the acclimatisation movement (Wallace 1911, Osborne 2000) that translocated familiar biodiversity into unfamiliar environments colonised by settlers [START_REF] Carruthers | A native at home and abroad: the history, politics, ethic and aesthetics of acacias[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF]. This was done to improve the productivity of the land for agriculture, hunting, or for nostalgic reasons (Osborne 2000). Thus, the appearance of this concept of acclimatisation was characterised at a local scale by an increase in species richness but was often detrimental to native ecological inheritance and its biodiversity. This effect was much greater on island ecosystems with species extinctions occurring after the introduction of exotic species at the start of each new sociocultural stage [START_REF] Wood | Island extinctions: processes, patterns, and potential for ecosystem restoration[END_REF]. Most recently, this cultural evolution has also led to the creation of game reserves and wildlife sanctuaries to improve survival of species and, maintaining and rationalising future hunting opportunity (Jepson and Whittaker 2002).

Change in native biodiversity communities following filtering and adaptations

Throughout each of these sociocultural regimes, human-induced habitat changes, such as altered land use, introduction of exotic species and exploitation of native species (Table 1), have acted as a 'filter' that some native species will pass through or persist, and others do not [START_REF] Kraft | Community assembly, coexistence and the environmental filtering metaphor[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF], Andersmann et al. 2020), leading to associated extinctions and extirpations (Braje and Erlandson 2013). The number of surviving species after these filtering periods will be the result of three factors: 1) the numbers of species initially present (biogeographic principles), 2) the adaptive capacities of native species to cope and persist with altered environments and climate change, which will be determined by their species-specific traits [START_REF] Cornwell | A trait-based test for Habitat filtering: Convex hull volume[END_REF], and 3) the response of human society to biodiversity decline [START_REF] Welch | Indigenous burning as conservation practice: Neotropical savanna recovery amid agribusiness deforestation in Central Brazil[END_REF][START_REF] Lightfoot | Rethinking the Study of Landscape Management Practices Among Hunter-Gatherers in North America[END_REF][START_REF] Lepofsky | Indigenous Marine Resource Management on the Northwest Coast of North America[END_REF]. So, following each filtering period, co-existing species sharing a trait or combination of traits, which allowed them to cope with anthropogenic change, tended to have a higher probability of survival; a process termed 'inheritance ecology' [START_REF] Kraft | Community assembly, coexistence and the environmental filtering metaphor[END_REF]. Thus, as human societies and their associated habitat alterations change through time, species with suitable life history strategies will tend to be selected and survive (Figure 1). Consequently, the current native species richness of an ecosystem and its community structure are the product of initial regional species richness mediated by a combination of phylogenetically conserved and convergent traits that are adaptive in the face of anthropogenic pressures [START_REF] Pavoine | Measuring biodiversity to explain community assembly: a unified approach[END_REF]Bonsall 2011, Andersmann et al. 2020).

The adaptive capacity of a species in responding to dynamic anthropogenic environments depends on life history and other complex genetic traits [START_REF] Helm | Human influence lowers plant genetic diversity in communities with extinction debt[END_REF]. Natural selection for life history and other traits under earlier environmental conditions (i.e., initial human arrival), therefore, shape evolutionary responses to later environmental changes [START_REF] Sih | Evolution and behavioural response to human-induced rapid environmental change[END_REF]. Species face three main outcomes in responding to rapid environmental change: (1) survival through adaptive traits already selected through evolutionary history without further evolutionary change, (2) survival without an evolutionary history of selection for adaptive traits or phenotypic plasticity or (3) extirpation or extinction in the absence of adaptive traits or phenotypic plasticity [START_REF] Sih | Evolution and behavioural response to human-induced rapid environmental change[END_REF][START_REF] Hendry | Evolutionary principles and their practical application[END_REF]. Similar adaptive patterns have been observed in environments undergoing urbanization and are categorised as exploiter, adapter and avoider [START_REF] Blair | Land use and avian species diversity along an urban gradient[END_REF]. Thus, the evolutionary past shapes the persistence of native species in rapidly changing anthropogenic habitats [START_REF] Pavoine | Measuring biodiversity to explain community assembly: a unified approach[END_REF], Essl et al. 2015a, Figueiredo et al. 2019).

Different types of societies act as different filters for native biodiversity and species traits, acting first to determine whether species go extinct or persist, and acting then on surviving species through extended periods of selection and adaptation within anthropogenic environments (Thompson et al. 2019). As the scale of societies has increased, so has human capacity for ecosystem engineering, such that anthropogenic filtering of biodiversity has changed substantially over human history, creating a cascading effect of different ecological constraints acting to filter the biological traits of persisting native species (Essl et al. 2015a). Mensing and Colleagues (2018) research have highlighted that the modern landscape shape of central Italy resulted from the legacies from multiple political regimes of forest ecosystem management. Each political regime or society have display different management of the forest ecosystem, through notably the use and selection of specific tree taxa, that drove in the persistence across the regime to the appearance of new forest type and led as a result of the subsequent regimes to the current forest ecosystem shape (e.g. soft hardwoods were favour by Lombard's society, Oaks tree was favour by Carolingian's society for pig production, see Mensing et al. 2018) Native species richness within continents like Africa, Asia and Europe received the earliest selective pressure from behaviourally modern humans (Box 1). The earliest anthropogenic filtering processes caused the extinction of many species, in particular large-sized species that were hunted to extinction (with some exceptions like Africa), with cascading consequences across entire ecosystems [START_REF] Grayson | The archaeological record of human impacts on animal populations[END_REF][START_REF] Barnosky | Assessing the causes of Late Pleistocene extinctions on the continents[END_REF][START_REF] Barnosky | Megafauna biomass tradeoff as a driver of Quaternary and future extinctions[END_REF][START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF].

This was followed by the first adaptive responses to anthropogenic environmental changes and pressures. For example, mollusc populations of Cape turban shell Turbo sarmaticus in South Africa (Klein andSteele 2013, Sullivan et al. 2018) declined in size and altered age structure due to harvesting pressure from prehistoric humans [START_REF] Grayson | The archaeological record of human impacts on animal populations[END_REF][START_REF] Boivin | Ecological consequences of human niche construction: Examining longterm anthropogenic shaping of global species distributions[END_REF], Sullivan et al. 2017). A similar evolutionary pattern was observed in other species of molluscs in Italy (Stiner et al. 1999) as well as in north America (Erlandson et al. 2011). A comparable evolutionary response have also been reported in other terrestrial vertebrates, like the spur-thighed tortoises, Testudo graeca where humeral shaft diameter has decreased, which is a proxy for decreased body size (Stiner et al. 1999). With agrarian and industrial societies, anthropogenic filtering pressures continue to increase and expand, driving continuing species extirpation and extinction, community shifts and increasing the rates, intensity and extent of anthropogenic ecological changes. Archaeological studies in Chile (Villavicencio et al. 2015) and in Madagascar (Crowley et al. 2016) have shown evidence that the settlement and the shift from a subsistence strategy to an agrarian regime, notably with associated land-use change, accelerated the loss of biodiversity (see table in supplementary material. The amount of time elapsed between major changes in society type also moderate the severity of filtering effects, by increasing or decreasing the time available for species to adapt to dynamic anthropogenic habitats. At the time of first colonisation of native habitats by Hominids, different types of human societies present different degrees of filtering (Balée 1998, O'Connell andAllen 2015), with larger scale societies (agrarian, industrial) tending to induce more rapid rates of environmental change and more extreme filtering than smaller scale societies (e.g., hunter-gatherer; Box 1).

Native species in African and European regions experienced the longest and most gradual forms of filtering pressures by preindustrial societies. This enabled traits adaptive to dynamic anthropogenic environments to become established and lessened rates of extinction both during societal regime transitions and during the period between the transitions (Box 1) as found by previous studies (Faubry et al. 2020, Andersmann et al. 2020). In North America, hunter-gatherer societies arrived much later than in Europe and Africa, driving a rapid phase of extinction at the time of first colonisation [START_REF] Martin | The Discovery of America: The first Americans may have swept the Western Hemisphere and decimated its fauna within 1000 years[END_REF][START_REF] Martin | Prehistoric overkill: the global model[END_REF], Faubry et al. 2020, Andersmann et al. 2020). This rapid filtering effect is even more pronounced in the very recent first arrival of hunter-gatherers to New Zealand, where the and their commensal introduced species (e.g., dogs, rats) [START_REF] Barnosky | Assessing the causes of Late Pleistocene extinctions on the continents[END_REF][START_REF] Barnosky | Megafauna biomass tradeoff as a driver of Quaternary and future extinctions[END_REF], see Figure 2), where there had an absence of pre-sapiens Hominid societies.

Species exposed to shorter periods between anthropogenic filtering events (Figure 1) would be expected to experience higher risks of extinction when faced with a subsequent filtering event, owing to inadequate prior evolutionary shifts in adaptive traits (Essl et al. 2015a, 2015b, Figueiredo et al. 2019). This may explain why North America's more recent establishment of larger scale agrarian and industrial societies is related to higher extinction rates than in Europe, as native species have had less time to adapt to anthropogenic environmental changes in the Americas (Figure 2 see Box 1).

To fully understand the long-term prospects for native species' persistence and adaptation in the face of anthropogenic pressures, rigorous comparative investigations of long-term anthropogenic filtering that focuses on the 1) timing and types of societal colonisations, 2) sociocultural regime shifts (Kinzig et al. 2006), and 3) the society response to biodiversity decline [START_REF] Welch | Indigenous burning as conservation practice: Neotropical savanna recovery amid agribusiness deforestation in Central Brazil[END_REF][START_REF] Lightfoot | Rethinking the Study of Landscape Management Practices Among Hunter-Gatherers in North America[END_REF][START_REF] Lepofsky | Indigenous Marine Resource Management on the Northwest Coast of North America[END_REF], may serve as critical observational laboratories [START_REF] Cartwright | Anthropogenic natal environment of effects on life histories in a wild bird population[END_REF]. While it seems fairly well established that more recent colonisations are associated with higher rates of native biodiversity loss, the causes of this are not fully understood, nor are they necessarily generalisable to other species. Nevertheless, it is clear that failure to consider the effects of societal legacies and sociocultural transitions on native species loss and the presence of adaptations that might prevent future losses can have major consequences across history [START_REF] Ramalho | Time for a change: dynamic urban ecology[END_REF].

We propose a framework to outline the importance to understand the sociocultural legacy role in shaping current biodiversity patterns. Indeed, we argue that current biodiversity and ecosystems are the outcomes of human sociocultural filtering and transitions and not only result from ecological or climatological processes. By integrating this framework, we hope to contribute to our understanding of biodiversity evolution patterns overtimes, as well as the context in which sociocultural filtering and transitions act on species. By presenting an integrated framework for investigating the processes of native species extinction and adaptation in response to anthropogenic filtering will further advance our understanding of native species' adaptations to anthropogenic environments (see potential applications and additional considerations, Table 2). -Add new understanding to the evolution of genetic response and phenotype variation in biodiversity.

-Understand the ability of species or individuals to cope with anthropogenic habitats.

-Develop conceptual framework to explain how past human history influences the evolution cue-responses relationships between organisms and environments.

-Lack of available data on early human colonisation history -Lack of understanding of the effect of human population growth and size on biodiversity.

-Lack of knowledge about interactions of multiple stressor interactions (e.g., climate change, invasive species, and habitat clearance).

Conservation

-Probability of extinction or extirpation of native species.

-Aid in the understanding of the causes of species population declines.

-Further understanding of the factors that lead to the establishment of invasive species.

-Current rate of habitat clearance.

-Population size / propagule pressure of invasive species.

-Rate of invasive species introduction.

-Vulnerability of native species (e.g. island species).

-Lack of inclusion of interdisciplinary approaches.

Ecology -Add to biogeographical parameter to help to refine the process of -Uncertainty about the effect of climate change.

species extinction.

-Understand biodiversity dynamic and response to novel cues.

-Understand behavioural response flexibility of species to anthropogenic habitats.

-Understand the behavioural response to ecological change.

-Habitat loss and fragmentation.

-Introduction of novel enemies (e.g. diseases, predator or parasites).

-Lack of knowledge about interaction of multiple stressors.

-Problem of interpretation due to complexity of biodiversity response and individual potential adaptations.

Anthroecology

-Understand human-induced ecological pattern changes and their influence on the complexities of socioecological systems.

-Understand the dynamic interplay between human society and ecological systems.

-Aid in understanding the evolution of the anthropogenic globalwarming system and its effect on biodiversity.

-Understand the relation between human sociocultural systems transitions, landscape alteration and their long-term impact.

-Required the time depth accessible on the area on human societies.

-Lack of knowledge of major anthropogenic roles in shaping biodiversity in each societies.

-More multidisciplinary research required with ecologists and archaeologists.

469 470

Figure captions Figure 1 .

 captions1 Figure captions

Figure 2 .

 2 Figure 2. Long-term global changes based on the percentage of pre-human anthropocene large-sized fauna

  465 in relation to anthropogenic transformation and their impact on local ecosystems and biodiversity ([a] Kuneš et al. 2008, [b] Grayson 2001, [c] Huston 2005, [d] 466 Smith 2011, [e] Cartwright et al. 2014, [f] Croci et al. 2008, [g] Stockwell et al. 2003, [h] Sullivan et al.

Table 1 :

 1 Human sociocultural systems classified by primary subsistence regime, in order of historical emergence (based on[START_REF] Nolan | Human Societies: An Introduction to Macrosociology[END_REF][START_REF] Nolan | Human Societies: An Introduction to Macrosociology[END_REF][START_REF] Ellis | Ecology in an Anthropogenic Biosphere[END_REF]

Table 2 .

 2 Potential applications and additional considerations following our framework 468

		Additional consideration (limitation, alternative hypothesis,
	Research fields	Suggested applications
		other factors)
		-Understanding of how human society affects species adaptation rate.
	Evolution

Acknowledgments

We thank Andrew E. Hollis, James Dale, Dianne Brunton and the anonymous referees for their valuable comments and discussion on the paper. This project was supported by the Institute of Natural and Mathematical Sciences of Massey University.

Figure 2

Box 1. Case study

Our planet has experienced a large mass extinction event caused by human activities over millennia through human interaction (e.g., hunting, harvesting) or transformation (e.g., human niche constructing activities like cleared lands, biotic exchange, erosion). Thus this model has been used in three different contexts of selective pressure from behaviourally modern human, (1) Europe and Africa, which had the earliest selective pressures from humans (> 45,000 years before present [YBP]; Fu et al. 2014, Suppl. 1), (2) North America (~10,000 YBP; Bourgeon et al. 2017, Suppl. 1) and (3) New Zealand, which observed a later selective pressure from humans (~737 YBP; Wilmshurst et al. 2008, Suppl. 1). The ultimate goal is to identify divergence between the change of human societies and their associated habitat alterations are affiliated to reduction of native large-sized fauna richness.

In these three contexts, similar process of sociocultural niche construction have been observed, also with similar sequences of changes sociocultural systems (i.e., hunter-gatherer, agrarian, industrial societies), but with different durations and overlaps between each type of system (Figure 2.2). Thus countries more recently colonised by humans have experienced shorter transitions times between sociocultural systems that could help explain the longer temporal delays in biodiversity responses to society change. The proportion of large-sized fauna species that went extinct following colonisation of hunter gatherer societies (Figure 2.2) in Europe (0.8% biodiversity reduction) and Africa (8.8%) were more fewer than those in North America (30.6%) and New Zealand (55.9%). Humans in Europe and Africa became established over a much longer period and had less impact on local large-sized fauna extinctions, a pattern that is clearly contrasted by North America and New Zealand. However, such extinction rates cannot fully be explained by human impacts alone, as other effects such as environmental impacts, either working alone or in tandem with human impacts can contribute to extinction or extirpation events. Similar results have also been observed during the later sociocultural niche construction during the onset of industrial societies (Figure 2.2), which are more characteristic of the Anthropocene period. Indeed, Europe (13.9%) and Africa (8.8%) had a lower extinction rate during the industrial period and the Anthropocene period than North America (18.7%) and New Zealand (25.0%) where extinction was more pronounced. The time-lagged between the onsets of a news human sociocultural system and the subsequent extinction events appears to vary with the duration of human association. In locations with longer periods of human association, the extinction rates of native biodiversity was lower (see Figure 2.2 for European and Africa) following transitions between sociocultural systems, 20 likely due to longer periods for evolutionary change to occur between transitions. However, in New Zealand, one of the last locations to be colonised by humans with particularly rapid transitions between sociocultural systems, the native community of large-sized fauna has undertaken a faster magnitude of extinction. So, the transition period between human sociocultural systems combined with climatic change may favour an evolutionary responses of biodiversity to anthropogenic impacts (i.e., past experience with humans provides the evolutionary history that could shape how biodiversity responds to human impacts; [START_REF] Sih | Evolution and behavioural response to human-induced rapid environmental change[END_REF].

The importance of temporal dynamics in ecology has been well recognised. Our approach highlights the importance of temporal effect of human societies of the study of human impacts on biodiversity and landscape.

Figure 1