Ecology

Trends in cephalopods tradeSince 2000, trade in fresh octopus has been constantly dominated by the flow from China to Korea, followed by Vietnam to Japan, Portugal to Spain and Spain to Italy. However, there has been a marked decrease in the traded volume and monetary value over time, with a 50% reduction in the top 5 traders (Tables S3–S5).Over the last 20 years, fresh octopus exports have been strongly dominated by China, followed by Spain, Vietnam, Portugal, and France, and recently by Morocco and Thailand (Table S3). While Vietnam was the most important exporter in the first period (2000–2005), it was not within the top 5 traders in the last 5 years. Imports have been dominated by Korea, Italy, and Portugal, with no notable changes in the whole period (Table S4). Regarding trade of processed octopus, the largest transactions have been performed from Morocco to Spain, Morocco to Japan, Mauritania to Japan (and more recently also to Spain) and China to Korea (Table S5). Since 2000, exports of processed products have been dominated by Morocco, Mauritania, China, Spain, and Vietnam (Table S3), while imports have been led by Japan, Spain, Italy, Korea, and the United States (Table S4).Trade in volume of fresh cuttlefish and squid includes fewer clear relationships over time, such as transactions from Malaysia to Singapore (2000–2004 and 2005–2009); from Myanmar to Thailand (2005–2009 and 2015–2019); and from Yemen to Vietnam (2010–2015) (Tables S6-S8). For the first 5 years, exports of fresh commodities were dominated by Vietnam. However, since 2005, India, Spain and France have increased their exports in both monetary value and volume, displacing Vietnam from the top rank (Table S6). The main importing traders were Spain and Italy. Although China was important in the first decade, it was replaced by Vietnam in the last decade (Table S7). The trade of elaborated cuttlefish and squid products was dominated by monetary value flow from Thailand to Japan and from Malvinas/Falkland Islands to Spain in the first decade, while in the last decade the flow from China to other traders (Japan, the USA, and Thailand) gained relevance. However, the volume follows a different pattern, with flow from the Malvinas/Falkland Islands to Spain and from Korea to China in the first decade, while in the last decade, flows from Peru to China and from China to Thailand were important (Table S8). A disparity exists between the top five traders in terms of flow of monetary value and volume in the first 15 years; but in the last 5 years the top positions are constantly represented by, China, Peru, India, and Spain. Italy, Japan, China, and the USA are important importers in terms of monetary value and volume, although in the last decade Thailand has increased its importance, replacing the USA in the top 5 in the last 5 years (Table S7).The CGTN involved 220 traders (countries or territories) from around the world with exports greater than or equal to 500 kg between 2000 and 2019 (Fig. 3). The remaining 32 traders either did not report exports or their exports were below 500 kg. The most important cluster of traders was composed by 8 countries that dominate the cephalopod global markets in Asia (China, India, Republic of Korea, Thailand, Vietnam), Europe (the Netherlands, Spain) and the USA. The second and third most relevant clusters were composed of 8 and 12 traders, respectively. These two clusters involve 9 developed countries (Belgium, Canada, Denmark, France, Germany, Italy, Japan, United Kingdom and Portugal) and 11 developing countries (Morocco, Malaysia, UAE, Senegal, South Africa, Peru, Indonesia, Philippines, Argentina, Chile and New Zealand). Some of these traders have the most productive cephalopod fisheries in the world (e.g., Patagonian shortfin squid in the Southwest Atlantic Ocean and Patagonian squid in the Southeastern Pacific Ocean).Figure 3The Cephalopod Global Trade Network. The top 220 traders of the CGTN as nodes (circles) and their trade links as lines. The colour and the size of the nodes represent, respectively, the cluster membership and relative importance of the trader in the CGTN, estimated from the number of trade links with other traders (i.e., degree). The colour of the edges represents the origin, destination and the proportion of trade links for all years between each pair of traders. The clusters were made using Ward’s method. The figure was created with R12 (https://cran.r-project.org) packages: “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com) and “ggtree” v3.0.232 (https://guangchuangyu.github.io/ggtree-book/chapter-ggtree.html).Full size imageOctopus trade networkLive, fresh, or chilled octopusThe normalised strength (Fig. 4) revealed the importance of China and Republic of Korea in the trade of fresh octopus in monetary value, with high importance of flows between these two traders over time (Supplementary Fig. S1). Other relevant traders over time were Spain, Portugal and Italy, in Europe; and Vietnam and Japan, in Asia from 2000 to 2004 (Supplementary Fig. S1). The network based in volume showed similar results.Figure 4Global trade network for octopus live, fresh or chilled between 1 January 2000, and 31 December 2019 in monetary value (USD). The numbers correspond to the normalised strength for the monetary value. Each node represents a trader, and each edge represents the export–import relationship between two traders. The size and colour of the node represent the relative importance of the trader in the network in terms of its strength. The width and colour of the edge represent the relative importance of the relationship between two traders in terms of their edge strength. The figure was created with R12 (https://cran.r-project.org) packages: “ggplot2” v.3.2.113 (https://ggplot2.tidyverse.org), “ggmap” v.3.0.029 (https://github.com/dkahle/ggmap) and “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com).Full size imageThe Betweenness identified important actors facilitating flow through the network. For fresh octopus, the most relevant traders in the last two decades were Spain, France, and Italy, followed by Thailand, Portugal and the USA. Again, no major differences exist between the monetary value and volume networks. However, the ranking of traders changed over time, with Italy replacing Spain in the first place during the period 2005–2009, and then Spain consolidating again in the first place in the following periods. Also, in the last 20 years there were many changes in Asia, with Vietnam losing and Korea gaining prominence over time (Supplementary Fig. S2).In a global trade network, there are countries that are essential to the network structure because they are connected to other countries critical to the network and those critical countries, in turn, have no other significant connections. PageRank is a centrality measure that identifies these important countries, resulting from an iterative algorithm that assigns higher values to countries with a greater number of import connections with other countries that move large quantities of goods or money33. In the last 20 years, Italy, Germany France and Spain have occupied those central positions in the global trade network of live, fresh or chilled octopus. Their dominance has not changed over the four 5-year periods analysed (Fig. 5).Figure 5Global trade network for octopus live, fresh or chilled between 1 January 2000, and 31 December 2019 in monetary value (USD). The numbers correspond to the normalised PageRank for the monetary value. Each node represents a trader. The size and colour of the node represent the relative importance of the trader in the network in terms of its PageRank. The figure was created with R12 (https://cran.r-project.org) packages: “ggplot2” v.3.2.113 (https://ggplot2.tidyverse.org), “ggmap” v.3.0.029 (https://github.com/dkahle/ggmap) and “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com).Full size imageElaborated octopusThe normalised strength revealed a diversified trade network for elaborated octopus products. This network has remained relatively stable over the last 20 years, with some exceptions. In the 2000s, there was an intense flow of exports from North Africa (i.e., Morocco and Mauritania) to Japan, although the most important flow was from Morocco to Spain. In the 2010s, Mauritania changed its preferential partner and exported large quantities of elaborated octopus to Spain, with the latter gaining dominance. In this last decade, flows from China and Vietnam to the Republic of Korea also became important. Other relevant actors were distributed globally (e.g., Italy, Portugal, Senegal, the USA). However, the most important routes showed a common pattern: the origin was in developing countries or territories (that emerged as producers) while developed countries showed a high and stable consumer demand (Supplementary Fig. S3). The network based on volume was highly similar to the monetary value network. However, Italy, China, Korea, Vietnam, and the USA reduced their importance compared to the top-ranked traders (i.e., Spain, Japan, and Morocco). The most important routes of the volume network were from China to Korea; Morocco to Spain; Morocco and Mauritania to Japan and Vietnam to Korea.The Betweenness measure highlighted the role of Spain as a facilitating actor in the trade network of elaborated octopus, followed by Italy, China, and the USA. These countries have maintained their importance as structurers of the world trade network in elaborated octopus over the past 20 years. Similarly, the routes from Italy to Spain, and from Spain to China and the USA emerged as relevant in the network structure, with a special mention to the route between Japan and China in the last 5-year period (Supplementary Fig. S4). There are no major differences between the most central traders in this network and the volume-based one.PageRank revealed the importance of Spain and Italy as leading traders in the elaborated octopus market. These two countries concentrated a large number of import relationships, which also concentrated a large monetary and volume flow. This importance has been maintained over time. Other reference actors were Greece, Japan, the USA and Portugal, in the first decade; and the USA, Portugal, Greece and Korea in the second decade. Note how in the second decade, Greece and Japan lost relevance, while Northern European countries and the Republic of Korea gained relevance (Supplementary Fig. S5).Squid and cuttlefish trade networkLive, fresh, or chilled squid and cuttlefishThe normalised strength revealed the importance of Spain, France, Italy, and India in the trade network of fresh squid and cuttlefish products, especially the route between east Asia and Spain (Fig. 6). The volume-based network is highly similar to the monetary value network. Over the four 5-year periods analysed, Vietnam and Japan have gradually lost relative importance in the network (Supplementary Fig. S6).Figure 6Global trade network for squid and cuttlefish live, fresh or chilled between 1 January 2000, and 31 December 2019 in monetary value (USD). The numbers correspond to the normalised strength for the monetary value. Each node represents a trader, and each edge represents the export–import relationship between two traders. The size and colour of the node represent the relative importance of the trader in the network in terms of its strength. The width and colour of the edge represent the relative importance of the relationship between two traders in terms of their edge strength. The figure was created with R12 (https://cran.r-project.org) packages: “ggplot2” v.3.2.113 (https://ggplot2.tidyverse.org), “ggmap” v.3.0.029 (https://github.com/dkahle/ggmap) and “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com).Full size imageFor fresh squid and cuttlefish, Betweenness identified Spain as the most important structurer of the network, in both monetary value and volume, over the time. Europe emerged as a major region structuring the global trade network. While in Asia, the exchange of countries with higher betweenness over time evidences the strong struggle for control of trade in the region and the more fragile sub-networks. In the monetary network, in the 2000s the bridge between the United Kingdom and Korea stood out, while in the 2010s the Europe-Asia bridge was established between Spain and India. In the volume-based network, it is noteworthy that in the 2010s there were many critical routes for the stability of the network, even in the Europe–Asia connection. Note, for example how the link between Netherlands and Myanmar stand out (Supplementary Fig. S7).PageRank revealed the importance of Italy, Spain, Germany and France as leading traders in the fresh squid and cuttlefish market. In the last 20 years, Europe concentrated the largest number of import relationships, which also concentrated a large monetary and volume flow. Europe leadership has been maintained over time (Fig. 7).Figure 7Global trade network for squid and cuttlefish live, fresh or chilled between 1 January 2000, and 31 December 2019 in monetary value (USD). The numbers correspond to the normalised PageRank for the monetary value. Each node represents a trader. The size and colour of the node represent the relative importance of the trader in the network in terms of its PageRank. The figure was created with R12 (https://cran.r-project.org) packages: “ggplot2” v.3.2.113 (https://ggplot2.tidyverse.org), “ggmap” v.3.0.029 (https://github.com/dkahle/ggmap) and “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com).Full size imageElaborated squid and cuttlefishThe trade networks based on monetary value and volume for elaborated squid and cuttlefish emerge as global and complex, where several far distant traders have relevant roles in the import/export network (Supplementary Fig. S8). Although the most important nodes in the volume-based network reflected important nodes in the monetary value network, the strengths of the links, i.e., the flow of value and volume, did not. For example, in the volume-based network, Peru exported the largest quantities of squid and cuttlefish to China (Supplementary Fig. S8b,d), but the flow of money for these transactions was less important (Supplementary Fig. S8a,c).The betweenness centrality metric (based both on monetary value and volume) showed the importance of China, the USA, and Spain (followed by Italy, Korea, and Thailand) as facilitators in the elaborated goods trade network (Supplementary Fig. S9). While the main important bridges in volume transactions were between Italy and Spain, Spain and China, and China and the USA (Supplementary Fig. S9b,d), the main monetary bridges were from the USA to China, followed by the routes from Spain to the USA and from Italy to Spain (Supplementary Fig. S9a,c). The key traders structuring the network were the same, but they follow different directions.Closeness centrality highlighted the main actors in a regional context (Fig. 8). In both the monetary value and volume-based networks, China, North and South Korea, India, Indonesia, Thailand, and Vietnam form a strong trade network for squid and cuttlefish elaborated in Asia. Key players include South America (Peru, Argentina, Chile, the Malvinas/Falkland Islands); the USA; the Mediterranean (Morocco, Spain); Africa (South Africa, Mauritania); and the West Pacific region (New Zealand, Japan). Note how the highest values of closeness were slightly different in the money-based network (Fig. 8a) and in the volume-based network (Fig. 8b), mainly for those countries that are historically large producers of elaborated squid and cuttlefish (e.g., Peru and Argentina).Figure 8Global trade network for squid and cuttlefish elaborated between 1 January 2000, and 31 December 2019 in monetary value (USD) above, and volume (kg) below. The numbers correspond to the normalised closeness for the monetary value (USD) and volume (kg) traded, respectively. Each node represents a trader. The size and colour of the node represent the relative importance of the trader in the network in terms of its closeness. The figure was created with R12 (https://cran.r-project.org) packages: “ggplot2” v.3.2.113 (https://ggplot2.tidyverse.org), “ggmap” v.3.0.029 (https://github.com/dkahle/ggmap) and “ggraph” v.2.0.030 (https://ggraph.data-imaginist.com).Full size imagePageRank revealed the importance of Greece, Spain, China, USA and Japan as leading traders in the global elaborated squid and cuttlefish market for the first 10-years (Supplementary Fig. S10a). Globally, in the last decade, China, the USA and Japan have declined in importance, while Europe has consolidated its importance. However, Spain’s relevance has declined over the last decade while Germany’s has increased (Supplementary Fig. S10b). More

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OBITUARY
10 January 2022

Edward O. Wilson (1929–2021)

Naturalist, conservationist and synthesizer who founded sociobiology.

Bert Hölldobler

0

Bert Hölldobler

Bert Hölldobler holds the Robert A. Johnson Chair in Social Insect Research and is Regent’s Professor in the School of Life Sciences at Arizona State University, Tempe. He began working with Wilson in 1970.

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Harvard University Professor E.O. Wilson in his office at Harvard University in Cambridge, MA. USACredit: Rick Friedman/Corbis via Getty

Edward (Ed) Wilson began by exploring the systematics, geographical distribution, social organization and evolution of ants. He became one of the great scholarly synthesizers, winning two Pulitzer prizes. A superb naturalist who enjoyed challenging dogma, he fought for conservation, brought ideas of biodiversity into the mainstream and set ecology on a rigorous conceptual footing. He has died aged 92.Wilson’s book Sociobiology, published in 1975, was the first to address the evolution and organization of societies in organisms ranging from colonial bacteria to primates, including humans. The final chapter, on human social interaction, ignited controversy. Wilson argued that human behaviour, although adaptable to environmental conditions, is rooted in a genetic ‘blueprint’. Opponents claimed that nothing in human behaviour is grounded in genetics, except sleeping, eating and defecation. In a letter to The New York Review of Books, a group of academics including evolutionary biologists Stephen Jay Gould and Richard Lewontin associated Wilson’s view with racism and genocide. Wilson responded with elegance and humour; in my view, most scholars now agree that he won this argument.
Conservation: Glass half full
Wilson was born in 1929 in Birmingham, Alabama, and grew up, as he admitted in his 2006 autobiography, Naturalist, “mostly insulated from its social problems”. After studying biology at the University of Alabama in Tuscaloosa, he did graduate studies at Harvard University in Cambridge, Massachusetts. He felt its Museum of Comparative Zoology, with the world’s largest ant collection, was his “destiny”.In 1955, he obtained his PhD on the systematics of the ant genus Lasius, which includes the widespread black garden ant. Systematic biology and the study of biodiversity remained his mission, but he made significant contributions to other fields, such as animal behaviour and chemical ecology. His early work on chemical communication in animals, particularly social insects, inspired a generation of scientists to explore a new area in behavioural physiology.In 1954, Wilson set out for Melanesia, including New Guinea, to study ant taxonomy and biogeography. On the basis of his data, he elaborated the critique that he and his Harvard colleague William Brown had previously developed on the idea of subspecies. They argued that the distinctions between species should be more clearly defined, allowing for variability within species. Equally influential was their thinking on character displacement — when similar species in the same area diverge genetically to avoid competing for resources.Through his fieldwork in Melanesia and later in the Caribbean, Wilson drafted a principle of biogeography that he called the taxon cycle. Species evolve back and forth between being able to live in marginal habitats, and thus disperse widely, and restricting their distribution to species-rich habitats in island interiors. He tested this and other original hypotheses in the Florida Keys in the 1960s, in collaboration with his former student Daniel Simberloff. With ecologist Robert MacArthur, he proposed that species maintain their populations through trade-offs between number of offspring and quality of parental care (the concept of r/K selection). Their 1967 book The Theory of Island Biogeography had far-reaching effects on studies of evolution and conservation.
A revolution in evolution
From early in his career, Wilson wondered about ways to understand the evolution of social organization, from primates to social insects (such as honeybees and ants). “A congenital synthesizer,” he wrote in his autobiography, “I held on to the dream of a unifying theory.” He developed a theory of adaptive demography — that certain kinds of social structure might increase reproductive fitness — and the evolution of division of labour between castes, such as insect queens and worker groups. First brought together in The Insect Societies (1971), these concepts were elaborated in Caste and Ecology in the Social Insects, with mathematical biologist George Oster, in 1978.Sociobiology was a much more far-reaching synthesis on the evolution of social systems. The furore that ensued stimulated Wilson to write an even more provocative book, On Human Nature (1978). This garnered his first Pulitzer. His highly original book Biophilia (1984) was the first to use the term to mean human empathy for the natural world. He argued that pleasure in being surrounded by diverse living organisms is a biological adaptation. These books prepared the ground for Consilience (1998), which one reviewer called a biologist’s dream of the unity of knowledge. It proposed the kind of intellectual annexation that occurs when one field can be explained in terms of a more fundamental discipline, and received a mixed response.To his and my utmost surprise, in 1990, the huge monograph The Ants, on which we worked for years, won another Pulitzer. Wilson continued to publish on human evolution and humanity’s relationship with the planet into his 90s. Half-Earth (2016) is a passionate plea to leave half of our world to nature.Ed was not a team builder. He preferred to work alone, although in a few cases he found colleagues who complemented his abilities. He thrived on controversy. In the past two decades, he had rejected the theory of inclusive fitness — the idea that the reproductive success of an individual increases when it helps to raise the offspring of its close relatives — that he once propagated. This led to heated debates, and I opposed some of his views. When we reached a compromise and submitted the manuscript of our book The Superorganism (2009), Ed’s concluding remark was: “Bert, there is one thing we agree on 100%. That is: my co-author is wrong.” One could disagree with Ed over scientific issues and remain good friends.

Nature 601 (2022)
doi: https://doi.org/10.1038/d41586-022-00078-7

Competing Interests
The author declares no competing interests.

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Study areaAll simulations were run at a 100 × 100 m resolution for the entire European Alps, which cover ~200,000 km². Elevations reach 4,810 m above sea level at the highest peak (Mont Blanc, elevational data were obtained from ref. 44). Mean annual temperature ranges from approximately −13 up to 16 °C and annual precipitation sums reach up to ~3,600 mm (climatic conditions were obtained from WorldClim45).Species dataTrue presences/absences were derived from complete species lists of 14,040 localized plots covering subalpine and alpine non-forest vegetation of the Alps, compiled from published46 and unpublished data sources. For more information see the supplementary information in ref. 21.Data on demographic rates as well as dispersal parameters were taken from ref. 21, Supplementary Table 1. Detailed values are provided in Supplementary Table 1.Environmental variablesCurrent climate dataMaps of current climatic conditions were generated on the basis of mean, minimum and maximum monthly temperature obtained from WorldClim45 and monthly precipitation sums derived from ref. 47 at a resolutions of 0.5 arcmin and 5 km, respectively. Temperature and precipitation data were downscaled to 100 m as described in ref. 21 and using ordinary kriging with elevation as covariable. As the reference periods of these two datasets do not match (temperature values represent averages for 1950–2000, while precipitation covers 1970–2005) temperature values were adapted using the E-OBS climate grids available online (www.ecad.eu/download/ensembles/download.php). We used these spatially refined temperature and precipitation grids to derive maps of mean annual temperature and mean annual precipitation sum. We chose only two climatic variables to keep models simple and, therefore, interpretation of results more straightforward. In fact, the climatic drivers of population performance and distribution can be more complex48 and vary among species, life history stages and vital rates49. However, as correlations between different descriptors of temperature (such as coldest month or warmest month temperature, Pearson correlation of 0.84) as well as between precipitation variables are high in the European Alps, we chose mean annual temperature and mean annual precipitation sum as they give the most basic description of how climatic conditions change over geographical and elevational gradients.Future climate dataMonthly time series of mean temperature as well as precipitation sums predicted for the twenty-first century were extracted from the Cordex data portal (http://cordex.org). We chose two IPCC5 scenarios from the RCP families representing mild (RCP 2.6) and severe (RCP 8.5) climate change to consider the uncertainty in the future climate predictions. Both scenarios were generated by Météo-France/Centre National de Recherches Météorologiques using the CNRM-ALADIN53 model, fed by output from the global circulation model CNRM-CM5 (ref. 50). The RCP 2.6 scenario assumes that radiative forcing reaches nearly 3 W m−2 (equal to 490 ppm CO2 equivalent) mid-century and will decrease to 2.6 W m−2 by 2100. In the RCP 8.5 scenario, radiative forcing continues to rise throughout the twenty-first century and reaches >8.5 W m−2 (equal to 1,370 ppm CO2 equivalent) in 210024.These time series were statistically downscaled (delta method) by (1) calculating differences (deltas) between monthly temperature and precipitation values hindcasted for the current climatic conditions (mean 1970–2005) and forecasted future values at the original spatial resolution of 11′; (2) spatially interpolating these differences to a resolution of 100 × 100 m using cubic splines and (3) adding them to the downscaled current climate data separately for each climatic variable21,36. Subsequently, we calculated running means (averaged over 35 years) for every tenth year (2030, 2040 through to 2080) for each climatic variable and finally derived, on the basis of the monthly data, mean annual temperature and mean annual precipitation sums for these decadal time steps. The application of 35-yr running means ensures that we use the same time interval for parameterization and prediction. Furthermore, for long-lived species such as alpine plants, running means over long time intervals appear most appropriate to characterize climatic niches33.Soil dataIn addition to the climatic data we used a map of the percentage of calcareous substrate within a cell (5′ longitude × 3′ latitude dissolved to 100 m resolution; further referred to as soil) as described in the supplementary information of ref. 21.Environmental suitability modellingWe parameterized logistic regression models (LRMs) with a logit link function using species presence/absence data as response and the three environmental (two bioclimatic and one soil) variables as predictors. All predictor variables entered the model as second-order polynomials in agreement with the standard unimodal niche concept.From the models, we also derived a threshold value to use for translating predicted probability of occurrence (as a measure of site suitability) into predicted presence or absence of each species at a site (called occurrence threshold, OT, henceforth). The threshold was defined such that it optimized the true skills statistic (TSS), a measure of predictive accuracy derived from comparing observed and predicted presence–absence maps51.Genetic model and niche partitioningSpecies-specific suitability curves for the annual mean temperature gradient derived from the LRMs were partitioned into suitability curves of ecologically different haplotypes of a species as defined by allelic variation in up to three diploid loci (Extended Data Fig. 6). The number of alleles was varied (n = 1, 2, 3, 5 and 10 alleles) as was the proportion of the entire species’ (temperature) niche covered by each haplotype. Models with more than one locus were run with diallelic loci, as otherwise computational efforts would have increased excessively (for each haplotype the number of seeds, juveniles and adults has to be stored and all seeds have to be distributed separately). Each combination of haplotype number and allelic niche size used in a particular simulation is further referred to as setting. Species-specific suitability curves along the other two dimensions (precipitation and soil) remained unpartitioned to ease interpretation of results. The implications of relaxing this assumption by modelling niche partitioning along different environmental gradients are hard to predict. Outcomes would probably depend on the direction and strength of individual specialization along these gradients, whether they are positively or negatively correlated, as well as on how both temperature and precipitation patterns will be affected by climate change. As a consequence, the patterns we found could be re-enforced, for example when the replacement of cold-adapted haplotypes within the species elevational range is further delayed, for example, because those haplotypes adapted to warmer conditions can cope less well with higher precipitation at higher elevations. Vice versa, maladaptation to the warming temperatures might be attenuated, for example, if temperature increase is paralleled by precipitation decrease and emerging drought stress. If, in this case, haplotypes from lower elevations can better cope with both higher temperatures and less water availability than those of median elevations, they may replace the latter faster at these median elevations and hence shorten the phase of maladaptation.Allelic effects were assumed to define the temperature optimum additively. Hence, the heterozygotes’ optimum is always exactly between the optima of the two corresponding homozygotes, corresponding to a codominant genetic model. Finally, all haplotypes corresponding to one setting were assumed to have constant (temperature) niche size, defined as a proportion (ω = 50%, 75% and 100%, for one haplotype only 100%) of the entire species’ (temperature) niche. The temperature niche was computed as the difference between the upper and lower temperature values at which the LRM-derived suitability curve predicted a suitability equal to OT (with precipitation and soil set to the respective optima of the species, also derived from the LRMs). To derive the same geographic distribution under current climatic conditions for each setting, the union of the niches of all haplotypes of one set has to approximate the niche of the single-species model (Extended Data Fig. 6). Note, however, that, the aspired equality of niches is impossible, as the niches resulting from a logistic regression with quadratic terms are always elliptic in shape. Therefore, the optima of the two extreme homozygotes (that is, those carrying haplotypes adapted to the coldest or warmest margins of the entire species’ niche) are fixed at: niche limits ± 1/2 × ω × niche size and all other optima are distributed between them at equal distances (Extended Data Fig. 6 gives a schematic illustration). As a consequence, the performance of the extreme haplotypes, both coldest and warmest, is modelled to be somewhat higher towards the cold and warm margins of the temperature niche, respectively, compared with the performance modelled for the species without intraspecific niche partitioning (compare the overlap of the black with the red and blue curves in Extended Data Fig. 6a). However, as haplotype number did not affect modelled range loss (compare Table 1), this marginal mismatch does not apparently impact our results. Homozygotes were ordered from the cold-adapted A1A1 up to the warm-adapted AnAn.Finally, the suitability curves of the genotypes were assumed to have the same value at their optimum as the species-specific suitability curve at this point (Extended Data Fig. 6).Artificial landscapesArtificial landscapes were defined as a raster of 50 × 112 cells (of 100 × 100 m). These rasters were homogeneous with respect to precipitation and soil carbon conditions which were set to the values optimal for each species according to the LRMs. With respect to temperature, by contrast, we implemented a gradient across the raster running from the minimum –9.1 °C to the maximum +3.8 °C temperature for which the LRM predicts values >OT across all six species. Buffering by 1 °C at both limits was done to avoid truncating simulation results. Further 4 °C at the lower limit were added to consider simulated temperature increase (below). The final temperature range represented by the raster ran from −14.1 to +4.8 °C. Temperature increased linearly along this axis by an increment of 0.171 °C per cell, derived from a 20° slope and a temperature decrease of 0.5 °C per 100 m of elevational change. Along the 50-cell breadth of the landscape, temperature values were kept constant. On the basis of these grids, we implemented a moderate and a severe climate change scenario, characterized by temperature increases of 2 and 4 °C, respectively, over 80 yr. Therefore, temperature of each raster cell increased annually by 0.025 and 0.05 °C, respectively.Simulations of spatiotemporal range dynamicsCATS21 is a spatially and temporarily explicit model operating on a two-dimensional grid (of 100 m mesh size in this case). CATS combines simulations of local species’ demography with species’ distribution models by scaling demographic rates relative to occurrence probabilities (suitabilities) predicted for the respective site or grid cell by the latter. Dispersal among grid cells is modelled as a combination of wind, exozoochoric and endozoochoric (that is, animal dispersal via attachment to the fur or ingestion and defecation, respectively) dispersal of plant seeds. Time proceeds in annual steps. The annually changing occurrence probabilities at each site affect the demography of local populations and hence, eventually, the number of seeds that are produced in each grid cell in the respective year. As a consequence, local populations grow or decrease, become extinct or establish anew and hence the species’ distribution across the whole study area changes as a function of the changing climate.Demographic modellingClimate-dependence of local demography was modelled by linking demographic rates (seed persistence, germination, survival, flowering frequency, seed yield and clonal reproduction) and carrying capacity to occurrence probabilities predicted by LRMs by means of sigmoidal functions. Furthermore, all rates were fixed at OT at a value ensuring stable population sizes; for more information see refs. 21,33. Demographic rates were confined by zero and a species-specific maximum value (Supplementary Table 1), which was assumed to be the same for all genotypes of a species. As a corollary, the demographic rates of all genotypes of a species are the same under optimal environmental conditions but their actual values for a particular site in a particular year differ due to different temperature optima of genotypes. In addition, germination, survival and clonal reproduction were modelled as density-dependent processes to account for intraspecific competition between genotypes. In our application, for all density-dependent functions, the species abundance is defined as the sum of all adult individuals in a given cell, regardless of their genotypes. Density dependence is commonly achieved by multiplying rates with (C – N)/C, where N is the species abundance and C is the (site- and genotype-specific) carrying capacity. This correction for density dependence causes the functions to drop to zero when N approximates C. To avoid the subsequent collapse of population sizes, we defined density-dependent rates as (C – N)/C × rate() + N/C × rate(OT), which ensures stable population sizes at densely populated sites occupied by only one genotype. To account for uncertainty in parameters of demographic rates, we assigned each species two value sets representing the upper and lower end of a plausible range of values on the basis of information derived from databases and own measurements (Supplementary Table 1).The simulations allowed for cross-pollination between genotypes. We used the relative amount of flowers (genotype-specific flowering frequency as defined by the sigmoid curve for the given suitability in the given raster cell for the given year × number of adults of that genotype in the population of that cell) to derive an estimate of the haplotype frequencies in the total pollen produced by the population within a grid cell. For the multiallelic case we allowed for recombination between loci with a recombination rate of 0.1%. These frequencies were set equal to the probability that particular haplotypes are transmitted to each year’s seed yield by pollination. Spatial pollen dispersal was accounted for in the following way: in each cell, 5% of the pollen involved in producing the annual seed yield, was assumed to stem from outside the respective raster cell. The proportions of different haplotypes in this 5% were derived from the overall pollen frequencies in all cells within a 700 m radius around the target cell (following estimates in ref. 52). Subsequently, produced seeds of each genotype were divided into resulting genotypes regarding the adult’s haplotype composition and the haplotype frequencies in the cells’ entire pollen load.Dispersal modellingFor wind dispersal of plant species we parameterized the analytical WALD kernel53 on the basis of measured seed traits and wind speed data from a meteorological station in the Central Alps of Austria. Exozoochorous and endozoochorous plant kernels were parameterized on the basis of correlated random walk simulations for the most frequent mammal dispersal vector in the study area, the chamois (Rupicapra rupicapra L.). For more details, see ref. 33. To account for uncertainties in species-specific dispersal rates, the proportion of seeds dispersed by the more far-reaching zoochorous kernels was assumed either as high (1–5%) or low (0.1–0.5%), setting upper and lower boundaries of a credible range of the dispersal ability of species.Simulation set up and simulation initializationTo test for the effects of climate change on genetic diversity in 2080, we ran CATS over the period 2000 to 2080 for each of the six study species across the entire Alps under a full factorial combination of (1) three niche sizes (50%, 75% and 100%); (2) six numbers of haplotypes (equal to two, three, five and ten alleles for one locus and four and eight for the diallelic two- and three-locus models, respectively); (3) three climate scenarios (current climate, RCP 2.6 and RCP 8.5); and (4) two sets of demographic and dispersal parameters. As a ‘control’ we also ran simulations for all climate scenarios and the two demographic and dispersal parameter sets for a setting with one genotype filling the whole niche of the species. To account for stochastic elements in CATS four replications were run for each combination of ‘treatments’.For simulations in artificial landscapes we used, instead of RCP 2.6 and RCP 8.5, ‘artificial’ climatic scenarios with an assumed warming of 2 and 4 °C, respectively, and no change in precipitation.All simulation runs were started with homozygotic individuals only. As initial distribution, for each simulation run each cell predicted to be environmentally suitable to the species (that is, occurrence probability of species >OT)—and within the real distribution range of the species28 (not relevant for simulations in artificial landscapes, of course)—was assumed to be occupied by an equal number of adults of each (homozygotic) genotype, with the total sum equal to the carrying capacity of the site. To accommodate this arbitrary within-cell genetic mixture of homozygotes (and missing heterozygotes) to actual local conditions we started simulations of range dynamics with a burn-in phase of 200 yr, run under constant current climatic conditions. To have a smooth transition from the burn-in phase under current climate (corresponding to the climate of the years 1970–2005; see current climate data) to future climate projections (starting with 2030) and to derive annual climate series, climate data were linearly interpolated between these two time intervals.Statistical analysisWe evaluated the contribution of climate scenario, haplotype number and haplotype niche size to overall species’ range change as well as the change in the frequency of the warm-adapted haplotype by means of linear models. In these models, log(range size in 2080/range size in 2000) and log(percentage of warm-adapted haplotype in 2080/percentage of warm-adapted haplotype in 2000), averaged over the four replicates and the two demographic and dispersal parameter sets, were the response variables. For the analysis of the change of the warm-adapted haplotype simulation settings with 100% niche size were ignored, as in this setting all haplotypes have the same temperature optimum (that is, neutral genetic variation). Approximate normality of residuals was confirmed by visual inspection.As indicator of the ‘topographic opportunity’ remaining to the species in the real world we calculated the area colonizable at elevations higher than those already occupied at the end of the simulation period. We therefore drew a buffer of 1 km around each cell predicted to be occupied in 2080 and then summed the area within these buffers at a higher elevation than the focal, occupied cell. Overlapping buffer areas were only counted once. This calculation was done for simulations conducted with one full-niche genotype per species only.Sensitivity analysisWe interpret the simulated relative decrease of warm-adapted haplotypes mainly as an effect of (1) the unrestricted expansion of cold-adapted haplotypes at the leading edge and (2) the resistance of the locally predominating haplotype that becomes increasingly maladapted with progressive climate warming, to recruitment by better-adapted haplotypes from below that are either rare or not present at all initially. However, the results achieved, and our conclusions, may be sensitive to assumptions about particular parameter values. Parameters that control the longevity of adult plants, and indirectly the rate of recruitment of new individuals, as well as those controlling gene flow via pollen (instead of seeds) may be particularly influential in this respect. We additionally ran simulations on artificial landscapes under alternative values of these parameters. In particular, we set the maximum age of plants to 10 yr instead of 100 yr and raised the proportion of locally produced pollen assumed to be transported up to 700 m to 10%. Both of these values are thus probably set to rather extreme levels: a maximum age of 10 yr is much shorter than the 30–50 yr assumed to be the standard age of (non-clonal) alpine plants31; and a cross-pollination rate between cells of 10% is high given that among the most important alpine pollinators only bumblebees are assumed to transport pollen >100 m regularly54,55. We add that we ran these additional simulations only in combination with the demographic species parameters set to high values because a short life span combined with low-level demographic parameters did not allow for stable populations of some species, even under current climatic conditions.For individual species, adapting plant age and cross-pollination rate between cells (Extended Data Fig. 7), did change the magnitude of loss of the warm-adapted haplotype. Nevertheless, for all of them the warm-adapted haplotype still became rarer when climate warmed and this effect increased with the level of warming. We are confident that our conclusions are qualitatively insensitive to variation of these parameters within a realistic range.Finally, in the simulations where we assumed a multilocus structure of the temperature niche, the recombination rate may also affect simulation results because it determines the rate by which new haplotypes can emerge. We also tested sensitivity of our simulations to doubling the recombination rate to 0.2%. Again, we found that a higher recombination rate had little qualitative effect on the results. Quantitatively, it resulted in a slightly pronounced relative decrease of the warmth-adapted haplotype in most species (Extended Data Fig. 8).Reporting SummaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More