Philippa Kaur

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This is the first study to evaluate the mercury content in the fur of Siberian tigers in the Far East of Russia. It is a commonly recognized fact that fish are the main source of mercury entering the organism of predators and the trophic network of the ecosystem. In some areas, the seasonal abundance of salmonids can provide tigers with protein: masu (Oncorhynchus masou) from April to early July, chum (O. keta) in late autumn and up to December and pink salmon (O. gorbuscha) in July–early October. However, our observations during 1976–2018 (15 (Poddubnaya, unpublished data)) and data on mercury content show that tigers do not eat salmon often. Tigers do not hunt the redfin dace (Tribolodon hakonensis) a cyprinid fish, moving up in huge swarms from April to June.Although tigers were never observed to consume fish in mass quantities, as is the case with bears, one would expect that the total concentration of mercury (THg) in the body of tigers from two sections of the Sikhote-Alin (basin drainage of the Sea of Japan and the catchment of the Amur River) would vary depending on the availability of anadromous fish. The rivers of the Sea of Japan basin are shorter and shallower than Amur’s tributaries. Therefore, the likelihood of a tiger catching fish here seems to be higher. However, Amur’s tributaries are richer in fish and the probability of catching fish has to be no less than that observed in the coast. Thus, it can be assumed that the proportion of fish in tiger’s diet on the coast and in the inland region should be similar. Apparently, the consumption of salmon by tigers can be neglected in this analysis.The minor role of fish in the Siberian tiger diet is further evidenced by comparing its average mercury content with other large felids known to consume fish and other aquatic animals. Thus, individuals of Florida panther (Puma concolor coryi), consuming aquatic and fish-eating animals have elevated levels of MMHg—1.62 ± 1.87 mg/kg or 1.84 mg/kg THg 20. The average mercury concentration in the jaguar (Panthera onca), mainly preying on fish and alligators, reaches even higher values of up to 4.27 mg/kg (from 2.13 to 7.26 mg/kg) 21. The average THg content in the Siberian tiger is 0.383 ± 0.062 mg/kg indicating that it eats little fish, if any.In the south of the Russian Far East, some ungulates caught by the tiger on the eastern macro slope of Sikhote-Alin periodically go to the sea to lick salt and eat algae. This can lead to some increase in the level of mercury in their tissues and in the tiger along the trophic chain. In addition, ungulates, especially deer, can eat various lichens, including Usnea in the temperate forests.As we found out, THg in lichens from the coast, where sea fog is observed, was 0.170 ± 0.017 mg kg−1 (n = 30), which is 2.6 times higher than the average value for inland areas (0.065 ± 0.004 mg kg−1 (n = 24) (Fig. 1A). The absolute values of THg in Usnea lichens from the coast in the south of the Russian Far East turned out to be higher than in the Ramalina menziesii lichens (from the same order Lecanorales and the same ecological form as Usnea) from the coast in California 3 (0.138 ± 0.012 mg kg−1). It is possible that such differences are related to species-specific features of their thalli. Different species of lichens from the same locality can accumulate different amounts of toxic substances in their thallus. Thus, Usnea contained 0.170 ± 0.017 mg kg−1, and the mesomorphic evernia (Evernia mesomorpha) collected on the same site—0.292 mg kg−1(n = 2).Figure 1Map of sampling sites and mean values of (A) THg concentrations in lichen (Usnea sp.) (site names correspond to data in Table 1), (B) THg in tiger fur. The blue triangles and circles represent the samples from coastal sub-region and the black—from inland sub-region. Lichen sampling was done in 2019 and tiger fur sampling was done in 2004–2014. The map was generated in Adobe Photoshop CS6, based on a map from the public domain on the site https://yandex.ru/legal/maps_termsofuse/?lang=en.Full size imageWeiss-Penzias et al. 3 do not give the average THg for inland lichens, but they show that the high MMHg content in lichens on the coast is obtained through coastal marine atmospheric fog. We compared our data with those for THg on Bathurst Island 22, where the spatial pattern in THg enrichment was very similar to that of MMHg, with enrichment highest at coastal sites and decreasing within 10 km, suggesting similar origins of atmospheric THg and MMHg to lichens. Potential sources of inorganic Hg and MMHg to lichens are diverse (e.g. 3,22). MMHg from THg can range from 4.4 to 23% 3, therefore special future studies are needed to understand the dynamics of Hg species in lichen.The average individual THg concentrations in tiger fur samples from the coast ranged from 0.115 to 0.918 mg kg−1 (n = 12), on average 0.434 ± 0.067 (Fig. 1B, Table 1), while tiger fur samples from the inland regions (n = 12) had lower concentrations of THg (range from 0.057 to 0.950 mg kg−1, average 0.239 ± 0.075); the differences between the means of the two sites were statistically significant (p = 0.01) (Fig. 1B, Table 1).Table 1 Statistics on the subsets of THg concentration data used in this paper.Full size tableThe average concentrations of THg in the tiger fur of two subregions were lower (0.434 ± 0.067 and 0.239 ± 0.075 mg kg−1) than similar values for pumas from California 3. In contrast to California, where the average THg content in pumas from the coastal area was three times higher than in animals from the inland areas, in the Russian Far East the average THg content in tiger fur sampled in the area influenced by the sea fog was two times higher than that in comparable samples from inland areas. Although, it is the inland area (the Amur River basin) that is subject to high levels of human activity, including the mining of coal and gold in the past and present, and we could expect higher levels of mercury in living components of ecosystems. The average concentrations of THg in the tiger fur were only from 0.056 to 0.232 mg kg−1 (n = 4) near coal and gold mining sites.Different age classes were sampled in both the coastal and inland areas, and THg concentrations increased with age (adult  > young) in both areas (Table 1). This pattern is typical for predatory animals in general and for pumas, in particular 3, which is natural due to the cumulative effect and increasing mercury content with age 22,23,24,25. Differences in the average individual mercury content in individual fur samples of young and adult tigers were significant (p = 0.04) (Table 1). Moreover, the differences between the average mercury content in young and adult tigers were insignificant in the inland site (p = 0.32), while being significant on the coast (p = 0.04) (Table 1).We did not observe any significant differences in THg concentrations between the sexes (p = 0.86) (Table 1) and between males from the coast and the inland (p = 0.25) (Table 1) as was noted earlier for puma 3,21. On the contrary, the average values of mercury in the fur of individuals from the coast were 3.1 times higher than from the inland sites within the group of females, this difference was statistically significant (p = 0.03) (Table 1). These data contradict to what was observed in puma by Weiss-Penzias et al. 3. Such feature of female tiger is apparently associated with their shorter migration routes and smaller individual territories compared to males 26,27. Unlike males, which can cross the main Sikhote-Alin ridge, females are usually located either on the territory in the zone of sea fog influence, or in the inland areas.In addition, young females often remain within the territory of their mothers during dispersal 15. Rather similar information was obtained for the wild European cat 28, where THg in females was about 1.4 times higher than in males, although the differences were not statistically significant. There were no significant differences in THg content between young tigers in coastal and inland areas, as well as in the samples of animals, which died in autumn–winter and spring-early summer (Table 1).Apparently, preying on land animals does not lead to the accumulation of high Hg levels in felids. The average Hg levels in the fur of a near-water species such as the ocelot (Felis pardalis) mainly preying on terrestrial animals, varied in the same range as the tiger: 0.5–1.25 mg kg−1 29. Tigers are mainly consumers of the second level and therefore the average content of mercury in their body (0.383 ± 0.062) (Table 1) is lower than in the fur of consumers of the third level such as the pine marten 1.80 ± 1.34 mg kg−1 30 or Daubenton’s bat 1.15 ± 0.27 mg kg−1 31.The only sample with a maximum mercury content of 1.402 mg kg−1 (age and gender unknown) was from the southwestern Primorye, which is located on the coast and where there are cinnabar deposits nearby. This sample was not used in the total analysis. Interestingly, the available sample of a young female Far Eastern leopard fur (Panthera pardus orientalis) from the same site had practically the same mercury content (1.456 mg kg−1). Local increased mercury content in the body of tigers can be associated with deposits of mercury-containing minerals. These data do not confuse our understanding of the sources of mercury in the ecosystem; they serve as a signal for a more profound study of natural processes.Our data on a higher mercury level (THg) in lichens and tigers of the sea coast compared to inland areas may be related to the effects of coastal sea atmospheric fog, a potential source of monomethylmercury (MMHg) produced in the ocean 3.The levels of mercury we found in Siberian tigers from the Russian Far East are about four times lower than the mercury content in the fur and vibrissa of puma from California 3. It seems that such differences are related to the position of these regions relative to the zones of deep faults of the mantle formation of the East Pacific platform 32. The maximum concentrations of Hg in the near-surface atmosphere are confined to such zones, and the concentrations decrease at a distance from them. California is located closer to such zones, while the south of the Russian Far East is further away.And the fact that different levels of mercury in ecosystems depend on the distance relative to the deep faults of the East Pacific platform is confirmed, for example, by pink salmon: fish from the Sea of Japan contain much less mercury than fish from the Kuril region closer to the fault zone (from 0.045 to 0.087 mg kg−1 wet weight 31. At the same time, we must not forget that California is the most populated and one of the most industrially developed states in the USA. However, it seems that natural processes currently play the main role in formation of heavy metal content in the discussed populations. Thus, lead concentrations in organs and tissues (liver, gonads, and muscle) of fish from Kuril oceanic waters was one and a half order of magnitude higher than that of pink salmon from the Sea of Japan 33.If the global anthropogenic mercury pollution of terrestrial and aquatic ecosystems continues, coastal food webs in the zone of influence of the East Pacific platform will be at most risk of toxicological effects. More

Abundant production of a type IV pilus in Synechococcus sp. WH7803We first detected an abundant PilA protein (i.e. SynWH7803_1795) in the extracellular proteomes of the model marine cyanobacterium Synechococcus sp. WH7803, accounting for up to 25% of the exoproteome19,20. Transmission electron microscopy confirmed the existence of the macromolecular pili structures (Fig. 1a and Fig. S1). Unlike Synechocystis sp. PCC6803 that simultaneously produces thick and thin pili14, this marine picocyanobacterium presented multiple pili of similar thickness (diameter of ~6 nm), each ~10 µm in length. The amino acid sequence of PilA revealed a typical Sec-targeting signal peptide and a conserved GFTLxE motif at the N-terminus of the protein (Fig. 1b) that is known to be cleaved in the cytoplasmic membrane by PilD before the protein is translocated to the base of the pili for assembly10. After cleavage, the N-terminal of PilA can be post-translationally modified, e.g. methylated, to increase the hydrophobicity and stability of the pilin10,21, although we were unable to detect this modified N-terminal tryptic peptide during proteomic analyses. In close proximity to pilA in the Synechococcus sp. WH7803 genome we found five other type IV-like pilin genes (Fig. 1c), all with the conserved GFTLxE motif (Fig. 1d).Fig. 1: Pilus in the marine cyanobacterium Synechococcus sp. WH7803.a Transmission electron microscopy images of wild-type Synechococcus sp. WH7803 (WT) and pili mutant (Δpili) obtained from late-exponential liquid cultures incubated in ASW medium under optimal growth conditions. Imaging of three independent cultures in different occasions consistently showed long pili appendages only in the wild-type strain (Fig. S1). Middle panel image, obtained with the same magnification as other panels, is from an intercellular region between wild-type cells to improve the visualisation of the pili. Scale bar represents 1 µm. b The amino acid sequence of PilA1 (SynWH7803_1795). Trypsin hydrolytic sites are indicated in blue. Red lines highlight tryptic peptides detected by shotgun proteomics. The conserved GFTLxE motif is shown and the cleavage site is indicated with an asterisk. c Genomic context of pilA1 in Synechococcus sp. WH7803. Numbers in each gene represent their ID number (SynWH7803_). In red are genes detected by proteomics. While PilA1 and PilE are abundantly detected in exoproteomes20, PilA2 has only ever been detected in cellular proteomes of this strain22. Blue dotted lines indicate genes encoding possible structural pilin pairs, i.e. PilA1-PilE, PilA2-PilV and PilA3-PilW. Question marks indicate genes encoding proteins of unknown function. d The N-terminal amino acid sequence of PilA1 and five other pilin-like proteins, all with the highly conserved GFTLxE motif. e Synechococcus sp. WH7803 structural pilus proteins identified by homology with S. elongatus PCC 794216 and assembled in the inner (IM) and outer membrane (OM) as modelled by Craig et al11. pilM and pilO were not identified by homology but the SynWH7803_2367 and SynWH7803_2365 genes are suggested because they form a standard pilMNOQ operon as found in other species. While S. elongatus PCC 7942 encodes three pilT, only two were found in Synechococcus sp. WH7803, one being part of the characteristic pilCTB operon.Full size imageUsing the pilus apparatus from the freshwater cyanobacterium S. elongatus PCC 7942 as a reference16 and the established architecture for type IV pilus machinery11, we were able to find all components necessary for pilus assembly in Synechococcus sp. WH7803 (Fig. 1e). We speculate that the genetic cluster encoding the six pilin-like proteins (Fig. 1c and 1d) may provide three distinct pili functions. Based on homology with the annotated genes from S. elongatus16 and conserved domains found using the CD-search tool in NCBI, we suggest the three pilin pairs: PilA1-PilE, PilA2-PilV and PilA3-PilW (Fig. 1c). Of these, shotgun proteomic analyses have only ever detected PilA1-PilE19,20 implying these are responsible for the pili observed in Fig. 1a, although PilA2 was also detected in low abundance in cellular—but not extracellular—proteomic datasets22. Unlike in S. elongatus, where PilA1 and the contiguously-encoded pilin-like protein are almost identical, the amino acid sequence of PilA1 and PilE in Synechococcus sp. WH7803 are clearly distinguishable. Although in much lower abundance, PilE seems to be correlated with PilA1 in the exoproteomes of this cyanobacterium19,20 and, therefore, it is possible that PilE and PilA1 form subunits of the same pilus apparatus.Pilus distribution amongst picocyanobacterial isolates and Single-cell Assembled Genomes (SAGs)Genomic analysis of sequenced marine picocyanobacterial isolates downloaded from the Cyanorak database23 revealed that 74% of sequenced Synechococcus (n = 46) and 33% of Prochlorococcus (n = 43) encoded pilA1 (Fig. 2 and Supplementary Data 1). In Synechococcus, pilA1 was prevalent in all clades (93%; n = 28) except for clades II and III where it was less abundant (44%; n = 18). Interestingly, all low light Prochlorococcus isolates from clades III and IV encoded pilA1 (n = 7; Fig. 2). Most of these pilA1-containing strains also encoded a pilE homologue in close proximity (Fig. 2). Genes pilA2 and pilA3 were also abundantly found in Synechococcus (59 and 74%, respectively), although were much less prevalent in Prochlorococcus (12 and 9%, respectively). As expected, all strains that encode at least one of the pilA types also possessed the transmembrane pilus apparatus, whereas this apparatus was completely absent or partially lost in strains lacking pilA (Fig. 2). PilA3 is known to be involved in DNA uptake and competence in S. elongatus, requiring additional competence proteins to do so16. Marine picyanobacteria are not known for being naturally competent but, interestingly, all strains encoding PilA3 also contained the competence genes encoding ComEA and ComEC (Fig. 2). Further work is needed to investigate the conditions under which the PilA3-type pilus becomes active in these organisms and, therefore, when exogenous DNA might be taken up.Fig. 2: The presence of pilus-related proteins in cultured marine picocyanobacteria strains.Pilus proteins from Synechococcus sp. WH7803 were used for the BLASTp search. Log10 E-value scales are shown (1 to More

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Spotted wing drosophila captures within the United StatesComparison between the raspberry field and wooded area, during pre-harvest and harvest periods to account for presence of developing and fully ripened fruit, SWD captures and selectivity per QB dry sticky trap is found in Fig. 1A,B. No difference was found in average capture per trap between either area during the pre-harvest period, nor was there a difference between these and the field during the harvest period. The wooded area during the harvest period captured the greatest amount of SWD/trap (F1,209 = 7.335, P = 0.007) (Fig. 1A). Dry sticky traps baited with QB had a significantly higher selectivity during the pre-harvest period in the raspberry field than in the wooded area but was not significantly different from the trap selectivity in the wooded area during the harvest period. The pre-harvest wooded area trap selectivity was not different from the harvest field trap selectivity. While the harvest field trap selectivity was lower than that of the wooded area trap selectivity during the same period (F1,203 = 23.6, P  More

Traits are attributes that speak to biophysical limitations, pressure on species, ecological functionality, and interactions. They have found their way to the forefront of many discussions and debates about ecosystem dynamics and, with a slight time lag, social-ecological systems1,2,3. The promise is that a traits framework can further our understanding of patterns, dynamics, interactions, and tipping points within and across complex social-ecological systems. But what will it take to make good on this promise, in particular for our cities, where change is fast and—being the places where the majority of humans live—human perceptions are particularly diverse? What kind of framing, what research, would allow traits—classically understood as a different representation and interpretation of well-established and known properties of the social-ecological system―to fully work as “mediators” for understanding the behavior, functions, and needs of urban systems under pressure?This perspective aims to contribute to the current wide-ranging discussion about traits in both theoretical and applied ecology, and parallel work on better understanding human connections to nature. To this end, we explore the potential of using an expanded conceptualization of traits as a platform for integrated approaches to understanding the different facets of people-in-nature relationships and dynamics4,5.Expanding from the original “characteristics which have demonstrable links to the organism’s function”6, we see traits as a nexus where different theories and conceptualisations about social-ecological systems can connect, intertwine and comprehensively allow us to assess the current state of a system—and even more importantly, evaluate the implications of change (Box 1 and Fig. 1). To make it an integrative and useful framework for urban studies and policy/practice, traits need to be easy to recognise and relevant to decision makers across scales and in different contexts. In addition, information on trait profiles—generic as well as site specific—need to be easily available through monitoring or in databases.Fig. 1: Traits within social ecological systems.Theoretical flow chart linking the entities of a social-ecological system to its traits, demonstrating how a traits framework—as outlined in this article—might be positioned to support the analysis, interpretation and governance of urban systems.Full size imageOur argument is threefold: The first dimension focuses on how to assess and anticipate change by establishing chains of interconnected traits that describe and causally connect sensitivity and response to different urban pressures such as heat, soil compaction, environmental toxicants, and stormwater runoff, understood through “response” traits7,8,9,10 to their functional consequences11, mediated by “effect traits”. The second dimension is grounded in human perceptions and appraisal of diversity to highlight the different cues and characteristics people use to detect change or articulate value narratives, and it is linked to the role of traits in ecological literacy. Here, we propose traits be viewed as boundary objects, i.e., features that carry meaning across society (although the meanings might be diverse and sometimes conflicting), and that this second dimension is essential for understanding the role of society and humans in a traits framework. The third dimension outlines how the first two dimensions connect to inform and support decision making and management at different scales, for example in different, multilayer, and multiactor governance processes12 (Fig. 2).Fig. 2: A traits framework for scientific study and practical application.The three dimensions of a social-ecological traits framework for understanding and governing urban systems. The first dimension is represented by observable traits of the urban environment, e.g., features of humans and other co-inhabiting species and their differing responses to pressures and selection, leading to functional consequences and finally, altered characters of an urban social-ecological system. The second dimension is characterized by feedback loops between those effect outcomes and individual and collective perceptions and decision making. Lastly, the third dimension is represented by urban ecosystem planning and management embedded in governance processes and instruments. Through its ability to connect different spheres and discourses, an expanded traits framework can aim for effective and inclusive decision support that is responsive and place-adapted. By expanding and bridging these three dimensions, we can connect different insights and knowledge about ecosystem function and human perceptions, values and interactions with the environment. This will support the development of a (meta-) theoretically grounded, practically applicable traits framework to interrogate reciprocal feedback linkages and nature-human relationships. The figure includes resources from Freepik.com.Full size imageBox 1 definitionsFunctional trait: A feature of an organism which has demonstrable links to the organism’s function69, and, as such, “determines the organism’s response to pressures (response trait), and/or its effects on ecosystem processes or services (effect trait). In plants, functional traits include morphological, ecophysiological, biochemical and regeneration traits, including demographic traits (at population level). In animals, these traits are combined with life history and behavioral traits (e.g., guilds, organisms that use similar resources/habitats)”70, p. 2779.Boundary object: “[…] those […] objects which both inhabit several intersecting social worlds and satisfy the information requirements of each of them. Boundary objects are objects, which are both plastic enough to adapt to local needs and the constraints of the several parties involving them, yet robust enough to maintain a common identity across sites. […] They have different meanings in different social worlds [and across cultures] but their structure is common enough to more than one world to make them recognizable, a means of translation.”71 p. 393, see also72.Social-ecological traits (expanded definition): An ecologically or socially (inter)active and demonstrable feature of the environment at any level or scale. A social-ecological trait either mediates reactions to selective social-ecological filtering (response trait) or determines effects on ecosystem processes or services (effect trait), or both. The aggregate trait profile of a given entity should ideally speak both to ecological functioning and socio-cultural meaning.The first dimension: response and its effect outcomesTrait-based approaches have been used for descriptive purposes13 to enable broader global comparisons that transcend the constraints of regional taxonomic diversity (e.g., see refs. 6,14) and allow for the types of generalizations sought in ecology15,16. Traits offer a way of looking at causality and change, and trait profiles can indicate whether emergent communities are functionally different from historic communities. To this end, traits can be divided into those that determine an organism’s sensitivity and response to environmental factors, and those that relate to its effect on the environment4,17. When combined, the two categories of traits can be used to detect, identify and monitor the current state of ecosystems, and to anticipate the outcomes of change8,10,17,18,19.An environment described through traits: The urban bio-physical environment includes hydrology and soils, as well as biotic elements (flora and fauna), and understanding the relationships among those components is necessary to measure and anticipate the profound effects of urbanisation. Currently, knowledge of plant traits is most developed4,20, although there is work emerging on traits for animals or other taxonomic groups8,21 as well as for soil and geodiversity22. Animal studies so far tend to focus on habitat modelling for birds, insects, invertebrates and a few on mammals (e.g., see refs. 3,8,16,23). Many studies have looked at the impact of different community assemblages on ecological functions through effect traits and, in particular, how altered or dynamically changing communities will affect ecosystem process through changes in representation of effect traits (but e.g., see ref. 23). However, the link between traits and ecosystem functions has largely been inferred (ibid.), and is, according to Cadotte et al.24, rudimentary (see also25 and26). As we indicated with our definition of traits (Box 1), we see a value in including soil properties as traits and not to leave them as “environmental filters”, as this may offer a more dynamic way of understanding one of the major urban processes of change—soil sealing and compaction—and thus help guide urban development.Traits at different levels and scales: Traits at the species level are by far the best known and most explored, but there are also studies that use traits from other ecological levels—gene, community, ecosystem and landscape—as indicators for tracking response to stress27 and calculating functional “performance”. A common approach to scale is to aggregate species level information. For example, the average values of aggregations of plant species traits at the ecosystem level provide a basis for calculating overall sensitivity to pressures28. This in turn, and drawing on different sets of traits, allows for estimations of changes in ecosystem function (e.g., see ref. 29). However, there are other characteristics that could also be understood as traits. At the landscape level the mosaic of ecosystems and the location and combination of patches are used to assess flows and exchange across larger areas (e.g., see ref. 30). A good example is a city in a river valley, where water flows and exact location within the drainage basin affect urban green spaces and their aggregated matter production, CO2 absorption or carbon sub-section31. Aggregate, or higher-level traits, such as structural composition and functional diversity of vegetation, matter flows, or species migration, are the most common traits analysed through remote sensing in order to track trends25. More work needs to be done to explore relevant traits at different levels of organisation to match the scale and nature of disturbances and the spatial and temporal scale at which different functions are most relevant. Being explicit about scale, and ensuring traits at different levels are nested, allows for tracking of processes across scales.Individual traits, trait combinations, and interlinked suites of traits: A key promise of traits is to provide mechanistic explanations of observed structure, patterns and functionality, which is usually demonstrated through statistical correlations. Further developing suites of response and effect traits could provide valuable input and indicators for assessment and monitoring frameworks. For example, traits could inform DPSIR (drivers, pressures, state, impact, and response) models by anticipating or measuring response to a pressure and the direct and indirect impact this response could have. At a more fundamental level, traits explain whether impacts may be causing a change in the functional state of the system. Interlinked traits, from those determining sensitivity, to those mediating response elicited by sensitivity, could improve mechanistic understanding by supporting the development of stepwise response-effect pathways17. For example, land conversion—like the soil sealing and compaction typical in cities—fundamentally alters soil properties, which in turn affects vegetation. Soil properties influence the growth and composition of plant communities. This translates into trait-mediated effects like reduction of total leaf area, which leads to cascading effects of early leaf senescence and limitation of stomatal transpiration. This reduces water exchange capacity, which in turn is key for mediating air cooling or shading and other functions/services plants may offer to humans.For this first dimension, trait databases, classical field inventories, and experiments, remote sensing data, and GIS-based information are crucial15,32. We see valuable developments from the past two decades of research towards achieving a traits response-effect library in both the ecology and remote sensing communities33,34, even if recent advances from remote sensing studies still rarely find entrance into urban planners’ work and policy decision-making35. In particular, the development in the technical dimensions of detecting traits and trait variation20,34, and tracking these over time, has recently rapidly developed. The progress in application of high-resolution hyperspectral data, light detection, and ranging (LiDAR) or the possibility of mounting the recently developed sensors on unmanned aerial vehicles (UAVs) equip the researchers with addditional tools that can not only expand the range of measurable traits but also allow easy access to data. This provides a powerful support for urban planning and, ultimately, urban governance. Moreover, applications for tablets or smartphones offer alternative ways to directly involve citizens in ecosystem monitoring and further develop citizen science (e.g., see refs. 22,36).The second dimension: traits as an interdisciplinary bridgeThe literature explicitly using the term traits tends to focus on soil, geodiversity, plant, and community trait profiles as an outcome of social-ecological selection through environmental conditions, species interactions, human preferences, management regimes etc. (e.g., see refs. 4,37). This approach has started to address not just how people filter traits (e.g., see ref. 38), but the reason(s) behind either individual or group decisions that lead to filtering (e.g., see refs. 39,40). Here, we propose that the environment, described through traits, could be considered a boundary object (Box 1), allowing for a multiplicity of views, disciplinary connections, engagements, and perceptions, and that speaks to the complexity of social-ecological systems. This will expand the range of functions used to describe a system, and the types of traits required to capture them.Ecological functions relative to ecosystem services: The plant and animal traits that people respond to may not be the same ones that mediate responses to environmental change. For example, seed mass and specific leaf area are important plant functional traits41 but are less likely to influence people’s preferences for urban vegetation (e.g., see ref. 42). Indeed, some esthetic traits promoted by human decision-making and management, such as selection for leaf variation and predominantly deciduous plants, may also lead to the predominance of woody plants that are strongly affected by water stress, fungal attack or insect infestation or trimmed canopies, and thus promote reduced fitness of individual organisms and communities43. On the other hand, a successful reproductive strategy such as the emission of high quantities of pollen might limit the suitability to human-dominated environments (including cities) due to allergenic potential44. Do we need more, or different traits to link ecosystem dynamics more strongly to the lived reality of people? Are traits too simplistic proxies, or perhaps too specific features, to express and understand people–nature interactions? Introducing humans and human appraisal into our trait framework encourages a broader definition of what might be relevant traits. Traits used in this way provide a specific link to interactions and feedback mechanisms between human wellbeing and functional ecology (and respective proxies that serve multiple relational (feedback) purposes).Traits as relational features: Trait lists already include features which are easy to understand and readily detected by human sensory organs, and thus find traction in society or connect to existing ethno-biological narrations39. Traits such as flower colour, leaf shape, and canopy density, which may not necessarily be considered central functional traits, are important drivers of people’s preferences37,39,45,46. Both size and colour of the flowers are plant traits affecting people’s perception47 and can thus be an important factor for gaining societal approval for more urban greenery48. Seasonality is another relevant trait; for example, an extended flowering season49. At the same time, there is a growing interest in flowers and blooming meadows among gardeners worldwide also to support insects in urban landscapes to counteract global biodiversity decline37,39.In this vein, we argue that traits are a formative force influencing human wellbeing and world views, giving shape to ecological systems and linked human affordances (through, e.g., shade and sensory stimuli), and social systems by shaping the context of human activities and experiences. For example, we know that people recognize and value a wide range of plant traits, and that this has even been identified as a useful way to speak about the state of nature and large scale change50. There is evidently a role for traits and trait composition as language for more “functional” ecological literacy36,50. This position as a boundary object needs to be further explored and linked to the responses of social-ecological urban systems, which are subject to a multitude of pressures, including climate change and soil sealing.Traits as boundary objects and connectors between knowledge systems: What is needed to better position and connect the concept of traits to multiple different literatures and disciplines and enable traits to be used as a useful boundary object? Many disciplines outside the ecological and environmental sciences have an interest in understanding ecosystem function and biodiversity, and how people relate to these ideas. Traits, and deeper meanings of some traits, can be found within environmental psychology, ethno-botany/zoology and environmental anthropology. Trait-based approaches may also be well suited to engage with other ways of knowing, such as traditional ecological knowledge and religious systems. This disciplinary and trans-disciplinary knowledge is needed if traits are to connect social-ecological attributes to diverse human values and wellbeing dimensions, and to ensure we do not produce trivial and culturally biased conclusions51,52. Based on the diverse use and potential meanings of the word “traits”, we argue that a traits framework, and traits-focused interdisciplinary discussions and projects, could support a dual ontological stance where some connections are more universal, while others are inherently interpretational or simply individual. Hence, this may help to effectively connect the social and cultural dimensions of traits to a deep ecological understanding of change and its multiple consequences. This would be an important development that allows for critical engagement with concepts like tipping points and system states and what they actually mean in a complex social–ecological urban system.The third dimension: traits for decision supportThe major purpose of the traits concept, as we present it here, is to develop an ontologically inclusive traits framework capable of addressing both the resilience of ecological functions and the experiential and relational aspects of human interactions with nature. On the applied side, this would be relevant to a wide range of decision-making processes, not least urban planning. Clearly visible and easy-to-map traits are well-suited as indicators to describe the state of urban landscapes relevant for biodiversity and society alike. To this end, there are still many questions that need answers. For example, how can the understanding of trait profiles help improve species selection in times of climate change, to inform management priorities and strengthen cross-community stewardship, especially where the diversity of response traits may be low? And which traits are incompatible and how are they best kept separate, a question particularly relevant in the light of zoonosis like the COVID-19 pandemic in 2020? And finally, what traits could best serve as reasonable proxies or indicators to provide either cues or early signals of species responses to (fundamental) change in urban environments?Supporting holistic decisions: Already now we see increasing use of traits in modelling and decision support tools like CiTree and iTree53,54. As cities strive to adapt to climate change by, for example, revising tree species selection (e.g., see ref. 55), an improved understanding of the relationship between detectable functional traits and the provision of ecosystem services can help avoid maladaptation56. For example, replacing shade trees with fine-foliaged trees may improve adaptation to future climates but would not provide the same levels of climate mitigation57. From a decision-making point of view, key traits are those determining the response of ecosystems to human-induced pressures such as air pollution, soil sealing, or urban heat islands, as well as those mediating the effects of these changes on ecosystem services and related benefits as perceived by people8,58.A traits framework that uses our social-ecological definition of traits might support informed decisions on trade-offs. For example, invasive or non-native plants are often seen as ecologically problematic, but certain traits such as high leaf coverage or flower colour and shape make them socially desirable48. Traits connected to more social-ecological dimensions will allow for a more holistic assessment of options and the potential trade-off implications of different choices. While decisions are often grounded, implicitly or explicitly, in considerations of multiple traits (e.g., see ref. 53), we need to ensure that traits considered in the plant selection include both traits related to broad and diverse preferences and desires for ecosystem services and traits, that ensure a resilient response to drivers of change that may impact their ability to provide these services (see, e.g., the scoring system for urban vegetation species proposed by Tiwary et al.59).Urban planning informed by an expanded traits framework and spatial-temporal patterns of trait profiles has the promise to be adaptive in the best sense and thus, resilient. More city and regional comparisons are needed to make target setting and threshold discussions grounded and allow for global discussion. This requires a targeted effort at broader inclusion of cases and trait data from different climates, biomes, multiple ecological levels but also cultures, and would move traits studies towards a truly transdisciplinary venture with real impact on how we plan and manage our cities.Feasible and easy to use: Indicator traits need to be robust, easy to measure and low-cost to assess, and have a causal link to relevant social-ecological processes and patterns (such as ecosystem services for recreation, cooling or food4,60). The potential use in planning and decision-making at multiple levels again point to the need to discuss the scales and levels for traits studies to make sure trait levels are nested and logically commensurable. Higher-level, larger-scale properties such as landscape morphology and water availability, the profile of pest communities or potential invasions can be further informed by the development of more detailed traits frameworks. This makes traits frameworks highly relevant also from an economic, social and health perspective, especially in intensely managed environments like cities, where combinations of multiple stressors and external factors create small scale heterogeneity and fast temporal change in pressures61,62.Trait selection can play that important role for assisting in the planning and design and then evaluation of the functionality of high-biodiversity green spaces63, and for trait-informed assessment of “performance”, e.g., of ecologically protected areas. A relevant example to this point is the ongoing debate about how to evaluate ex-ante, and then monitor, the implementation of nature-based solutions62,64, which remains a challenge65. Could this be done using traits instead of commonly used area-based indicators? Could traits become the basis to design and assess the impacts of offsets and compensation measures, thus increasing their efficacy? From this perspective, we see in a traits framework the potential to support a shift towards more flexible and effective planning approaches, more suitable to address today’s urban challenges and to promote greater well-being, sustainability and resilience of present and future cities.Conclusion and looking aheadThrough their direct relation to ecosystem services such as cooling and fresh air, easy-to-understand traits can be an entry-point for nature awareness and, subsequently, ecological knowledge in decision-making both at the citizen and the societal level66. However, to make traits successful indicators of global, regional, or local environmental changes, it is vital that urban society is understood as diverse across characteristics such as cultural background, physical mobility, gender, age, degree of formal or informal education, access to information and communication, purchasing power, and political influence67. All these factors affect the needs, preferences, and values of individuals and groups, and the way each interpret human-nature relationships. Only by taking these factors into account, planning for spatial-temporal diversity in traits across an urban landscape will create more inclusive urban systems that foster multiple benefits for both people and biodiversity68.The expansion and implementation of a traits-based approach for urban systems is impeded by availability of traits data. For example, trait databases are usually a primary data source in studies on urban ecology, however, these data have mainly been collected in natural areas or controlled environments such as laboratories, where organisms may display different trait values than those in urban environments. Studies have also been concentrated in the global north, and there are major challenges with potentially transferring and adapting thinking mostly developed in the Global North to rapidly urbanising areas in Africa, Asia and South America.To enable a social-ecological traits framework for interdisciplinary discussion and for guiding urban planning and decision making, we suggest a three-pronged approach for building a social-ecological understanding of trait mediated interactions and their implications, and make this understanding useful to practice (Table 1). Large-scale monitoring needs to be coupled with in-depth understanding of response mechanisms and their impact on ecosystem functions as well as services, and a deeper connection between traits and human perception as well as sense-making of the world we live in. Application to human perception and sense-making requires more data, theory and empirical work, and especially the way people relate to traits will likely vary considerably across cities and contexts across the globe. All branches of investigation need to be embedded in an interdisciplinary discussion about the role that traits play for social-ecological interactions and mutual exchange. Drawing on this broad evidence base, synthesized knowledge will offer a more comprehensive support for urban decision making, not least in anticipation of future change.Table 1 Research agenda.Full size table More