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The data reported in this study expands upon the present knowledge concerning the mineralogy of coralline algae species worldwide, encompassing for the first time coralline algae species data from the Southwest Atlantic Ocean, where this group is the main frame-builders in coral reefs and the major inner component in rhodoliths16,26.Mineralogical analysis revealed that coralline algae species of the Brazilian Shelf were mainly formed of high-Mg calcite. Six coralline algae species in this study had the same range of high-Mg calcite, between 80 and 100%, than the same species from different regions of the world: Lithophyllum corallinae, Lithophyllum kaiseri (as Lithophyllum congestum), Lithophyllum stictaeforme, Lithothamnion crispatum, Melyvonnea erubecens (as Lithothamnion erubecens) and Sporolithon episporum (Table S2). This result confirms that species from different families, such as Corallinaceae, Hapalidiaceae and Sporolithaceae have a CaCO3 skeleton formed mainly of high-Mg calcite.In agreement with earlier studies, the average high-Mg calcite content in Corallinaceae was very similar to the results compiled by Smith et al.11 (96.7 wt.% and 96.2 wt.%, respectively). This pattern was also observed for Hapalidiaceae, which presented a mean value of 88.9 ± 3.6 wt.% in our study and 90.2 wt.%. However, Smith et al.11 registered a high-Mg calcite content of 98 wt.% for Sporolithaceae, while in our study this polymorph had a mean occurrence of 86.2 ± 6.5 wt.%. This percentage can be attributed mainly to the lower content of high-Mg calcite found in Sporolithon yoneshigueae, which is an endemic species of the Brazilian Shelf27.The high similarity between the mineralogy (% high-Mg calcite, % aragonite and % dolomite) of the species belonging to three encrusting algae families, revealed by the cluster analysis, emphasizes the lack of CaCO3 disparities over skeleton mineralogy of coralline algae at family level. This aspect was also evidenced by several studies concerning coralline algae mineralogy11,21,22,23,24,25. This fact was confirmed in the cluster analysis between the mineralogy of the studied coralline species, in which samples from different families were grouped. Considering these findings, the mineralogical pattern exhibited by the crustose algae may not be driven by taxonomic classification, as was first proposed by Chave28. Therefore, the skeletal mineralogy from Brazilian coralline algae species can not be used as a taxonomic character, not even for higher taxonomic levels.In this sense, the mineralogical analysis from L. crispatum, the most common rhodolith-forming species on the Brazilian Shelf16, revealed that samples from the Abrolhos Bank presented higher high-Mg calcite in their composition, and the highest % of Mg substitution in the calcite lattice than the species from the other four regions studied. One of the possible explanations is that the Abrolhos Bank has the highest seawater temperature compared to the other four sites, which influences CCA mineralogy. This result corroborates the hypothesis that coralline algae species do not have a strict control over Mg precipitation as stated by Stanley et al.29. In addition to seawater temperature, Mg/Ca ratio in seawater can also affect the incorporation of magnesium into coralline algae skeletons11,29.In relation to other CaCO3 polymorphs, previous studies have registered some species with up to 20% aragonite11,12. Meanwhile, in this study, S. yoneshigueae presented CaCO3 skeletons formed of more than 30% of aragonite, which expands the range found in coralline algae for this polymorph. The high percentage of aragonite found in S. yoneshigueae could be related to the fact that this species presents larger overgrown calcified empty tetrasporangial compartments, in comparison with other Sporolithaceae species27, which could be filled with aragonite. This feature has mainly been described in the overgrown conceptacles of Lithothamnion sp.30 and in cell infills of Porolithon onkodes31. The presence of aragonite could be also attributed to the use of aragonite granules in the sediment to repair any damage in the alga-substrate attachment32.Raman mapping showed the presence of high-Mg calcite in the bulk of the cell wall with little aragonite in its inner part, which seems to form an inner “shell”, closer to the cell membrane. To date, this is the first study that has utilized Raman maps to show the localization of aragonite in cell walls of coralline algae. The maps consisted of the cellular living layer from the coralline algae crust, right beneath the epithelial cells, which indicates that the mineralization of aragonite occurred in live cells and it was probably not a remineralization process.Aragonite inside cell bodies was first seen by Nash et al.12 using Backscattered Scanning Electron Microscopy. They also reported the presence of dolomite or protodolomite, which were not observed herein by Raman spectroscopy, probably because of the low amount of this polymorph.Previous studies considered that the inclusion of dolomite into carbonate skeletons is a microbial-mediated process after cell death upon the discovery of microbial-associated dolomite formation in anoxic marine33 and freshwater environments34. The presence of several calcium carbonate polymorphs found in coralline algae raises the question of whether all these polymorphs are in fact synthesized by the algae.Indeed, the role of coralline algae in the different forms of calcium carbonate crystal precipitation is a crucial issue that should be addressed. Nowadays, studies calculate the production of CaCO3 by coralline algae based on CCA coverage35, without considering that not all CaCO3 produced in that structure is related to coralline algae biomineralization processes (e.g. secondary calcification processes such as infilling of the older skeleton and skeletal dissolution vs newly deposited carbonate). Therefore, it would be misleading to presume the net CaCO3 accretion of coralline algae structures without knowing the origin of the CaCO3 processes. This is also valid in relation to studies on the influences of atmospheric [CO2] rise on coralline algae, based on weight changes36,37,38 and its impacts on the mineralogy of the existing crust21.Concerning Mg2+ substitution in the high-Mg calcite lattice, we found that Brazilian encrusting algae possess a higher Mg-substitution (46.3% more Mg2+ than the global average) in calcite than specimens collected worldwide. A possible explanation for the higher mean Mg2+ content might be related to the high seawater temperatures39, as this was also observed along the tropical Brazilian Continental Shelf. This can be exemplified by the high Mg2+ content found in fourteen species that occur in warmer waters of the Brazilian Shelf, where the mean surface seawater temperature (SST) ranged between 26.4 and 29.8 °C (from 2008 to 2016), between 17°S and 3°N. The lower Mg2+ amounts presented in L. margaritae and L. attlanticum could also be explained by the temperature, as these species were collected at the southernmost site (27°S) in the temperate zone, where the mean SST (from 2008 to 2016) varied between 22.5 and 25 °C (NOAA Comprehensive Large Array-Data Stewardship System-CLASS: SST50). A relationship between the Mg2+ content and temperature has already been proposed in previous works39 and is widely accepted. Nash and Adey40, when plotting the data collected using XRD, found a very strong correlation coefficient (R2 = 0.975) between mol% MgCO3 in coralline algae and temperature. Moreover, the Mg/Ca rate in coralline algae is used as a proxy archive41 and to generate multicentury-scale climate records from extratropical oceans42.Although seawater temperature is loosely associated with latitude, the New Zealand species, for example, are subjected to lower temperatures (2012 annual maximum and minimum surface seawater temperatures: 21 and 18.7 °C, respectively), while Caribbean and Cocos Island algae grow at higher temperatures (2008–2016 annual maximum and minimum surface seawater temperatures: 29.5 and 23.4 °C, respectively) (NOAA Comprehensive Large Array-Data Stewardship System – CLASS: SST50). If we consider the differences in temperature (≅ 6 °C) and Mg2+ content difference (7.67 wt.%) between the sampling sites along the Brazilian Shelf, we can infer that there is an average increase of 1.27 wt.% of Mg2+ per °C. This value is in the range from 0.4 to 2 wt.% Mg per °C reported previously, both in experimentally and in situ studies39.This relationship between Mg substitution and temperature is also critical in face of the temperature risen episodes that we are seeing all over the world43, including the Brazilian Shelf44. If coralline algae produces High Mg calcite with more Mg substitution in higher seawater temperatures, these thermal anomalies could force the production of a highly soluble polymorph, making coralline algae skeleton even more prone to dissolution.It is well known that high-Mg calcite is the most soluble CaCO3 crystalline polymorph under acidified conditions and that this dissolution is more evident when Mg substitution in the calcite lattice is higher45. In our study 70% of the coralline algae species presented a Mg substitution in the range of 12 to 24% and the mean Mg substitution was 21.1%, which reinforces the susceptibility of Southwestern Atlantic coralline algae to future high [CO2] scenarios.Even though previous experiments using synthetic calcium carbonate showed that the rise of seawater temperature increases Mg substitution, making high-Mg calcite more stable46 and other studies claiming that coralline algae with higher Mg substitution (more than 24% in average) presented less dissolution when exposed to high [CO2]13, Southwestern Ocean coralline algae are already living in a limit situation, where seawater can reach temperatures up to 28 ºC. Since we have a correlation between Mg substitution and temperature around 1.27% Mg per 1 ºC, it would take 2.4 to 6.2 ºC rise so the alga starts to produce a more stable calcite polymorph. Such a temperature rise could be lethal to these algae, also promoting a surface microbial shift that could be crucial to sucectional processes (e.g. settlement) involving other marine organisms, such as corals, which is critical for reef regeneration and recovery from climate-related mortality events47. The comparisons of results obtained through assays with synthetic calcium carbonate must be done with caution, because it should be take into account that the complex calcium carbonate biomineralization processes performed by marine organisms are highly dependent of a narrow range of environmental conditions.In face of the dependency of these environmental conditions, the broad range of Mg content in temperate coralline algae25, a high inter species variability in the % Mg in this study (Abrolhos Bank; 14.5 to 28.8% Mg), as well as an anatomical difference in Mg content in coralline algae40, suggest that other environmental parameters (e.g. Mg/Ca in seawater, light, salinity, etc.) could also drive Mg substitution in coralline algae. Furthemore, coralline algae biological processes might exert some kind of control over Mg-calcite calcification which make them more resilient under rising CO239.Long-term projections of ocean acidification and the CaCO3 saturation state indicated that high-latitude seawater will be undersaturated with respect to high-Mg calcite in the second half of this century45. Early results with coralline algae Sonderophycus capensis and Lithothamnion crispatum in a subtropical mesocosm in Brazil showed that an increase in seawater pCO2 (1000 ppm) enabled both species to continue photosynthesizing but did cause carbonate dissolution48.However, coralline algae from the North Atlantic Ocean, where the temperatures are lower, presented the lowest Mg substitution mean (11.91%), with some algae presenting only 8% of Mg substitution. This fact confers a more stable calcite skeleton to face ocean acidification then individuals from tropical environments. In addition, coralline algae from the Southwestern Atlantic Ocean are already living at temperatures that can be considered a limit for their survival. In fact, for cold water species, a subtle temperature increase could be beneficial in terms of their metabolism, photosynthesis and biomineralization.By the year 2100, surface seawater in all climatic zones could be undersaturated or at metastable equilibrium, with a high-Mg calcite phase containing ≥ 12 mol% Mg45. This could be catastrophic to coralline algae from the Southwest Atlantic Ocean, which produce CaCO3 crystals with more than 20% of Mg substitution in average as shown by the present study and for all the carbonate structures (e.g. rhodolith beds, coralline reefs, etc.) that depends on these skeletons to maintain and grow.It is worth to mention that coralline algae are present since the Mesozoic, in particular Sporolithaceans, which were already abundant in Cretaceous shallow waters49 and have already been submitted to bigger climate change events in the past, such as the Paleocene-Eocene Thermal Maximum (PETM), in which the deep-water temperature increased ∼5 ºC and a massive carbon cycle change took place with a large amount of CO2 absorbed by the oceans50. One of the possible explanations for the survival of coralline algae is that their biomineralogical control is limited to polymorph specification and would be ineffectual in the regulation of skeletal Mg incorporation51. In this sense, in past geological eras, such as the Cretaceous and Paleogene, the Mg/Ca ratio of the oceans favors the precitation of low Mg calcite29,52, which are more stable to dissolution. In a parallel to present day, other fundamental aspect we should take into account is the speed of progression of these changes. Actually, we know that the fast evolution of temperature and acidification present scenarios may result in significant impact on marine biodiversity and in marine calcium carbonate cycle players, as reef organisms and CCA.Carvalho et al.53 proposed that there would be a suitable area for rhodolith occurrence around 230,000 km2, providing a new magnitude to Brazilian Continental Shelf relevance as a major world biofactory of carbonate. In fact, this work confirms the estimation from previous studies, which indicated that this area would correspond to a 2 × 1011 tons of carbonate deposit of the Brazilian coast53. Among the most critical regions in the Brazilian coast, the Abrolhos Bank encompasses the largest continuous latitudinal rhodolith beds registered to date6, which is responsible for the production of approximately 0.025 Gt−1 year−1 of calcium carbonate, similar to those values reported for major tropical reef environments54,55. Another recently described important reef area on the Brazilian Shelf is an extensive carbonate system (≅ 9500 km2) off the Amazon River mouth56, which is composed of mesophotic carbonate reefs and rhodolith beds. These huge carbonate reservoirs and biodiversity hotspots may undergo a major decline if global ocean acidification and temperature rise take place in the near future. More

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Experimental designTo test whether an 8-year treatment of recurrent summer droughts would change biodiversity effects and species interactions of grassland plants when facing a subsequent drought event, we grew ambient- vs. drought-selected plants of 12 species in a glasshouse. The plants were grown from seeds collected from 40 plots (Supplementary Data 2) under 8-year treatments of yearly summer droughts vs. ambient precipitation in a biodiversity field experiment in Jena, Germany11,41.The Jena Experiment was established in 2002 using a common seed pool of 60 grassland species, with 80 (20times 20,{{{{{rm{m}}}}}}) large plots of species richness levels of 1, 2, 4, 8, 16, and 60 species40. Most of the species are perennial and capable of outcrossing (Supplementary Table 1). The Jena Drought Experiment11,41 was initiated in 2008. Two (1times 1,{{{{{rm{m}}}}}}) subplots were set within each large plot, designated as either drought treatment or ambient control. For the drought treatment, rainout shelters were set up to exclude natural rainfall in mid-summer for 6 weeks. The ambient control treatment got the same shelter construction but rain water was reapplied to not confound the results with artifacts from the shelter60. We repeatedly harvested the aboveground biomass per year, once before and once after the summer drought treatment11,41. The design of the Jena (Drought) Experiment did not allow the exclusion of cross pollination or gene flow between subplots or large plots in the field. Such gene flow may have reduced the possibility for genetic differentiation and for the observed effect sizes of the selection treatment23. We collected seeds from drought and control subplots throughout the 2016 growing season (Fig. 1). We obtained seeds of 17 species, but only used 12 of them, because the other five species had either few seeds or low germination rates. Seeds per species per selection treatment were collected from 4 to 23 (interquartile range: [8.50, 17.00]) maternal plants distributed across 2–10 (interquartile range: [4.75, 9.00]) large plots in Jena Experiment, in which the functional group richness ranged from 1 to 4 (Supplementary Data 2). The 12 plant species represented four functional groups (grass, small herb, tall herb, and legume) (Supplementary Table 1). The detailed classifications of the functional grouping can be found in the design of the Jena Experiment40. Eleven of the 12 species were perennial, and one was annual (Trifolium dubium). The average longevity of the perennial species in the Jena Experiment has been estimated at 3–4 years61, so that multiple generations and sexual reproduction cycles could occur during the 8-year drought treatment. Although each subplot was small, population sizes of each species were estimated to range from 100 to 1000 individuals m−2 in ambient and drought subplots at the beginning of the drought treatment in the field62.We germinated the seeds in Petri dishes and transplanted the seedlings into pots in February 2017 in a glasshouse (day temperature range 20–25 °C, night temperature range 15–21 °C, and humidity range 60–80%) at the University of Zurich, Switzerland. Seedlings were planted individually, in monocultures, or in 2-species mixtures in the glasshouse (Fig. 1). In the glasshouse experiment, both monocultures and mixtures contained four plants within a pot. The pots were (11times 11times 11.5) cm in size and filled with soil composed of 50% collected from a sugar-beet field, 25% sand and 25% perlite. We randomly assigned the pots into four blocks in the glasshouse. To test the effects of drought-induced selection on plant traits, we planted individual seedlings of the 12 species in a fifth block. Within the first 2 weeks, dead individuals were replaced, thereafter dead individuals were not replaced anymore. In total, we established 958 pots: 257 pots of mixtures, 217 pots of monocultures, and 484 pots of individual plants (244 pots of individuals in blocks 1–4, and 240 pots of individuals in block 5; Supplementary Methods). For mixtures, there were 21 species pairs (Supplementary Table 1). Species pairs composed of Crepis biennis or Lotus corniculatus had low numbers of replicates (Supplementary Table 1). However, including or excluding these communities produced qualitatively similar results. Thus, we present the results including these two species in this paper. We provide detailed explanations on the choices of species pairs and regarding the biodiversity treatment history in the Jena Experiment in Supplementary Methods.During a first phase of 3 months in the glasshouse (Fig. 1), pots were watered regularly (“before drought”). After 14–16 weeks, when most of the species had reached peak aboveground biomass, we harvested all individuals in each pot by cutting them 3 cm above the ground, allowing regrowth from the left plant bases (first harvest, “before drought”). The time span for the first harvest included both the time for trait measurements (section “Plant traits” below) and for the immediately following biomass harvest. We completed the biomass harvest of each block within 1–2 days. This allowed us to account for the larger time differences between blocks by fitting block effects in the statistical analyses. After the first harvest of each block, plants were watered regularly and allowed to regrow until the 18th week from planting. This was followed by a second phase of 2 weeks without watering. Soil moisture decreased from more than 40% to less than 10% after 10 days since drought initiation. At the end of the second phase, that is after 20 weeks from planting, we made a second aboveground harvest at 3 cm above the ground (second harvest, “during drought”). During a third phase of 7 weeks, pots were watered regularly again for recovery until most plants reached a new aboveground biomass peak again. At the end of the third phase, that is after 27 weeks from planting, we harvested both above- and belowground plant biomass (third harvest, “after drought”). We checked and confirmed that most plants had reached the full-grown state and peak biomass before each harvest by monitoring their flowering. After each harvest, we cleaned and dried the harvested plant material at 70 °C for 48 h to obtain the dry biomass. We used the aboveground biomass as a proxy for productivity. Although clipping may affect plant responses to the experimental drought in the glasshouse, clipping had the advantage that all plants were “standardized” in height before the experimental drought, thus reducing carry-over effects of differential growth before the experimental drought.Additive partitioningWe used the additive partitioning approach (Eq. 1)17 to decompose the net biodiversity effect (NE) on aboveground biomass into the complementarity effect (CE) and the sampling effect (SE):$$triangle Y={Y}_{O}-{Y}_{E}=N,overline{triangle {RY}},{bar{M}}+N,{{{{{{rm{cov}}}}}}}left({{triangle }}{{{{{bf{RY}}}}}},,{{{{{bf{M}}}}}}right),$$
(1)
where (triangle Y) is the NE; ({Y}_{O}) is the observed yield (productivity) in a mixture; ({Y}_{E}) is the expected yield in the mixture, calculated from the observed yield in monocultures and their corresponding species proportions planted in the mixture, here 0.5; the two additive terms at the right side of the equation represent CE and SE, respectively; N is the number of species in the mixture, here 2. The partitioning is based on the observed and expected relative yield (RY) of species in the mixture. The expected RY of species in the mixture is the proportion planted. ∆({{{{{bf{RY}}}}}}) is the difference between observed and expected RY of species in the mixture; (overline{triangle {RY}}) is the average of ∆({{{{{bf{RY}}}}}}). A positive (overline{triangle {RY}}) indicates a positive CE; a positive covariation between monoculture yield (M), and ∆({{{{{bf{RY}}}}}}) indicates a positive SE. More details about the calculation can be found in Loreau and Hector17. We conducted the partitioning separately for each harvest, selection treatment, and block. We did not perform the partitioning for mixtures with zero biomass63. For monocultures with zero biomass in the second or third harvest, we kept the ones which had positive biomass in the previous harvest but excluded the ones which had zero biomass in the previous harvest. For example, when performing the partitioning for the second harvest, we kept the monocultures that had zero biomass in the second harvest but non-zero biomass in the first harvest; we excluded the monocultures that had zero biomass already in the first harvest. This was to assure that communities that died before the drought could not reappear during or after the drought, and communities that had died during the drought could not reappear after the drought.We used mixed-effects models to assess the influences of drought vs. ambient-selection treatments on biodiversity effects (NEs, CEs, and SEs) separately for each harvest (Fig. 2; Table 1). Block and selection treatment were set as fixed-effects terms, while species composition (identity of species pair) and its interaction with selection treatment were set as random-effects terms. This conservative approach was used to allow for generalizations across all possible species compositions, although a more liberal approach with species composition and its interactions as fixed-effects terms could also have been applied (see Schmid et al.64 for a discussion of defining terms as fixed- vs. random-effects terms, including a justification of preference for treating block as a fixed-effects term). We square-root transformed the CEs and SEs with sign reconstruction (({{{{{{rm{sign}}}}}}}(y)sqrt{y})) prior to analysis to improve the normality of residuals17. The mixed-effects model did not converge in the analysis with CE after the drought event. In this case, we used a general linear model, in which we fitted block, species composition, selection treatment, and species composition by selection treatment interaction in this order. Then we tested the significance of selection treatment using its interaction with species composition as an error term. This procedure is an alternative to mixed-effects models that estimate variance components for random-effects terms with maximum likelihood64.To test whether biodiversity effects on productivity differed from zero, we additionally tested the significance of NEs, CEs, and SEs separately for each selection treatment and harvest (Supplementary Table 3). We set block and species composition as fixed- and random-effects terms, respectively. The model corresponding to CE for ambient-selected plants during the drought event did not converge so that we fitted it with a general linear model, in which we tested the significance of the overall mean (intercept) using species composition as an error term. All statistical analyses were conducted in R 3.6.365. The mixed-effects models were conducted with asreml-R package 4.1.0.11066.Finally, we also tested whether the effects of drought selection on biodiversity effects (NEs, CEs, and SEs) in the glasshouse depended on the history of biodiversity treatment in the Jena Experiment. Most plants in the 2-species communities in the glasshouse originated from mixtures in the Jena Experiment (Supplementary Data 2; whether mixtures in the glasshouse composed of plants originating from monoculture field plots did not affect the effects of drought-selection on biodiversity effects on productivity (Supplementary Data 3)). To increase statistical power, we used functional group richness, ranging from 1 to 4, instead of species richness of the field plots as explanatory variable (Supplementary Methods). We fitted functional group richness either in linear (Supplementary Data 4) or log-linear (Supplementary Data 5) form. We did not detect significant effects of field treatment of functional group richness nor significant interactions between field treatment of functional group richness and the drought-selection history. Therefore, we excluded the history of biodiversity treatments in the field from further analyses.Biomass stability to the drought event in the glasshouseTo assess the temporal responses of community aboveground biomass to the drought event, we calculated three indices representing different facets of stability: biomass resistance, recovery, and resilience (see van Moorsel et al.43 for an example). We calculated resistance as the biomass ratio during vs. before the drought, recovery as the ratio after vs. during the drought and resilience as the ratio after vs. before the drought (see also Isbell et al.9). We log-transformed the indices (plus a half of the minimum positive value to allow taking logs of indices that were originally zero) prior to statistical analyses to improve the normality of residuals. Excluding index values that were originally zero produced qualitatively similar results.To assess the effects of drought-selection on biomass stability, we fitted mixed-effects models with block and selection treatment as fixed-effects terms, and species composition and its interaction with selection treatment as random-effects terms (Supplementary Fig. 3; Supplementary Table 4). We fitted the models separately for mixtures and monocultures. We included the log-transformed biomass at the first harvest as a covariate because biomass stability in response to droughts often depends on plant performance under ambient conditions.In the same way as net biodiversity effects on productivity were calculated for additive partitioning, we calculated biodiversity effects on biomass stability as the difference between each mixture and its corresponding monocultures. Then, we tested the influence of selection treatment on the biodiversity effects on biomass stability. Block and selection treatment were set as fixed-effects terms; species composition and its interaction with selection treatment were set as random-effects terms (Fig. 3; Supplementary Table 5). The log-transformed biomass at the first harvest was also included as a covariate43. To assess the significance of biodiversity effects on biomass stability for each selection treatment, we fitted another set of simplified models, with block and log-transformed biomass as fixed-effects terms, and species composition as random-effects term (Fig. 3).Neighbor interactionsWe assessed interactions between neighboring plants within pots using the metrics of neighbor interaction intensity with multiplicative symmetry (NIntM)44:$${NIn}{t}_{M}=2frac{triangle P}{{P}_{-N}+{P}_{+N}+left|triangle Pright|},$$
(2)
where ({P}_{-N}) and ({P}_{+N}) are the productivities without (individual plant) and with neighbors (monocultures or mixtures), respectively; (triangle P={P}_{+N}-{P}_{-N}). Negative values of NIntM indicate competition and positive values indicate facilitation. NIntM is bounded between –1 (competitive exclusion) and 1 (“obligate” facilitation). For monocultures, we first calculated the per-plant biomass as the ratio between total biomass and planting density, and then used the per-plant value to compare with the corresponding individuals (without neighbor) of the same species with the same selection treatment in the same block. Note that under the reciprocal yield law45, an individual grown alone in a pot should be four times larger than an individual grown with three others in a pot, resulting in a NIntM of –0.75. For 2-species mixtures, we calculated the per-plant biomass separately for each species and took the average NIntM of the two species to measure the interaction intensity of the mixture. We set zero biomass for dead plants in the calculation. Again, if mixtures would also follow the reciprocal yield law independent of species identity, then NIntM = –0.75 would be expected. Values greater than –0.75 indicate some sort of overyielding due to higher density or higher density and higher diversity.To assess how selection treatment modified interactions between plants, we tested the effects of selection treatment on neighbor interaction intensity separately for monocultures and mixtures. We included block and selection treatment as fixed-effects terms, species composition and its interaction with selection treatment as random-effects terms (Supplementary Fig. 4; Supplementary Table 6).We calculated the difference between the heterospecific interaction in a mixture and the conspecific interactions in its two corresponding monocultures. A positive value of this difference indicates a weaker heterospecific than conspecific competition (i.e., niche differentiation) or stronger heterospecific than conspecific facilitation, which may lead to a positive complementarity effect. We tested the effects of selection treatment on interaction difference for each harvest by fitting block and selection treatment as fixed-effects terms, and species composition and its interaction with selection treatment as random-effects terms (Fig. 4; Supplementary Table 8). We also tested the significance of the interaction difference for each selection treatment by fitting block and species composition as fixed- and random-effects term, respectively (Fig. 4; Supplementary Table 7).Plant traitsTo assess whether drought selection would change plant traits, we measured six traits (Supplementary Table 9) closely related to plant usages of water or carbon on plants in pots with one individual from blocks 1–5. We focused on the traits on individual plants without neighbor to evaluate the influence of selection treatment on traits without the impacts of plasticity induced by plant interactions. We measured leaf relative chlorophyll content, leaf area (LA), leaf mass per area (LMA) and leaf osmometric pressure before the drought; leaf stomatal conductance both before and during the drought; and dry biomass ratio between root and shoot after the drought (in the third harvest). Leaf relative chlorophyll content was measured for three mature, fully expanded leaves per plant by using a SPAD-502 Plus chlorophyll meter from Konica Minolta. LA was obtained by scanning 3–4 mature, fully expanded leaves per plant with a LI-3100C Area Meter from LI-COR. LMA was calculated as the ratio between leaf dry mass (oven-dried at 70 °C for 48 h, using the same leaves that for LA) and LA. Leaf osmotic potential at full hydration was considered as an important trait associated with plant tolerance to drought30. We measured leaf osmotic potential with freeze-thaw leaf pieces cut from 1 to 2 mature, fully expanded leaves per plant by using a Wescor vapor pressure osmometer VAPRO (Model 5520) according to the method by Bartlett, et al.30. Plants were fully hydrated 1 day before the leaf sampling for osmotic potential measurement. Leaf stomatal conductance is a measure of exchange rate of carbon dioxide and water vapor through the stomata67. It was measured for 3–5 healthy mature leaves per plant by using a SC-1 Leaf Porometer from Decagon Devices. For grass species, 3 blades were placed adjacent to each other to have a large enough area for the measurement of stomatal conductance. For stomatal conductance during the drought event, we measured the individual plants from block 5 only due to limited time during the drought phase. We harvested aboveground and belowground plant biomass separately for alive individuals at the end of the experiment (after the complete recovery from the drought). The oven-dried (70 °C for 48 h) aboveground and belowground biomass were used to calculate the biomass ratio between root and shoot. We took the average value of each trait of each plant for statistical analyses. Each trait was measured for each block in turn.We used linear mixed-effects models to assess the influence (generalized across species) of selection treatment on trait values (red lines in Supplementary Figs. 5–7). Block and selection treatment were set as fixed-effects terms; species and its interaction with selection treatment were set as random-effects terms. Alternatively, we set species, selection treatment and their interaction as fixed-effects terms to assess whether species responded differently to the selection treatment (Supplementary Table 9). To test whether effects of selection treatment on traits differed between the five trait groups (leaf relative chlorophyll content, leaf area, leaf mass per area, leaf osmometric pressure, and leaf stomatal conductance) measured before the drought event in the glasshouse, we conducted two alternative analyses. First, we performed a principal component analysis with all traits and retained the first two principal axes (PC1 and PC2), which accounted for 39.06% and 22.3% of the total variation, respectively. Then we used PC1 and PC2 as response variables in mixed-effect models, separately. We fitted the models with the same fixed- and random-effects terms as those using each separate trait as the response variable. Effects of selection treatment on PC1 or PC2 were not significant. Second, we pooled the five traits as a single response variable in a mixed-effect model (corresponding to multivariate analysis of variance). Block, trait group (a factor with five levels), selection treatment, and the interaction between trait group and selection treatment were set as fixed-effects terms; species and its interactions with trait group and selection treatment and their three-way interaction were set as random-effects terms. We did not detect significant effects of selection treatment nor its interaction with trait group. Therefore, we did not present the results associated with these multivariate analyses in this paper. LMA, LA, leaf osmotic potential, leaf stomatal conductance, and root-shoot biomass ratio were log-transformed to improve normality of residuals.We also measured leaf relative chlorophyll content, LA and LMA in mixtures before the drought event (Supplementary Table 10) to evaluate the influence of selection treatment on trait dissimilarity between interacting species within communities. We calculated the absolute trait distance between two species in each mixture both separately for each trait and jointly with the three traits. For multi-trait-based dissimilarity, we standardized each trait to mean zero and unit standard deviation and calculated the Euclidean trait distance in standardized three-dimensional trait space.We used linear mixed-effects models to assess the effects of selection treatment on trait dissimilarity in mixtures (Supplementary Table 10). Block and selection treatment were set as fixed-effects terms; species composition and its interaction with selection treatment were set as random-effects terms. The model for LA dissimilarity did not converge so that we fit it with a general linear model, in which we tested the significance of selection treatment using its interaction with species composition as an error term. For the models with LA, LMA, and the joint three traits as dependent variables, we removed one pot (B1P674) because the LA value of Alopecurus pratensis in this pot was extremely small (about 1/3 of the second minimum value of the same species in mixtures). However, including or excluding this pot produced qualitatively similar results.Reporting summaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More

TRY meta-­analysisWe used the TRY plant trait database27 to identify traits that show systematic differences between the tree and liana growth forms, as a way to narrow the scope of the rest of the analysis. We chose traits to represent major trade­offs within the “economic spectrum” framework, which places plants along a spectrum of strategies from acquisitive, fast return on investment to conservative, slow return on investment according to key functional trait values30. We narrowed traits to those that had observations for at least four tree and liana species. We then compiled our dataset using the following steps during November and December 2019. For each trait, we downloaded the dataset for all species available globally and averaged the observations of the trait to the species level to avoid statistical biases introduced in our growth form comparison due to a high density of observations in a few commercially valuable species. We matched the species ID number with the most frequently used growth form identifier using the TRY “growth form” trait and kept the species with the most frequent identifier of “tree,” “liana,” or “woody vine.” We subsetted the data to keep only species with a majority of observations ascribed to the tree and liana growth forms (i.e., no herbaceous species, ferns, etc.), resulting in observations for 44,222 total species. Finally, we filtered the dataset of 44,222 species by hand to remove species misclassified as trees or lianas; species occurring entirely in temperate to boreal biomes; species from all gymnosperm lineages except the order Gnetales; and entries for taxonomic classifications broader than the genus level (e.g., taxonomic families). We found that hydraulic functional traits in the TRY database (i.e., Ks,max and P50) show systematic differences between growth forms (Supplementary Fig. 1; Supplementary Tables 3 and 4), while there is mixed evidence for differences in the acquisitiveness of trees and lianas in terms of stem anatomical traits (Supplementary Fig. 1; Supplementary Tables 3 and 4) and leaf functional traits (Supplementary Fig. 6; Supplementary Tables 3 and 4), and no evidence of differences between tropical trees and lianas with respect to root functional traits (Supplementary Fig. 7; Supplementary Tables 3 and 4).Extended meta­-analysisWe conducted an additional literature search to supplement the hydraulic trait observations from the TRY database. The additional literature search served two purposes: (1) to fill a major gap identified during our TRY analysis in terms of liana trait observations, and (2) to address the methodological inconsistency of measuring Ks,max and P50 on liana branches shorter than the longest vessel, which incorrectly measures Ks,max and P50 without accounting for end wall resistivity59,60.We conducted a literature search using Web of Science and Google Scholar. We searched the following phrases in combination with “liana:” “hydraulic conductivity,” “hydraulic trait,” “hydraulic efficiency,” and “hydraulic K.” Of the literature we found, we kept only the studies that met the following criteria: (1) reported Ks,max measurements for lianas, (2) measured Ks,max instead of computing Ks,max from xylem conduit dimensions, (3) measured Ks,max on sunlit, terminal branches of mature individuals or saplings, and (4) measured Ks,max on a branch longer than the longest vessel. We considered the authors to have used a branch length longer than maximum vessel length if the authors measured or reported maximum vessel length for the species and a longer branch was used. Because the best methodological practice for measuring P50, especially in species with long vessels, is currently a matter of debate, we additionally removed all observations of P50  > ­0.75. This filtering was performed to reduce the probability that falsely high (i.e., less negative) P50 values were retained in our analysis because of improper measurement technique and is consistent with the P50 filtering performed by Trugman et al.61. Improper measurement technique is a particular concern for lianas, whose wide and long vessels require cautious implementation of the traditional measurement techniques developed for trees. We note that retaining all liana P50 observations (i.e., not filtering out observations  > −0.75) results in a significant difference between trees and lianas (Mann­–Whitney test statistic = 1029, ntree = 61, nliana = 46, p  More

Chloroplast and repetitive nuclear DNA enrichment in the sedaDNA extractsTo the best of our knowledge, we generated the first large-scale target enriched dataset using sedaDNA extracted from sediments of multiple lakes. Sequencing of two datasets produced 325.5 million (M) quality-filtered paired-end DNA sequences. The first target enriched dataset, targeting both the chloroplast and a set of nuclear genes of Larix on 64 sedaDNA extracts and 19 negative controls from seven lake sediment records resulted in 324 M quality-filtered paired-end sequences. The second target enriched dataset, targeting only the set of nuclear genes of Larix on four samples and two negative controls from an additional lake (Lake CH12) resulted in 1.5 M sequences. Quality-filtering of an additional published target enriched dataset29, targeting the Larix chloroplast genome on the same CH12 samples as applied for the second dataset, added another 54 M sequences.For the chloroplast enrichment, 390 thousand (K) sequences (1%) were classified as Larix at the genus or species level. The average coverage of bait regions was 19% at a mean sequence depth of 0.8. Sequencing of 19 library and extraction blank (negative control) samples resulted in 597 K paired-end sequences, of which 58% quality-filtered and deduplicated sequences remained. Of these, 38% were classified, with 0.03% of them (463 sequences) corresponding to the genus Larix. Negative controls from library preparation resulted in no to very few (0 to 5) sequences mapping to the Larix chloroplast reference genome. Negative controls from DNA extractions, which were in several cases pooled to one library, showed a low number of sequences mapped to Larix (0 to 94 sequences, except 237 sequences in one case). Excluding all sequences in negative controls from the sample analysis had no impact on the patterns resulting from the analysis of sample data. Detailed results and evaluation of negative controls are included in the Supplementary Information (Fig. S5) and Supplementary Data 1 and 2. Samples of all lake records with sufficient sequence coverage showed damage patterns typical of ancient DNA (see Supplementary Data 3).These results are comparable to the results obtained by Schulte et al.29, where 36% of quality-filtered sequences were classified as Viridiplantae with 9% assigned to Larix. In contrast to29, we raised the confidence threshold of taxonomic classification (a parameter defining the number of k-mers needed to produce a match against a taxon in the database), which drastically reduced the number of classified sequences, but increased the confidence in the analysis36.To analyze the enrichment obtained by the nuclear gene bait set, taxonomic classification was repeated using a plant genome database including available Pinaceae genomes. The classification resulted in 716 K sequences assigned to Larix, increasing the previous results by 325 K sequences. However, almost no sequences were mapped against the targeting baits (a maximum of five sequences for some samples). A closer inspection of unmapped sequences assigned to Larix revealed a high content of repetitive DNAs. More specifically, taxonomically classified Larix sequences could be assembled to EulaSat1, the most abundant satellite repeat in the nuclear genome of Larix32,37. This short repeat with a 173 bp long motif is arranged in large arrays of tandemly repeated motifs and is exclusively present in larches32. Analysis of modern L. sibirica, and L. gmelinii (western and eastern range) genomes reveals that EulaSat1 occurs in all species, contributing to 0.62% (L. sibirica), 2.52% (western range L. gmelinii), and 2.39% (eastern range L. gmelinii), of the genomes, respectively (Fig. S2). A comparison of the sequence proportions mapping to the repeat motif in the different datasets of Lake CH12 showed a specific enrichment of the repeat motif by the nuclear gene hybridization probe set (Fig. S3).In total, 17 K sequences mapped to the repeat motif of EulaSat1. The abundance of all sequences mapped per sample is in agreement with the abundance of sequences mapped to the chloroplast genome, confirming the general history of forest development (Fig. 2). Analysis of the nucleotide frequencies in the repeat motif showed a high constancy over all samples (Fig. S4). This suggests high conservation of the EulaSat1 motif in Siberian larches over time and space. Although satellite repeats are reported to have a high sequence turnover, for larches it has been shown that repeat profiles between two geographically well-separated species—the European larch (L. decidua) and the Japanese larch (L. kaempferi)—are very similar32. The main satellite in all larches, EulaSat1, is believed to have greatly multiplied after the split of Larix from Pseudotsuga32. Given the ongoing hybridization between the three Siberian larch species, it is not surprising to find a consistent pattern of nucleotide frequencies in all samples.Fig. 2: Comparison of target enrichment with available DNA metabarcoding and pollen datasets.From left to right: Larix-classified sequence counts mapping to (1) the Larix chloroplast and (2) the EulaSat1 satellite repeat motif, (3) percentage of Larix counts in metabarcoding data, (4) percentage of Larix pollen in pollen assemblages. All data from this study, except metabarcoding data from lakes CH1213 and Bolshoye Shchuchye55 and all pollen data except for several samples of Lake Kyutyunda which were produced in this study56,57,71. Pollen data of Lake Lama and the Holocene part of Lake Kyutyunda are based on parallel sediment cores PG1111 and PG2022, respectively. No available data are marked with crosses, asterisk marks a single Larix pollen grain found in the Bolshoye Shchuchye sediments.Full size imageOff-target sequences in target enriched datasets have already been demonstrated to be useful for the analysis of high-copy DNA such as ribosomal DNA or plastomes34,38,39. A recent study on five modern sedges showed that target enriched sequencing data originally targeting a set of gene exons can also be used to study the repetitive sequence fraction and even infer phylogenetic relationships based on repetitive sequence abundance35. Another study showed that also sequence similarities between homologous repeat motifs can be used to reconstruct phylogenetic relationships among closely related taxa40,41. In the case of Larix satellite EuLaSat1 in our study, no change in nucleotide frequencies, neither related to locations nor in time, could be detected. However, our results show that the off-target fraction in target enriched sedaDNA datasets can hold valuable information and that repeat motifs in more diverse taxon groups could even be a target for enrichment. Specifically enriching for repeat motifs in sedaDNA extracts could enable the study of satellite repeat evolution as well as giving additional information on species abundance and phylogeography.In the two target enriched datasets, sequences taxonomically classified to the genus Larix and mapping to the chloroplast and to the repeat sequence, respectively, show similar patterns of abundance (see Fig. 2). Compared with published metabarcoding and pollen data from the same locations, the Larix abundance patterns can be globally reproduced, underpinning the notion that sequence abundances in target enriched data can be used as good estimates of plant abundances. For older parts of the lake records, target enriched data show Larix where metabarcoding data were unable to detect a clear signal (see Fig. 2, lakes Billyakh, Bolshoye Shchuchye, Kyutyunda, and Lama). This shows that target enrichment is superior to metabarcoding when analyzing one taxonomic group in-depth, as it is less prone to errors by DNA degradation, which can impede primer binding if the molecule becomes too short. Also, independent of age, rare taxa mostly need multiple PCR replicates to be detected by metabarcoding42,43. Target enrichment, however, is more sensitive in identifying one focal taxon group, as the total target length can be much larger (e.g., a complete organellar genome) than for metabarcoding, and the DNA damage patterns are put to use to authenticate ancient DNA. Also, it is limited by molecule length only by the applied threshold in the bioinformatic analysis, for which we used 30 base pairs (bp) as opposed to a minimum of 85 bp molecule length for the Larix metabarcoding marker (for the plant-specific trnL g/h marker44). Similarly, compared to traditional pollen analysis, target enrichment is more accurate at tracing a specific target group, as it is not dependent on pollen productivity. Especially in the case of Larix, pollen productivity is low and preservation poor, resulting in rare findings of its pollen in the sediments22,45. This could explain why for Lake Bolshoye Shchuchye, only a single Larix pollen grain was retrieved throughout the core, whereas target enrichment and metabarcoding show a strong signal in the Holocene sediments (last ~12 ka BP). Target-enriched data also records signals in MIS 2 sediments, however, sequence counts are extremely low, and as it is the only record, where both of the other proxies fail to report a signal, it should be interpreted with caution.A wider pre-glacial distribution of L. sibirica
Chloroplast genomes of L. gmelinii and L. sibirica differ at 157 positions, which can be used to differentiate species in target enriched sedaDNA29. Here, we applied this approach to lake sediment records, which are distributed across Siberia (Fig. 1) and have time ranges back to MIS3, and thereby were able to track species composition in space and time for wide parts of the species ranges.In lakes Billyakh and Kyutyunda, ca. 1500 km east of L. sibirica current range (Fig. 1), we found evidence for a wider distribution of L. sibirica around 32 and 34 ka BP in MIS3 (Fig. 3). Billyakh is situated in the western part of the Verkhoyansk Mountains, and Kyutyunda on the Central Siberian Plateau. Both lakes have low counts of Larix DNA sequences in their oldest samples dated to 51 ka BP (Billyakh) and 38 ka BP (Kyutyunda) with variants of L. gmelinii, but there is a sudden rise in variants attributed to L. sibirica at 34 ka BP (Billyakh) and 32 ka BP (Kyutyunda), which persists in the following samples, but strongly decreases in younger samples (Fig. 3). The rise in the L. sibirica DNA sequence variants coincides with a peak in sequence counts for Lake Kyutyunda. These signals suggest a rapid invasion of L. sibirica into the ranges of L. gmelinii in climatically favorable times and a local depletion or extinction of L. sibirica during the following harsher climates. Lake Billyakh pollen data suggest a moister and warmer climate around 50–30 ka BP than in the latter part of the Last Glacial associated with the MIS3 Interstadial in Siberia46.Fig. 3: Percentage and sequence counts at variable positions along Larix chloroplast genome assigned to species.Left: Alignment of Larix-classified DNA sequences against the chloroplast genome at the 157 variable positions between the species. For each position, the percentage of sequences assigned to a single species is displayed. Each row represents one sample named according to the calibrated age before present. Gray background indicates no coverage at the respective position. Right: Total number of sequences assigned to each of the species per sample.Full size imageStrong support for a wider pre-glacial distribution of L. sibirica comes from genetic analyses which show that it is genetically close to L. olgensis, today occurring on the Korean Peninsula and adjacent areas of China and Russia27,47. It is assumed that the L. sibirica-L. olgensis complex used to share a common range, which was disrupted and displaced when the better cold-adapted L. gmelinii expanded south and southwest during the more continental climatic conditions of the Pleistocene47,48. Furthermore, modern and ancient genetic studies suggest that the L. sibirica zone was recently invaded by L. gmelinii from the east in the hybridization zone of the species, as the climate cooled after the mid-Holocene thermal maximum13,23. Today, pure stands of L. sibirica do not form a continuous habitat, but occur in netted islands5 and morphological features of L. sibirica can be found in populations of L. gmelinii located at least a hundred kilometers east of the closest L. sibirica populations49. Macrofossil findings of L. sibirica in Scandinavia dated to the early Holocene, point to the capability of rapid long-distance jump dispersal of this species50. Fossil L. sibirica cones dated to the end of the Pliocene and in the Pleistocene have also been found far east of its current range in several river valleys including Kolyma, Aldan, and Omolon, and even in the basin of the Sea of Okhotsk9. These indicate long-distance seed dispersal by rivers which may also have assisted in successful establishment since the active-layer depth is deeper close to rivers51,52. As mentioned earlier, L. sibirica is sensitive to permafrost and waterlogged soils. A warmer phase with a deeper thawed layer above the permafrost could have enabled L. sibirica to spread and establish in regions that today are part of the geographic range of L. gmelinii, as L. sibirica is reported to have higher growth rates than L. gmelinii13.
Larix gmelinii formed northern LGM refugia across SiberiaThe possible survival of Larix in high latitude glacial refugia during the LGM is still under discussion4,53 although more and more evidence is reported in favor of the existence of such refugia17,20,21. The question of which of the Larix species formed these populations has hitherto been unanswered, as both pollen and established metabarcoding markers are not able to distinguish between species in the genus Larix, and findings of fossilized cones identifiable to species are rare. By enriching sedaDNA extracts for chloroplast genome sequences, we are, to the best of our knowledge, for the first time, able to distinguish between L. sibirica and L. gmelinii in glacial refugial populations.From Lake Lama, located at the western margin of the Putorana Plateau (Taymyr Peninsula), we obtained a continuous record extending from 23 ka BP to today with varying sequence counts with minima around 18–17 ka BP and 13 ka BP. All samples prior to the Holocene show variations predominantly assigned to L. gmelinii (Fig. 3). Our results suggest a local survival of L. gmelinii in the vicinity of Lake Lama throughout the LGM, which is supported by low numbers of Larix pollen detected through this period. Both target enriched sequence data and pollen indicate an increase from ca. 11 ka BP54. Sparse Larix pollen in the bottom part of the record could be an indication of a possible refugial population (Fig. 2; ref. 54).In Bolshoye Shchuchye, the westernmost lake of the study, situated in the Polar Ural Mountains, all Pleistocene samples show similarly a dominance of L. gmelinii sequence variations (Fig. 3). However, sequence counts for some samples are extremely low and samples from 18 and 10 ka BP had so low counts of mapped DNA sequences that none of the variable positions between the species was covered. Although sequences mapped to the satellite repeat of Larix also showed a Pleistocene signal, this was not repeated in pollen or metabarcoding (Fig. 2) which instead indicates a treeless arctic-alpine flora for the late Pleistocene55,56. Especially for the sample of 20.4 ka, Larix sequence counts are extremely low and new investigations would be needed to confirm a local presence of Larix during the LGM.The record of Lake Billyakh situated in the western Verkhoyansk Mountain Range, likewise shows extremely low counts of sequences mapped to the reference for a range of samples with no sequences covering the studied variable sites (45, 42, and 15 ka BP, 11–56 sequences mapped to non-variable sites). However, the pollen record for the same core shows a quasi-continuous record of Larix with a gap only occurring during the early LGM46 (25–22 ka BP, Fig. 2). Considering the known short-distance dispersal ability and poor preservation of Larix pollen, this strongly supports the presumed existence of a local glacial refugium at Lake Billyakh during that time20. Our samples also show a low but steady presence of Larix throughout the rest of the record, thus making glacial survival probable. The sample closest to the LGM (24 ka BP) indicates a clear dominance of L. gmelinii type variations.The only exception to this general pattern is the record from Lake Kyutyunda, which is located on the Central Siberian Plateau west of the Verkhoyansk Mountain Range. In this record, LGM samples have extremely low counts but show variations assigned to L. sibirica and not to L. gmelinii as in the other lakes. In addition, the preceding sample dated to the MIS3 interstadial shows L. sibirica variation. A possible explanation is that relics of L. sibirica survived during the LGM, but were unable to spread after climate warming, possibly due to genetic depletion or later local extinction. The presence of reworked sediment material can also not be excluded, as suggested by reworked pollen in the record57.In conclusion, our data show almost exclusively L. gmelinii variation for samples covering the most severe LGM climate conditions. This is in agreement with the ecological characteristics describing the species as adapted to extreme cold. In contrast to L. sibirica, it can grow in dwarf forms and propagate clonally and potentially survive thousands of years of adverse climatic conditions58.Postglacial colonization history—differences among larch speciesOf great interest in the Larix history is not only the location and extent of possible high latitude glacial refugia but also if and to what extent these refugia contributed to the recolonization of Siberia after the LGM. Northern refugial populations could have functioned as kernels of postglacial population spread and recolonization, or spreading could have been driven by populations that survived in southern refugia. There are only a few studies on modern populations that report evidence for possible recolonization scenarios of Larix23,27,28. Here, we show that patterns differ between L. sibirica and L. gmelinii.In the western part of our study region, two lakes are situated in the current distribution range of L. sibirica (Figs. 1, 4): Lake Bolshoye Shchuchye in the Polar Ural Mountains and Lake Lama on the Taymyr Peninsula. Despite this, both lakes show L. gmelinii for all Pleistocene samples, and a strong signal of L. sibirica variants only in the Holocene samples, with ages of 5.1 ka BP in Lake Bolshoye Shchuchye and 9.7 ka BP in Lake Lama (Fig. 3). The peak in L. sibirica also coincides with a peak of sequence counts in the respective sample, with a Larix pollen peak in Lake Lama sediments54, and metabarcoding for Lake Bolshoye Shchuchye55. This points to a migration of L. sibirica in its current northern area of northern distribution in the course of climate warming during the early Holocene, whereas glacial refugial populations were consisting of L. gmelinii. Although the local survival of L. gmelinii around Lake Bolshoye Shchuchye remains uncertain due to extremely low sequence counts, it is clear that L. sibirica did not form a refugial population at this site.Fig. 4: Percentage of DNA sequences assigned to references displayed on the geographical locations of the lakes investigated.Samples in the same time frame are averaged. Lake names and current species ranges are annotated in the middle plots. Colors indicate current species distribution (adapted from Semerikov and Lascoux72). The base map is done with ggmap73, map tiles by Stamen Design, under CC BY 3.0. Data by OpenStreetMap, under ODbL.Full size imageA range-wide genetic study of L. sibirica analyzing chlorotypes and mitotypes of individuals23 found strong indications for rapid colonization of the West Siberian Plains from populations originating from the foothills of the Sayan Mountains in the south, close to the border of Mongolia, with only limited contribution from local populations. According to our results, the local populations could have been L. gmelinii populations, while the rapid invasion could have been L. sibirica.In the eastern range of the study region, in the current range of L. gmelinii, namely at lakes Emanda, Satagay, and Malaya Chabyda, genetic variations throughout the records are less pronounced. Of the three lake records, only that from Lake Emanda reaches back beyond the LGM, but with a sampling gap for the time of the LGM. Therefore, it remains uncertain whether populations survived the LGM locally, or whether they were invaded or replaced by populations coming from the south with Holocene warming. The restricted variations throughout the record, however, hint at stable populations, which is supported by scarce pollen findings (Fig. 2).Our data suggest that postglacial recolonization of L. sibirica was not started from high latitude glacial refugia, but from southern populations. In contrast, northern glacial populations of L. gmelinii could have potentially enhanced rapid dispersal after the LGM in their current area of distribution.Environment likely plays a more important role than historical factorsThe current boundaries of boreal Larix species arranged from west to east suggest a possible strong influence of the historical species distribution on the current distribution, whereas the gradient of increasing continental climate towards the east assumes a strong influence on the environment. By tracking species distribution in the past, spanning the time of the strongly adverse climate of the LGM, we can give hitherto unprecedented insights into species distribution history.Several lines of evidence suggest a strong influence of the environment on species distribution: (1) Signals for L. sibirica appeared in its current area of distribution as late as the Holocene warming, whereas cold Pleistocene samples are dominated by L. gmelinii type variation; (2) in lakes far east of its modern range, signals of variation typical for L. sibirica coincide with peaks in sequence counts (29 ka BP, Lake Billyakh; 32 ka BP Lake Kyutyunda), which point to more forested vegetation around the lakes and consequently a more favorable climate at that time; and (3) samples dated to the severely cold LGM are dominated by variations of the L. gmelinii type.This is in accordance with the different ecological characteristics described for the species. L. sibirica is sensitive to permafrost and only occurs outside of the zone of continuous permafrost5. In addition, L. sibirica achieves substantially higher growth rates and longer growth periods than L. gmelinii9,13 and can also produce more than twice as many seeds5. This potentially gives L. sibirica the ability to quickly react to climate change and outcompete the other species when the climate becomes more favorable.In contrast, L. gmelinii is adapted to extremely low soil and air temperatures and is able to grow on permafrost with very shallow thaw depths. It’s distribution almost completely coincides with continuous permafrost5, and even a restriction to permafrost areas is discussed as it does not grow well in field trials on warmer soils or where there is a small temperature gradient between air and soil9. Due to this ecology, L. gmelinii is more likely to survive in a high latitude refugium, even during the severe continental climate of the LGM, which was most probably connected to continuous permafrost of low active-layer depths.A study combining mitochondrial barcoding on sedaDNA and a modeling approach on Larix distribution in the Taymyr region around Lake CH12 concluded that the distributions of L. gmelinii and L. sibirica are most strongly influenced by stand density and thus by competition between the species, with L. gmelinii outcompeting L. sibirica at high stand densities13. As our study includes sediment cores reaching further back in time, we see a different trend. Instead of L. gmelinii, it was L. sibirica, which dominated samples with high sequence counts, suggesting high stand density and a more favorable climate. A possible explanation for the different outcomes is the use of different organelle genomes. Epp et al.13 used a marker representing the mitochondrial genome, which is known to introgress more rapidly and as a consequence might show a long past species history59,60.Our findings have potentially important implications for the projections of vegetation-climate feedback. A warming climate in conjunction with a greater permafrost thaw depth could enable the replacement of L. gmelinii by L. sibirica. In contrast to L. gmelinii, L. sibirica is not known to stabilize permafrost thus potentially further promoting permafrost thaw and with it the release of greenhouse gases, creating positive feedback on global warming11. On the other hand, the substantially higher growth rates of L. sibirica in comparison to L. gmelinii would increase carbon sequestration, thus mitigating global warming13. This shows the importance of understanding species-specific reactions to climate change, which can result in great shifts in distribution. Target enrichment applied on sedaDNA is able to reveal the impact of past climate change on populations and the increasing availability of modern reference genomes will further enhance its value of information. More