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arising from J. Cheng et al. Nature Communications https://doi.org/10.1038/s41467-021-22087-2 (2021)Recently, Cheng J. et al.1 cited and simulated the relative percentage tree cover to interpret the ~3000–4000 years lag between tree cover and East Asian Summer Monsoon (EASM) rainfall. They concluded that vegetation feedback has caused a lagged ecosystem response to EASM rainfall during the Holocene (11.7–0 ka). Here, we question the feasibility of using the relative percentage tree cover to measure vegetation feedback to climate. First, the land cover in northern China includes forests, grasslands, and bare land2. Cheng J. et al.1 did not consider the role of bare land in climate feedback models. Absolute land cover, including forest, grassland, and bare land can accurately reveal feedback to climate3. Second, the biome reconstructions they cited represent changes in vegetation type only, whereas the relationship between vegetation type and vegetation cover is altered by many other factors3,4,5. Third, the paper they cited6 averaged the vegetation types on a millennium scale with an interval of 1000 years, so the view that vegetation has a ~3000–4000 years lag in EASM rainfall is not credible as the lag can be enlarged by data resolution. Therefore, absolute vegetation cover, not relative cover, is a prerequisite for studying ecosystem feedback.Our previous work3 was the first to reconstruct the absolute vegetation cover in northern China based on pollen concentrations in two well-dated sediment cores. Using a random forest method, the vegetation cover at Dali Lake in the forest-steppe transition in northern China was determined for the period from 19,000 cal. yr BP to the present with a resolution of approximately 200 years. Han et al.3 showed that tree cover peaked during the early Holocene and it has gradually declined since the middle Holocene. Pollen percentages are widely used in vegetation reconstruction, but they are challenging to analyze because they can be similar in composition, despite being produced by very different flora7,8. This means they represent the relative fractions of vegetation type and not the absolute vegetation cover (Fig. 1A, B). Han et al.3 suggested that pollen concentration data are suitable for the reconstruction of absolute vegetation cover, particularly in arid and semi-arid regions. Dali Lake represents a typical lake in the semi-arid area of northern China. The random forest model showed that the area had a high tree cover during the early Holocene, which suggests that tree cover was a timely response to Holocene monsoon rainfall and there was no time lag at this specific location. Therefore, the data at Dali Lake challenge the conclusion that tree cover has a ~3000–4000 years lag to EASM rainfall.Fig. 1: Trends in pollen percentage, absolute vegetation cover and fire history during the Holocene at Dali Lake, a typical lake in the semi-arid area of northern China.Changes in the percentages of arboreal and non-arboreal pollen (A)3. Reconstructed absolute tree and grass cover (B)3, with gray, yellow, and green shaded areas indicating the standard deviation of 1000 random forest model results for total cover, grass cover, and tree cover, respectively. Normalized fire activity index in the northern region of eastern monsoonal China (C)10. Z-score of transformed charcoal value showing fire activity trend in the temperate steppe of northern China (D)11.Full size imageMoreover, Cheng J. et al.1 used −17 °C as the threshold for tree and grass transition, but this could be an incorrect citation from Bonan et al.9. In the original text by Bonan et al.9, they showed that both temperate deciduous broadleaved trees and C3 grasses have a tolerance of −17 °C for the coldest month for their survival, but this temperature is not the favored threshold for the shift from grasses to trees. Actually, the trend in absolute vegetation cover was mainly driven by summer temperature, annual precipitation, and fire incidents, which is in line with the vegetation-climate relationships at Dali Lake3. That is, higher monsoon rainfall could increase the competitiveness of trees, while increased fire could increase the competitiveness of grasses as grasses are mostly annual and perennial, and they renew faster than trees after a fire. Between 10,000 and 8000 cal. yr BP, monsoon rainfall peaked and there were relatively few fires, which led to a significant increase in absolute tree cover. Since 6500 cal. yr BP, monsoon rainfall decreased and fire increased, resulting in stronger competition by grasses, which has led to an increased grass cover and reduced tree cover3 (Fig. 1B).The impact of secondary disturbances on vegetation dynamics requires careful consideration, particularly the impact of fires on vegetation cover in semi-arid areas of China, even though vegetation growth is strongly constrained by rainfall in this region. Both the normalized fire index in the northern region of eastern monsoonal China10 and the charcoal value in the temperate steppe of northern China11 show a clear antiphase relationship with the absolute forest cover of the Dali Lake region (Fig. 1B, C, D). However, Cheng J. et al.1 did not discuss fire incidence on vegetation evolution in northern China. Fire has occurred frequently through the Holocene10,12 and it plays an important role in vegetation dynamics based on observational evidence13,14. Particularly, fire is considered as a triggering disturbance that can reduce a forest’s resilience to drought under a drying climate during the mid to late Holocene. For Dali Lake, fire and drying climate has co-driven the evolution of vegetation cover since 6500 cal. yr BP3. Moreover, in northern China, data from Daihai Lake and Hulun Nuur Lake also suggest that fire has accelerated the decline of forest cover and the transition from forest to grass during the Holocene13,14. Unfortunately, Cheng J. et al.1 did not discuss the effect of fire when citing and interpreting the observation data, as scale-dependent fires could make the transition between forest and grassland and their interactions variable.In summary, we believe there are three flaws in the data interpretation of Cheng J. et al.1. (1) They contradict the observed evolution of absolute tree cover at Dali Lake in northern China, which is representative of the marginal area of EASM. Relative percentage tree cover cannot accurately reflect a forest’s response and feedback to past climate change. (2) Both temperate deciduous broadleaved trees and C3 grasses have a tolerance of −17 °C for the coldest month for their survival. Thus, −17 °C is not a correct threshold for the shift from grasses to trees. (3) They contradict the ecological theories of secondary disturbance on vegetation dynamics. Fire and other secondary disturbances may be crucial to the transition between forest and grassland. Interpretations of vegetation feedback might be biased if these important factors are not fully considered. Based on the evidence, their conclusion that vegetation feedback causes lagged ecosystem response to EASM rainfall during the Holocene could be problematic. More

Elderflora: A Modern History of Ancient Trees Jared Farmer Basic (2022)About 45 million years ago, when the Arctic was ice-free, the world’s earliest known mummified trees flourished on what is now Axel Heiberg Island in Canada’s Qikiqtaaluk Region. In 1986, palaeobotanists identified the megaflora as members of Metasequoia occidentalis, an extinct redwood species. They had been buried in silt, then frozen, their wood preserved.The lead palaeontologist “celebrated his eureka by kindling a fire with 45-million-year-old twigs and boiling water for tea time,” writes historian Jared Farmer in Elderflora, his expansive global history of grand and venerable trees. Granted, these plants had been dead since the Eocene epoch. Nevertheless, as the author describes, the incident is part of a troubling pattern in which scientists rejoice at their discovery of the ‘oldest’ tree of their time — and then destroy it.In 1957, for example, Edmund Schulman at the University of Arizona in Tucson spent the summer seeking ancient bristlecone pines in California’s White Mountains. He found three more than 4,000 years old, and named them Alpha, Beta and Gamma. Then, in the interests of tree-ring science, he chose to “sacrifice” Alpha, taking snapshots as his nephew and a colleague sawed it down. When the University of Arizona issued a press release titled ‘UA Finds Oldest Living Thing’, Farmer writes, “they say nothing about the thing being dead”.Schulman’s aim was dendroclimatology — the reconstruction of climates using tree-ring data. That lofty motive cannot be ascribed to those who, in 1881, bored a tunnel into the 2,000-year-old Wawona tree in Yosemite National Park, allowing tourists to drive their cars through the 71.3-metre-high giant sequoia (Sequoiadendron giganteum), since toppled.Arboreal legendsAs Elderflora shows, big, old trees are objects of veneration and vandalism, appearing “in the oldest surviving mythologies and the earliest extant texts”. They were associated with gods and heroes, prophets and gurus: they had pivotal roles in the Mesopotamian Epic of Gilgamesh and in the Polynesian legend of Rātā, who fells a noble tree to carve a canoe. In more recent times, European settlers “dispossessed Indigenous peoples and cleared forests with abandon”. Research shows that, for 8,000 years after the glaciers of the last ice age retreated, forests in the Midwestern United States doubled in biomass (A. M. Raiho et al. Science 376, 1491–1495; 2022). Just 150 years of industrial logging and agriculture erased this carbon accumulation.
It takes a wood to raise a tree: a memoir
“Imperial conquests and industrial revolutions relied on timber,” Farmer writes. “Wood-stock long guns for capturing lands and peoples; naval vessels with mighty masts for transporting the enslaved and the harvests of their labor.” In New Zealand, European settlers decimated the majestic kauri trees, which can live for up to 2,000 years and that once covered 1.2 million hectares of land. The trees’ 50-metre-trunks became ships’ masts; their resin was made into varnish and linoleum.Like pines, firs, spruces, cedars, cypresses and redwoods, kauri (Agathis australis) is a gymnosperm. These flowerless plants with naked seeds tend to grow slower and live longer than angiosperms, flowering plants that bear fruit. About 25 plant species — most of them conifers — can live for more than a millennium without human assistance, surviving in restricted, vulnerable habitats.Farmer also offers a global survey of ancient trees that have been protected and exalted. They include olive trees of the Levant (Olea europaea); research published this year shows that these were domesticated about 7,000 years ago for their fruit and oil (D. Langgut and Y. Garfinkel Sci. Rep. 12, 7463; 2022). In Africa, the baobab (Adansonia sp.) is both the longest-lived tree and the largest, offering shade and shelter, foods, medicines and textiles. Enslaved Africans planted baobabs in the Caribbean; some survive still. Ginkgo biloba, a species that dates back 390,000 years, survived only in China, whence it was spread around the world in the past millennium. A grove of ginkgo trees survived the atomic bombing of Hiroshima in Japan in August 1945, pushing out new buds the following spring.The planet’s current tree cover, Farmer writes, includes 3 trillion large plants covering about 30% of all land. It is, in fact, expanding. But the new cover consists mostly of shelter belts (trees planted to protect crops or animals), temperate-zone timber crops and tropical plantations of eucalyptus and palm oil. A shrinking proportion of tree cover is made up of species-rich old-growth communities.Epic loss“What would humans and nonhumans stand to lose if these survivors all died prematurely? A world of things,” Farmer writes. “Old trees sustain forest communities” with their seeds and litter. Other plants grow on them, and animals live in them. Their roots share nutrients with other organisms via underground fungi. Groups of “Old Ones” are carbon sinks. Large-scale monocultures are shorter-lived and take less greenhouse gas out of circulation.But even bygone trees of the once-tropical Arctic might offer lessons for a warming world. Palaeobotanist Hope Jahren, in her 2016 memoir Lab Girl, describes how she spent three summers on Axel Heiberg Island, digging “through a hundred vertical feet of time”. Fir, cypress, larch, redwood, spruce, pine and hemlock trees populated this lush conifer forest, with an understory of angiosperms: maple, alder, birch, hickory, chestnut, beech, ash, holly, walnut, sweetgum, sycamore, oak, willow and elm. These plants thrived even through three months of winter darkness and three of constant summer light.“Here stood one of the great forests of all time,” Farmer writes. Today, as the Arctic warms nearly four times as fast as any other place on Earth, the genomes of species related to the trees of this mummified forest might be adaptable enough for the trees to flourish in a rewarmed planet, he says. Old trees have much to teach us: we would be wise to listen. More

replying to Y. Cheng et al. Nature Communications https://doi.org/10.1038/s41467-022-33958-7 (2022)We welcome the comment from Matters arising from Cheng Y et al.1, which provides us an opportunity for further clarification of some of our points2. The Comment raised interesting and important issues about our paper, that undoubtably could enhance our understanding for the Holocene vegetation evolution in the northern China and its relationship with East Asian Summer Monsoon (EASM). In particular, the results from Dali lake pose the questions on the timing of the peak of tree cover, that invokes the further investigation to understand this complex tree changes over Holocene period. However, these comments do not impact the key result in our original study2, that is the periodical asynchronous evolutions between EASM and northern China ecosystem under specific conditions.Main points of our paper are: First, we propose that the EASM and its rainfall over northern China mainly followed the variation of the summer insolation and peaked in the early Holocene, while the relative tree cover of temperate deciduous broadleaf tree peaked in the mid-Holocene; the delayed tree cover peak is caused by the winter warming, and peak soil moisture also in the mid-Holocene, which could be related to a hydrological impact from vegetation shift from grass to tree and the positive feedback between this vegetation shift and soil wetting.Second, this asynchronous evolution between the EASM rainfall, which peaks in the early Holocene, and the northern China ecosystem, which peaks in the mid-Holocene, is caused mainly by the opposing effect of residual ice sheet retreat on the decreasing summer insolation. The declining summer insolation does cause a substantial decrease of EASM rainfall from the early to mid-Holocene. However, 2/3 of this rainfall decrease is canceled by the rainfall increase forced by the retreat of residual Laurentide ice sheet, resulting in a weak decreasing trend of rainfall over this period.Third, under this background of weak rainfall changes, winter warming, induced by increased winter insolation and ice sheet retreat, raised the coldest temperature to above −17 °C, the threshold for the survival of temperate deciduous broadleaf tree3, and then favored an increase in tree, meanwhile induced a decrease in grass for reasons of its lower competitiveness than that of tree. This vegetation shift then supported the wetting of northern China through its hydrological effect2. The vegetation shift and soil wetting could reinforce each other. Furthermore, the dominant effect of winter warming on vegetation from the early to mid-Holocene is supported by our sensitive experiments with an off-line land-vegetation model.As stated in Cheng Y’s Comment1, the land cover in northern China includes forests, grass and bare land. In our interpretation, the process of “the vegetation feedback to climate” is mentioned as a possible feedback that enhances this asynchronous response, but is not critically involved in the mechanism. As such, whether the absolute or relative vegetation cover is not a major issue in our discussion. It’s sure that the reconstructed absolute tree cover, which based on pollen concentration, could enrich our understanding of the vegetation changes over the Holocene period in northern China. Indeed, the hydrological impact of bare land (evaporation) had been considered in our hydrological analysis of northern China soil moisture, and the results indicated its impact is important but not critical to the Holocene long-term changes of soil moisture over this region. The relative tree cover, the percentage of cool mixed tree (COMX4) in fossil pollen which is consistent with that of temperate deciduous broadleaf tree in simulation2, that we cited4 is a synthesis of 31 records, which represented the general evolution of vegetation over a large part of northern China, and its main result is consistent with records from other part of northern China such as the 6 ka peak in Gonghai Lake5. In spite of its low time resolution, the general trend over the millennium scale seems to us clear.It’s true that the −17 °C of the coldest month temperature is the survival threshold for the temperate deciduous broadleaved tree. While, the temperature threshold for C3 grass and C3 arctic grass are complex, its direct impact on the changes of grass proposed in our paper is somewhat not strict. However, considering the different competitiveness between tree and grass, increased temperate deciduous broadleaved tree, which derived by the winter warming, could induce a decrease in the grass from the early to mid-Holocene. Indeed, summer temperature, annual rainfall and fire incidents are all the important factors determining the Holocene changes of vegetation over northern China, but series of sensitivity experiment proposed the key impact of winter temperature on the vegetation shift and soil moisture evolution, which is consistent with the results of transient coupled climate simulation and geological records. This grass-to-tree shift for this period is evidenced in the pollen percentages and well simulated by the climate model shown in our paper2.Fire is an important factor for the long-term changes of vegetation cover over semi-arid regions, and its emergence and impact on vegetation are already incorporated into our model6, then, in turn, the simulation. Future works could assess the impact of fire on the long-term changes of semi-arid vegetation through the combination of reconstruction and process-based simulation of fire7.Focusing on the contrary views of Holocene EASM within proxy records, we proposed an asynchronous evolution of EASM rainfall and northern China ecosystem for the period of early to mid-Holocene. Our proposal is based on a state-of-the-art transient climate simulation, which reproduced the diverse evolution of EASM proxies reasonably well. The mechanisms proposed for this asynchronous evolution appear to us consistent with the current evidences available. There are, however, uncertainties in models and proxies. Meanwhile, the northern China is a broad region with large gradient in rainfall and ecosystem, that could induce the possible diverse evolutions in climate and ecosystem under Holocene climate change. Therefore, we believe further studies using other models and new proxies are important to further improve our understanding of this issue. More

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