{"title":"鹿与虎,森林与碳","authors":"Oswald J. Schmitz","doi":"10.1111/gcb.70198","DOIUrl":null,"url":null,"abstract":"<p>Healthy, verdant forest ecosystems are indicated by an intact complement of highly abundant vegetation along with diverse and abundant populations of herbivore species and their carnivore predators (Wang et al. <span>2025</span>). Keeping forest ecosystems verdant is considered vital to ensuring planetary resilience to climate change (Watson et al. <span>2018</span>). This is because the highly abundant vegetation takes up atmospheric carbon that is then stored in vegetation biomass and in soils of those ecosystems (Pan et al. <span>2024</span>).</p><p>But what would happen to climate change resilience if forest ecosystem intactness was disrupted by, say, the loss of predators? The answer depends on how intactness is sustained. Ecological science has two general views on this (Schmitz et al. <span>2018</span>). One—the bottom-up control view—holds that ecosystems are verdant because vegetation abundance, which is strongly determined by soil nutrient and moisture levels, supports but limits the abundance of herbivore populations, and in turn, populations of their carnivore predators. In this view, climate change resilience would not be disrupted by predator losses (or herbivores for that matter) because animal abundance and diversity do not drive vegetation abundance. The other—the top-down view—holds that predators, by virtue of limiting the abundance of their herbivore prey, keep ecosystems verdant by preventing herbivore overexploitation of vegetation. In this view, climate change resilience would be disrupted by the loss of predators because they ultimately drive vegetation abundance. An important and challenging research frontier for both ecological and global change science is resolving which view of control best explains the climate resilience of verdant forest ecosystems, especially across vast landscapes over which large animals live and roam (Schmitz et al. <span>2018</span>). This challenge is compounded by variation in biophysical conditions across those vast landscapes because the amount of carbon captured and stored among geographic locations becomes highly dependent upon biophysical context, including climatic conditions, the kinds, diversity, and abundances of plant and animal species, and the nutrient contents and physical properties of soils (e.g., Sobral et al. <span>2017</span>; Schmitz et al. <span>2018</span>; Schuldt et al. <span>2023</span>).</p><p>In newly published research in <i>Global Change Biology</i>, Roberts et al. (<span>2025</span>) address this formidable challenge to reveal how varying abundance and outright loss of a large predator—the tiger (<i>Panthera tigris</i>)<i>—</i>and the abundance of its “deer” (i.e., ungulate) prey species are related to the capture and storage of carbon in forests across the tiger's geographic range throughout Asia. This vast landscape has myriad biophysical dimensions that create much context dependency. It contains four broadly different forest ecosystem types, including boreal, temperate, subtropical dry, and subtropical moist. The ecosystems are arrayed across different elevations from lowland to montane and across different wet to dry regimes. They harbor many varieties and abundances of plant and herbivore prey species.</p><p>Resolving the role of tigers in forest climate resilience requires comparing carbon dynamics in places where tigers are present with similar environmental conditions where they are absent (Schmitz et al. <span>2018</span>). The hallmark scientific approach would be to conduct a manipulative Case (tiger-present target forest)–Control (tiger-empty forest) experiment that pairs biophysical conditions of forest locations and systematically excludes tigers from half of them, replicated across the different forest ecosystem contexts across the vast landscape (Schmitz et al. <span>2018</span>). This is altogether logistically impossible. Roberts et al. (<span>2025</span>) overcame this critical limitation with the next best approach (Schmitz et al. <span>2018</span>): they amassed a large multivariate data set gathered from satellite remote sensing and extensive on-the-ground sampling across landscape locations where tigers have been persistently present versus similar landscape conditions where they are absent—their loss arising from known historical extirpation by humans. The amassed data included measures of carbon in vegetation biomass and soil, net ecosystem carbon exchange, and aspects of context dependency including forest ecosystem type, regional climate, and human disturbance. These are exactly the kinds of data used in the conventional accounting of carbon capture and storage in forest ecosystems (Pan et al. <span>2024</span>). As well, they accounted for context dependency in top-down control due to variation in deer diversity and abundance, which according to classical theory (Polis and Strong <span>1996</span>) should be stronger in more linear food chains (i.e., deer diversity is low) and dissipate along the branching network of linkages in food webs with higher prey diversity. There was sufficient replication within and among forest ecosystem types to control for underlying spatial context dependency in these biophysical factors in both tiger present and absent sites. Hence, Roberts et al. (<span>2025</span>) were able to undertake extensive direct and counterfactual analyses to fully and robustly interrogate the veracity of the top-down view that tigers play a role in controlling forest carbon dynamics across their geographic range.</p><p>The short answer to the question about the consequences of predator loss is that it mattered. Tiger presence was generally linked to higher vegetation carbon stocks and higher net carbon exchange (i.e., forest carbon uptake exceeded carbon emissions) among forest types. But the longer answer is that there was also nuance due to context-dependency. Certainly, bottom-up control prevailed across all forests due to geographic variation in biophysical properties that created variation in vegetation biomass and carbon exchange and storage among the forest ecosystems. But top-down effects of tiger presence also prevailed, resulting in higher vegetation and soil carbon stocks among most contexts than in the absence of tiger, the exceptions being tropical dry forest and forests in lowlands and especially wetlands. These exceptions extended to net ecosystem carbon exchange, with one context (tropical swamp forest) even switching from being a slight net carbon sink to a carbon source in the presence of tigers. Tiger presence was impactful in forests with low to intermediate vegetation biomass but not high biomass, suggesting that the biomass dense forests might be completely bottom-up controlled. Yet prey diversity also seems to have played a hand by mediating tiger effects. Forests with low to intermediate vegetation biomass had less deer biomass and diversity than did high vegetation biomass forests. Hence, a second reason for weak, if any, tiger control in biomass dense forests is that the top-down effects may have dissipated along the many food web linkages, suggesting again that both top-down and bottom-up control persisted across contexts but with different relative importance.</p><p>In <span>1967</span> wildlife ecologist George Schaller published a seminal treatise <i>The Deer and Tiger</i> in which he provided foundational scientific understanding about the ecological interplay between tigers and their key prey species. It further described how those relationships were being eroded due to rural land use change for agriculture, persecution of tigers out of fear for human safety and loss of wellbeing, and exploitation for body parts that had perceived medicinal value—problems which continue to this day (Roberts et al. <span>2025</span>). Schaller's book was published at a time when ecological science had barely started to imagine that predator–prey dynamics could cascade to impact the properties and functions of entire ecological communities, let alone the kinds of ecosystem processes such as the carbon cycle that also have implications for human wellbeing. In retrospect it should have been a portent of ramifying ecosystem impacts. But at the time the scientific community was ill-equipped conceptually to even recognize this as a bigger issue because it had yet to embrace a holistic perspective on what it means to sustain verdant forest ecosystems. The portent is still unheeded in many areas of climate change science today, even while scientific understanding of the role of animals in controlling the carbon cycle has advanced considerably (Schmitz et al. <span>2018</span>). By expanding the scientific story of the Deer and the Tiger to the story of the Forest and the Carbon, Roberts et al. (<span>2025</span>) have provided a leap in scientific understanding about how broadly impactful an animal species can be.</p><p>These findings are sure to excite those working on the frontlines of tackling biodiversity loss and climate change by showing how both looming problems can begin to be solved together. There is, however, a risk that arguments about the carbon benefits of tigers will be used to justify headlong action to restore this iconic and endangered species everywhere across its geographic range. Here the deeper, and critical, lesson of the Roberts et al. (<span>2025</span>) article is that by deliberately addressing context-dependency it provides the kind of sober analysis that has been recently called for to temper policy and conservation from overpromoting animal restoration and conservation as a universal win-win for mitigating biodiversity loss and climate change together (Burak et al. <span>2024</span>). The science is clear that tigers can have impactful effects on forest ecosystem carbon storage. But most importantly, Roberts et al. (<span>2025</span>) have provided unprecedented scientific insight into which forest ecosystems across the tiger's vast geographic range should be considered candidates for tiger population conservation and restoration for carbon capture and storage; and which forests definitely should not. The research is exemplary for showing how to advance an evidence-based approach that can be applied in the service of restoring and sustaining verdant forest ecosystems for the purpose of maintaining resilience to climate change.</p><p>The author declares no conflicts of interest.</p><p>This article is a Invited Commentary on Guangshun Jiang et al., https://doi.org/10.1111/gcb.70191.</p>","PeriodicalId":175,"journal":{"name":"Global Change Biology","volume":"31 5","pages":""},"PeriodicalIF":10.8000,"publicationDate":"2025-05-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://onlinelibrary.wiley.com/doi/epdf/10.1111/gcb.70198","citationCount":"0","resultStr":"{\"title\":\"The Deer and the Tiger, the Forest and the Carbon\",\"authors\":\"Oswald J. Schmitz\",\"doi\":\"10.1111/gcb.70198\",\"DOIUrl\":null,\"url\":null,\"abstract\":\"<p>Healthy, verdant forest ecosystems are indicated by an intact complement of highly abundant vegetation along with diverse and abundant populations of herbivore species and their carnivore predators (Wang et al. <span>2025</span>). Keeping forest ecosystems verdant is considered vital to ensuring planetary resilience to climate change (Watson et al. <span>2018</span>). This is because the highly abundant vegetation takes up atmospheric carbon that is then stored in vegetation biomass and in soils of those ecosystems (Pan et al. <span>2024</span>).</p><p>But what would happen to climate change resilience if forest ecosystem intactness was disrupted by, say, the loss of predators? The answer depends on how intactness is sustained. Ecological science has two general views on this (Schmitz et al. <span>2018</span>). One—the bottom-up control view—holds that ecosystems are verdant because vegetation abundance, which is strongly determined by soil nutrient and moisture levels, supports but limits the abundance of herbivore populations, and in turn, populations of their carnivore predators. In this view, climate change resilience would not be disrupted by predator losses (or herbivores for that matter) because animal abundance and diversity do not drive vegetation abundance. The other—the top-down view—holds that predators, by virtue of limiting the abundance of their herbivore prey, keep ecosystems verdant by preventing herbivore overexploitation of vegetation. In this view, climate change resilience would be disrupted by the loss of predators because they ultimately drive vegetation abundance. An important and challenging research frontier for both ecological and global change science is resolving which view of control best explains the climate resilience of verdant forest ecosystems, especially across vast landscapes over which large animals live and roam (Schmitz et al. <span>2018</span>). This challenge is compounded by variation in biophysical conditions across those vast landscapes because the amount of carbon captured and stored among geographic locations becomes highly dependent upon biophysical context, including climatic conditions, the kinds, diversity, and abundances of plant and animal species, and the nutrient contents and physical properties of soils (e.g., Sobral et al. <span>2017</span>; Schmitz et al. <span>2018</span>; Schuldt et al. <span>2023</span>).</p><p>In newly published research in <i>Global Change Biology</i>, Roberts et al. (<span>2025</span>) address this formidable challenge to reveal how varying abundance and outright loss of a large predator—the tiger (<i>Panthera tigris</i>)<i>—</i>and the abundance of its “deer” (i.e., ungulate) prey species are related to the capture and storage of carbon in forests across the tiger's geographic range throughout Asia. This vast landscape has myriad biophysical dimensions that create much context dependency. It contains four broadly different forest ecosystem types, including boreal, temperate, subtropical dry, and subtropical moist. The ecosystems are arrayed across different elevations from lowland to montane and across different wet to dry regimes. They harbor many varieties and abundances of plant and herbivore prey species.</p><p>Resolving the role of tigers in forest climate resilience requires comparing carbon dynamics in places where tigers are present with similar environmental conditions where they are absent (Schmitz et al. <span>2018</span>). The hallmark scientific approach would be to conduct a manipulative Case (tiger-present target forest)–Control (tiger-empty forest) experiment that pairs biophysical conditions of forest locations and systematically excludes tigers from half of them, replicated across the different forest ecosystem contexts across the vast landscape (Schmitz et al. <span>2018</span>). This is altogether logistically impossible. Roberts et al. (<span>2025</span>) overcame this critical limitation with the next best approach (Schmitz et al. <span>2018</span>): they amassed a large multivariate data set gathered from satellite remote sensing and extensive on-the-ground sampling across landscape locations where tigers have been persistently present versus similar landscape conditions where they are absent—their loss arising from known historical extirpation by humans. The amassed data included measures of carbon in vegetation biomass and soil, net ecosystem carbon exchange, and aspects of context dependency including forest ecosystem type, regional climate, and human disturbance. These are exactly the kinds of data used in the conventional accounting of carbon capture and storage in forest ecosystems (Pan et al. <span>2024</span>). As well, they accounted for context dependency in top-down control due to variation in deer diversity and abundance, which according to classical theory (Polis and Strong <span>1996</span>) should be stronger in more linear food chains (i.e., deer diversity is low) and dissipate along the branching network of linkages in food webs with higher prey diversity. There was sufficient replication within and among forest ecosystem types to control for underlying spatial context dependency in these biophysical factors in both tiger present and absent sites. Hence, Roberts et al. (<span>2025</span>) were able to undertake extensive direct and counterfactual analyses to fully and robustly interrogate the veracity of the top-down view that tigers play a role in controlling forest carbon dynamics across their geographic range.</p><p>The short answer to the question about the consequences of predator loss is that it mattered. Tiger presence was generally linked to higher vegetation carbon stocks and higher net carbon exchange (i.e., forest carbon uptake exceeded carbon emissions) among forest types. But the longer answer is that there was also nuance due to context-dependency. Certainly, bottom-up control prevailed across all forests due to geographic variation in biophysical properties that created variation in vegetation biomass and carbon exchange and storage among the forest ecosystems. But top-down effects of tiger presence also prevailed, resulting in higher vegetation and soil carbon stocks among most contexts than in the absence of tiger, the exceptions being tropical dry forest and forests in lowlands and especially wetlands. These exceptions extended to net ecosystem carbon exchange, with one context (tropical swamp forest) even switching from being a slight net carbon sink to a carbon source in the presence of tigers. Tiger presence was impactful in forests with low to intermediate vegetation biomass but not high biomass, suggesting that the biomass dense forests might be completely bottom-up controlled. Yet prey diversity also seems to have played a hand by mediating tiger effects. Forests with low to intermediate vegetation biomass had less deer biomass and diversity than did high vegetation biomass forests. Hence, a second reason for weak, if any, tiger control in biomass dense forests is that the top-down effects may have dissipated along the many food web linkages, suggesting again that both top-down and bottom-up control persisted across contexts but with different relative importance.</p><p>In <span>1967</span> wildlife ecologist George Schaller published a seminal treatise <i>The Deer and Tiger</i> in which he provided foundational scientific understanding about the ecological interplay between tigers and their key prey species. It further described how those relationships were being eroded due to rural land use change for agriculture, persecution of tigers out of fear for human safety and loss of wellbeing, and exploitation for body parts that had perceived medicinal value—problems which continue to this day (Roberts et al. <span>2025</span>). Schaller's book was published at a time when ecological science had barely started to imagine that predator–prey dynamics could cascade to impact the properties and functions of entire ecological communities, let alone the kinds of ecosystem processes such as the carbon cycle that also have implications for human wellbeing. In retrospect it should have been a portent of ramifying ecosystem impacts. But at the time the scientific community was ill-equipped conceptually to even recognize this as a bigger issue because it had yet to embrace a holistic perspective on what it means to sustain verdant forest ecosystems. The portent is still unheeded in many areas of climate change science today, even while scientific understanding of the role of animals in controlling the carbon cycle has advanced considerably (Schmitz et al. <span>2018</span>). By expanding the scientific story of the Deer and the Tiger to the story of the Forest and the Carbon, Roberts et al. (<span>2025</span>) have provided a leap in scientific understanding about how broadly impactful an animal species can be.</p><p>These findings are sure to excite those working on the frontlines of tackling biodiversity loss and climate change by showing how both looming problems can begin to be solved together. There is, however, a risk that arguments about the carbon benefits of tigers will be used to justify headlong action to restore this iconic and endangered species everywhere across its geographic range. Here the deeper, and critical, lesson of the Roberts et al. (<span>2025</span>) article is that by deliberately addressing context-dependency it provides the kind of sober analysis that has been recently called for to temper policy and conservation from overpromoting animal restoration and conservation as a universal win-win for mitigating biodiversity loss and climate change together (Burak et al. <span>2024</span>). The science is clear that tigers can have impactful effects on forest ecosystem carbon storage. But most importantly, Roberts et al. (<span>2025</span>) have provided unprecedented scientific insight into which forest ecosystems across the tiger's vast geographic range should be considered candidates for tiger population conservation and restoration for carbon capture and storage; and which forests definitely should not. The research is exemplary for showing how to advance an evidence-based approach that can be applied in the service of restoring and sustaining verdant forest ecosystems for the purpose of maintaining resilience to climate change.</p><p>The author declares no conflicts of interest.</p><p>This article is a Invited Commentary on Guangshun Jiang et al., https://doi.org/10.1111/gcb.70191.</p>\",\"PeriodicalId\":175,\"journal\":{\"name\":\"Global Change Biology\",\"volume\":\"31 5\",\"pages\":\"\"},\"PeriodicalIF\":10.8000,\"publicationDate\":\"2025-05-07\",\"publicationTypes\":\"Journal Article\",\"fieldsOfStudy\":null,\"isOpenAccess\":false,\"openAccessPdf\":\"https://onlinelibrary.wiley.com/doi/epdf/10.1111/gcb.70198\",\"citationCount\":\"0\",\"resultStr\":null,\"platform\":\"Semanticscholar\",\"paperid\":null,\"PeriodicalName\":\"Global Change Biology\",\"FirstCategoryId\":\"93\",\"ListUrlMain\":\"https://onlinelibrary.wiley.com/doi/10.1111/gcb.70198\",\"RegionNum\":1,\"RegionCategory\":\"环境科学与生态学\",\"ArticlePicture\":[],\"TitleCN\":null,\"AbstractTextCN\":null,\"PMCID\":null,\"EPubDate\":\"\",\"PubModel\":\"\",\"JCR\":\"Q1\",\"JCRName\":\"BIODIVERSITY CONSERVATION\",\"Score\":null,\"Total\":0}","platform":"Semanticscholar","paperid":null,"PeriodicalName":"Global Change Biology","FirstCategoryId":"93","ListUrlMain":"https://onlinelibrary.wiley.com/doi/10.1111/gcb.70198","RegionNum":1,"RegionCategory":"环境科学与生态学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"","PubModel":"","JCR":"Q1","JCRName":"BIODIVERSITY CONSERVATION","Score":null,"Total":0}
引用次数: 0
摘要
健康、郁郁葱葱的森林生态系统是由高度丰富的植被以及丰富多样的食草动物种群及其食肉捕食者组成的(Wang et al. 2025)。保持森林生态系统郁郁葱葱被认为对确保地球对气候变化的适应能力至关重要(Watson et al. 2018)。这是因为高度丰富的植被吸收了大气中的碳,然后储存在植被生物量和这些生态系统的土壤中(Pan et al. 2024)。但是,如果森林生态系统的完整性因捕食者的减少而受到破坏,那么气候变化的适应能力会发生什么变化?答案取决于如何保持完好。生态科学对此有两种普遍观点(Schmitz et al. 2018)。一种是自下而上的控制观点,它认为生态系统是青翠的,因为植被的丰富程度很大程度上取决于土壤的养分和湿度水平,它支持但限制了食草动物种群的丰富程度,反过来也限制了食肉动物捕食者的数量。在这种观点中,气候变化的适应能力不会被捕食者(或食草动物)的减少所破坏,因为动物的数量和多样性不会驱动植被的数量。另一种自上而下的观点认为,捕食者通过限制食草动物猎物的数量,防止食草动物过度利用植被,从而保持生态系统的翠绿。在这种观点中,气候变化的适应能力将被食肉动物的消失所破坏,因为它们最终会推动植被的丰富。生态和全球变化科学的一个重要且具有挑战性的研究前沿是解决哪种控制观点最好地解释了绿色森林生态系统的气候适应能力,特别是在大型动物生活和漫游的广阔景观中(Schmitz et al. 2018)。这一挑战由于这些广阔景观中生物物理条件的变化而变得更加复杂,因为在地理位置之间捕获和储存的碳量高度依赖于生物物理环境,包括气候条件、动植物物种的种类、多样性和丰度,以及土壤的营养成分和物理性质(例如,Sobral等人,2017;Schmitz et al. 2018;Schuldt et al. 2023)。罗伯茨等人(2025年)在《全球变化生物学》(Global Change Biology)上新发表的研究中,解决了这一艰巨的挑战,揭示了大型捕食者老虎(Panthera tigris)的丰度变化和完全消失,以及它的“鹿”(即有蹄类)猎物物种的丰度与整个亚洲老虎地理范围内森林中碳的捕获和储存之间的关系。这片广阔的土地有无数的生物物理维度,产生了很大的环境依赖性。它包含四种不同的森林生态系统类型,包括北方、温带、亚热带干燥和亚热带湿润。生态系统分布在不同的海拔上,从低地到山地,从湿润到干燥。它们拥有多种丰富的植物和食草猎物。要解决老虎在森林气候适应能力中的作用,需要比较有老虎存在的地方与没有老虎的类似环境条件下的碳动态(Schmitz et al. 2018)。标志性的科学方法将是进行一个操纵案例(有老虎的目标森林)-控制(没有老虎的森林)实验,该实验将森林位置的生物物理条件配对,并系统地将老虎排除在其中一半之外,在广阔景观的不同森林生态系统背景下复制(Schmitz et al. 2018)。这在逻辑上是完全不可能的。Roberts等人(2025)利用次优方法克服了这一关键限制(Schmitz等人,2018):他们收集了大量的多元数据集,这些数据集来自卫星遥感和广泛的地面采样,跨越了老虎一直存在的景观地点,以及没有老虎的类似景观条件——它们的损失是由于人类已知的历史灭绝造成的。积累的数据包括植被生物量和土壤碳测量、生态系统净碳交换以及环境依赖方面,包括森林生态系统类型、区域气候和人为干扰。这些正是森林生态系统碳捕获和储存的传统核算中使用的数据(Pan et al. 2024)。此外,他们还解释了由于鹿的多样性和丰度变化而导致的自上而下控制中的情境依赖性,根据经典理论(Polis and Strong 1996),这种依赖性在更线性的食物链中(即鹿的多样性较低)应该更强,并在猎物多样性较高的食物网中沿着分支网络的联系消散。 在森林生态系统类型内部和类型之间存在足够的复制,以控制这些生物物理因子在老虎存在和不存在的地点的潜在空间背景依赖性。因此,Roberts等人(2025)能够进行广泛的直接和反事实分析,以全面而有力地质疑老虎在其地理范围内控制森林碳动态方面发挥作用的自上而下观点的准确性。对捕食者减少的后果这个问题的简短回答是,这很重要。在森林类型中,老虎的存在通常与较高的植被碳储量和较高的净碳交换(即森林碳吸收超过碳排放)有关。但更详细的回答是,由于情境依赖,也存在细微差别。当然,由于生物物理特性的地理差异造成了植被生物量和森林生态系统间碳交换和储存的差异,自下而上的控制在所有森林中普遍存在。但老虎存在的自上而下的影响也普遍存在,导致大多数情况下的植被和土壤碳储量高于没有老虎的情况,除了热带干燥森林和低地森林,特别是湿地。这些例外扩展到生态系统的净碳交换,其中一个环境(热带沼泽森林)甚至在老虎存在的情况下从一个轻微的净碳汇转变为碳源。在低到中等植被生物量而不是高植被生物量的森林中,老虎的存在是有影响的,这表明生物量稠密的森林可能完全是自下而上控制的。然而,猎物的多样性似乎也起到了调节老虎效应的作用。低至中等植被生物量森林的鹿生物量和多样性低于高植被生物量森林。因此,在生物量密集的森林中,老虎控制薄弱(如果有的话)的第二个原因是,自上而下的影响可能已经随着许多食物网的联系而消散,这再次表明,自上而下和自下而上的控制在不同的环境中持续存在,但相对重要性不同。1967年,野生动物生态学家乔治·夏勒(George Schaller)发表了一篇影响深远的论文《鹿与虎》(The Deer and Tiger),在这本书中,他对老虎与其主要猎物物种之间的生态相互作用提供了基础的科学理解。报告进一步描述了这些关系是如何被侵蚀的,原因是农村土地用于农业的改变,出于对人类安全和福祉的恐惧而对老虎的迫害,以及对具有药用价值的身体部位的开采——这些问题一直持续到今天(Roberts et al. 2025)。沙勒的书出版的时候,生态科学刚刚开始想象捕食者-猎物的动态可能会影响整个生态群落的特性和功能,更不用说影响碳循环等生态系统过程,这些过程也会影响人类的福祉。回想起来,这应该是生态系统影响扩大的一个预兆。但当时科学界在概念上还没有充分的准备,甚至没有意识到这是一个更大的问题,因为它还没有从整体的角度来看待维持翠绿的森林生态系统意味着什么。尽管对动物在控制碳循环中的作用的科学理解已经取得了相当大的进步,但在当今气候变化科学的许多领域,这一预兆仍然没有得到重视(Schmitz et al. 2018)。通过将鹿和老虎的科学故事扩展到森林和碳的故事,罗伯茨等人(2025)在科学理解上提供了一个飞跃,即动物物种的影响范围有多广。这些发现肯定会让那些在应对生物多样性丧失和气候变化的前线工作的人感到兴奋,因为它们表明了这两个迫在眉睫的问题是如何开始一起解决的。然而,有一种风险是,关于老虎的碳效益的争论将被用来证明轻率地采取行动,在其地理范围内恢复这种标志性的濒危物种。在这里,Roberts等人(2025)的文章更深刻、更关键的教训是,通过刻意解决环境依赖性,它提供了一种清醒的分析,这种分析最近被呼吁缓和政策和保护,避免过度促进动物恢复和保护,作为共同减轻生物多样性丧失和气候变化的普遍双赢(Burak等人,2024)。科学很清楚,老虎可以对森林生态系统的碳储存产生重大影响。但最重要的是,Roberts等人(2025)提供了前所未有的科学见解,即在老虎广阔的地理范围内,哪些森林生态系统应该被视为老虎种群保护和恢复碳捕获和储存的候选者;哪些森林绝对不应该。 该研究展示了如何推进一种基于证据的方法,该方法可用于恢复和维持青翠的森林生态系统,以保持对气候变化的适应能力。作者声明无利益冲突。本文为蒋光顺等人的特邀评论,https://doi.org/10.1111/gcb.70191。
Healthy, verdant forest ecosystems are indicated by an intact complement of highly abundant vegetation along with diverse and abundant populations of herbivore species and their carnivore predators (Wang et al. 2025). Keeping forest ecosystems verdant is considered vital to ensuring planetary resilience to climate change (Watson et al. 2018). This is because the highly abundant vegetation takes up atmospheric carbon that is then stored in vegetation biomass and in soils of those ecosystems (Pan et al. 2024).
But what would happen to climate change resilience if forest ecosystem intactness was disrupted by, say, the loss of predators? The answer depends on how intactness is sustained. Ecological science has two general views on this (Schmitz et al. 2018). One—the bottom-up control view—holds that ecosystems are verdant because vegetation abundance, which is strongly determined by soil nutrient and moisture levels, supports but limits the abundance of herbivore populations, and in turn, populations of their carnivore predators. In this view, climate change resilience would not be disrupted by predator losses (or herbivores for that matter) because animal abundance and diversity do not drive vegetation abundance. The other—the top-down view—holds that predators, by virtue of limiting the abundance of their herbivore prey, keep ecosystems verdant by preventing herbivore overexploitation of vegetation. In this view, climate change resilience would be disrupted by the loss of predators because they ultimately drive vegetation abundance. An important and challenging research frontier for both ecological and global change science is resolving which view of control best explains the climate resilience of verdant forest ecosystems, especially across vast landscapes over which large animals live and roam (Schmitz et al. 2018). This challenge is compounded by variation in biophysical conditions across those vast landscapes because the amount of carbon captured and stored among geographic locations becomes highly dependent upon biophysical context, including climatic conditions, the kinds, diversity, and abundances of plant and animal species, and the nutrient contents and physical properties of soils (e.g., Sobral et al. 2017; Schmitz et al. 2018; Schuldt et al. 2023).
In newly published research in Global Change Biology, Roberts et al. (2025) address this formidable challenge to reveal how varying abundance and outright loss of a large predator—the tiger (Panthera tigris)—and the abundance of its “deer” (i.e., ungulate) prey species are related to the capture and storage of carbon in forests across the tiger's geographic range throughout Asia. This vast landscape has myriad biophysical dimensions that create much context dependency. It contains four broadly different forest ecosystem types, including boreal, temperate, subtropical dry, and subtropical moist. The ecosystems are arrayed across different elevations from lowland to montane and across different wet to dry regimes. They harbor many varieties and abundances of plant and herbivore prey species.
Resolving the role of tigers in forest climate resilience requires comparing carbon dynamics in places where tigers are present with similar environmental conditions where they are absent (Schmitz et al. 2018). The hallmark scientific approach would be to conduct a manipulative Case (tiger-present target forest)–Control (tiger-empty forest) experiment that pairs biophysical conditions of forest locations and systematically excludes tigers from half of them, replicated across the different forest ecosystem contexts across the vast landscape (Schmitz et al. 2018). This is altogether logistically impossible. Roberts et al. (2025) overcame this critical limitation with the next best approach (Schmitz et al. 2018): they amassed a large multivariate data set gathered from satellite remote sensing and extensive on-the-ground sampling across landscape locations where tigers have been persistently present versus similar landscape conditions where they are absent—their loss arising from known historical extirpation by humans. The amassed data included measures of carbon in vegetation biomass and soil, net ecosystem carbon exchange, and aspects of context dependency including forest ecosystem type, regional climate, and human disturbance. These are exactly the kinds of data used in the conventional accounting of carbon capture and storage in forest ecosystems (Pan et al. 2024). As well, they accounted for context dependency in top-down control due to variation in deer diversity and abundance, which according to classical theory (Polis and Strong 1996) should be stronger in more linear food chains (i.e., deer diversity is low) and dissipate along the branching network of linkages in food webs with higher prey diversity. There was sufficient replication within and among forest ecosystem types to control for underlying spatial context dependency in these biophysical factors in both tiger present and absent sites. Hence, Roberts et al. (2025) were able to undertake extensive direct and counterfactual analyses to fully and robustly interrogate the veracity of the top-down view that tigers play a role in controlling forest carbon dynamics across their geographic range.
The short answer to the question about the consequences of predator loss is that it mattered. Tiger presence was generally linked to higher vegetation carbon stocks and higher net carbon exchange (i.e., forest carbon uptake exceeded carbon emissions) among forest types. But the longer answer is that there was also nuance due to context-dependency. Certainly, bottom-up control prevailed across all forests due to geographic variation in biophysical properties that created variation in vegetation biomass and carbon exchange and storage among the forest ecosystems. But top-down effects of tiger presence also prevailed, resulting in higher vegetation and soil carbon stocks among most contexts than in the absence of tiger, the exceptions being tropical dry forest and forests in lowlands and especially wetlands. These exceptions extended to net ecosystem carbon exchange, with one context (tropical swamp forest) even switching from being a slight net carbon sink to a carbon source in the presence of tigers. Tiger presence was impactful in forests with low to intermediate vegetation biomass but not high biomass, suggesting that the biomass dense forests might be completely bottom-up controlled. Yet prey diversity also seems to have played a hand by mediating tiger effects. Forests with low to intermediate vegetation biomass had less deer biomass and diversity than did high vegetation biomass forests. Hence, a second reason for weak, if any, tiger control in biomass dense forests is that the top-down effects may have dissipated along the many food web linkages, suggesting again that both top-down and bottom-up control persisted across contexts but with different relative importance.
In 1967 wildlife ecologist George Schaller published a seminal treatise The Deer and Tiger in which he provided foundational scientific understanding about the ecological interplay between tigers and their key prey species. It further described how those relationships were being eroded due to rural land use change for agriculture, persecution of tigers out of fear for human safety and loss of wellbeing, and exploitation for body parts that had perceived medicinal value—problems which continue to this day (Roberts et al. 2025). Schaller's book was published at a time when ecological science had barely started to imagine that predator–prey dynamics could cascade to impact the properties and functions of entire ecological communities, let alone the kinds of ecosystem processes such as the carbon cycle that also have implications for human wellbeing. In retrospect it should have been a portent of ramifying ecosystem impacts. But at the time the scientific community was ill-equipped conceptually to even recognize this as a bigger issue because it had yet to embrace a holistic perspective on what it means to sustain verdant forest ecosystems. The portent is still unheeded in many areas of climate change science today, even while scientific understanding of the role of animals in controlling the carbon cycle has advanced considerably (Schmitz et al. 2018). By expanding the scientific story of the Deer and the Tiger to the story of the Forest and the Carbon, Roberts et al. (2025) have provided a leap in scientific understanding about how broadly impactful an animal species can be.
These findings are sure to excite those working on the frontlines of tackling biodiversity loss and climate change by showing how both looming problems can begin to be solved together. There is, however, a risk that arguments about the carbon benefits of tigers will be used to justify headlong action to restore this iconic and endangered species everywhere across its geographic range. Here the deeper, and critical, lesson of the Roberts et al. (2025) article is that by deliberately addressing context-dependency it provides the kind of sober analysis that has been recently called for to temper policy and conservation from overpromoting animal restoration and conservation as a universal win-win for mitigating biodiversity loss and climate change together (Burak et al. 2024). The science is clear that tigers can have impactful effects on forest ecosystem carbon storage. But most importantly, Roberts et al. (2025) have provided unprecedented scientific insight into which forest ecosystems across the tiger's vast geographic range should be considered candidates for tiger population conservation and restoration for carbon capture and storage; and which forests definitely should not. The research is exemplary for showing how to advance an evidence-based approach that can be applied in the service of restoring and sustaining verdant forest ecosystems for the purpose of maintaining resilience to climate change.
The author declares no conflicts of interest.
This article is a Invited Commentary on Guangshun Jiang et al., https://doi.org/10.1111/gcb.70191.
期刊介绍:
Global Change Biology is an environmental change journal committed to shaping the future and addressing the world's most pressing challenges, including sustainability, climate change, environmental protection, food and water safety, and global health.
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