Global Human “Predation” on Plant Growth and Biomass
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Received: 8 July 2019 | Revised: 2 February 2020 | Accepted: 18 February 2020 DOI: 10.1111/geb.13087 RESEARCH PAPER Global human “predation” on plant growth and biomass David G. Jenkins1 | Helmut Haberl2 | Karl-Heinz Erb2 | Andrew L. Nevai3 1Department of Biology, University of Central Florida, Orlando, FL, USA Abstract 2Institute of Social Ecology, University of Aim: Ecological theory is not often applied to human appropriation of net primary Natural Resources and Life Sciences, Vienna, production (HANPP), which estimates reduction of natural net primary production Vienna, Austria 3Department of Mathematics, University of (NPP) due to harvest and land use. Here we use predator–prey theory to evaluate Central Florida, Orlando, FL, USA HANPP as “predation”. Macroecology and adaptive life history strategies also help Correspondence evaluate relationships among global terrestrial HANPP, NPP, and plant biomass (B). David G. Jenkins, Department of Biology, Location: Lands worldwide. University of Central Florida, 4110 Libra Drive, Orlando, FL 32816, USA. Time period: 2000. Email: [email protected] Major taxa studied: Terrestrial plants. Editor: Irena Simova Methods: HANPP and potential NPP allometric scaling were estimated for terrestrial ecoregions (N = 819, for 86% of global land surface area) in the year 2000. HANPP and NPP scaling were compared and projected to current and year 2050 conditions. NPP scaling for potential versus actual conditions were also compared, as were bio- mass turnover rates (T; per year). Results: Global HANPP scales predictably with B; consistent with predator–prey the- ory, HANPP scaling is not clearly satiated at greater B. NPP scaling supports adaptive life history strategies theory. HANPP scaling is c. 16% of NPP scaling; a conserva- tive estimate compared to a grid-based 22%. HANPP scaling could become 25–35% of potential NPP scaling by 2050 due to population growth, or be constrained to 20–26% of potential NPP scaling if resource use efficiency improves. However, B is more sensitive than NPP to human effects, and human population size and HANPP now dominate as predictors of T. Main conclusions: Three ecological theories converged here to broadly support prior empirical estimates and enable novel insights. B and T are more sensitive to global human impacts than is NPP and should be priorities for carbon budgets and conser- vation. Human population growth and resource use efficiency strongly affect terres- trial plant HANPP, B and T, and thus global carbon budget. Both human “top-down” effects (evaluated here) and “bottom-up” drivers (e.g., climate, nutrients, CO2) need to be incorporated into global carbon models. KEYWORDS adaptive strategies theory, allometric scaling, biomass turnover rate, HANPP, macroecology, metabolic theory, NPP, plant biomass, predator–prey theory Global Ecol Biogeogr. 2020;00:1–13. wileyonlinelibrary.com/journal/geb © 2020 John Wiley & Sons Ltd | 1 2 | JENKINS ET AL. 1 | INTRODUCTION global carbon analyses because NPP is related to removal of C from the atmosphere by plants, whereas HANPP contributes to increased While modifying and using >75% of terrestrial land area (Pandit et al., atmospheric C. 2018), humans appropriate a substantial portion of global terrestrial Global HANPP has been estimated for over 40 years using differ- annual plant growth (Haberl, Erb, & Krausmann, 2014; Imhoff et al., ent definitions and increasingly sophisticated methods (see review 2004; Vitousek, Ehrlich, Ehrlich, & Matson, 1986). For example, an- by Haberl et al., 2014). Those differences mean that estimates can- nual crop growth per unit area is often less than in primary forests not be easily compared, but recent values (i.e., after 1970s in table 1 cleared for those crops, and plant growth in urban areas is typically of Haberl et al., 2014) average 24 ± 10% (standard deviation) of reduced by paved and roofed areas. Plant growth is known as net potential NPP; a large impact by humans. Human effects on global primary production (NPP), and is quantified as a rate (g C/m2/year; plant biomass (B; g C/m2; Table 1) and turnover have also been quan- Table 1). Human appropriation of NPP (HANPP; also g C/m2/year; tified (e.g., Erb et al., 2016, 2018). Table 1) quantifies human impacts on NPP, including the sum of Analyses of global HANPP and human effects on B quantify land harvest and land use–related changes (e.g., agriculture, timber har- use changes and effects on the global carbon cycle, but have not vesting, urban development; Haberl et al., 2014). This definition of been evaluated through the lens of ecological theory. If ecological HANPP builds on earlier work (Vitousek et al., 1986; Whittaker & theory is general (i.e., not peculiar to specific ecological systems), Likens, 1973; Wright, 1990) and is more inclusive than most earlier it should: work at global scales; apply to humans; predict relation- definitions (Haberl et al., 2014). In turn, the portion not appropri- ships among HANPP, NPP and B; and offer an explanatory basis for ated by humans quantifies NPP available for the planet’s remaining models of these variables that are vital to ecosystem ecology, global biodiversity and sustained ecosystem function. Accordingly, HANPP carbon cycling and sustainable human use of natural resources. In research has influenced discussions of sustainable population turn, using ecological theory to evaluate global HANPP, NPP and growth, conservation planning, planetary “tipping points”, and the B tests that theory and helps to unite it with sustainability science Anthropocene (e.g., Barnosky et al., 2012; Ellis et al., 2013; Foley (e.g., Opdam, Luque, Nassauer, Verburg, & Wu, 2018). et al., 2005; Mellody et al., 2014; Running, 2012). HANPP focuses Here we apply predator–prey theory to evaluate global HANPP on terrestrial ecosystems and complements other analyses of non- as “top-down” regulation of plants in the sense of a food chain, renewable resources (e.g., Brown et al., 2014; Meadows, Meadows, where humans are the predator and “predation” on plants includes Randers, & Behrens III, 1972; Sabin, 2013) as a measure of our col- all effects in the definition of HANPP. This view complements “bot- lective impact. Comparison of NPP and HANPP directly relates to tom-up” estimates of NPP based on plant biology, CO2 enrichment TABLE 1 Terms used in this work, with definitions, acronyms, units, meanings and uses Term Acronym Units What it measures How it is used 2 Potential net primary NPPpot g C/m /year Potential plant growth rate, for natural NPPpot ~ B; natural NPP production conditions scaling 2 Actual net primary production NPPact g C/m /year Actual plant growth rate, after human NPPact ~ B; recent (year land use effects 2000) NPP scaling 2 Ecological net primary NPPeco g C/m /year Actual plant growth, after human land use As part of HANPP estimates production effects and harvest (HANPP = NPPpot–NPPeco) Human appropriation of net HANPP g C/m2/year Human use of plant growth, including land HANPP ~ B; estimate primary production use (HANPPlu) and harvest (HANPPharv) of human functional response, in the sense of predator–prey theory 2 Potential plant biomass Bpot g C/m Potential, aboveground + belowground As the predictor axis for plant mass in an area, for natural allometric scaling of conditions NPPpot, HANPP and Tpot 2 Actual plant biomass Bact g C/m Actual, aboveground + belowground plant As the predictor axis for mass in an area, after human effects allometric scaling of NPPact and Tact Potential plant biomass turnover Tpot Per year Potential rate that biomass changes (as Allometric scaling of natural rate in a slope coefficient for NPP/B), for T; how T is predicted natural conditions by plant B in natural conditions Actual plant biomass turnover Tact Per year Actual rate that biomass changes (as in Allometric scaling of actual rate a slope coefficient for NPP/B), after T; how T is predicted by human effects plant B and human effects JENKINS ET AL. | 3 and climate conditions (e.g., Körner, 2015; Malhi et al., 2015). curve occurs when predators switch to the subject prey (here plants) Ultimately, both perspectives will likely prove valuable in more in- when it reaches greater abundance. Here Types I or II are possible, tegrative global ecosystem models. We use a confluence of three but Type III would need a prior resource and does not apply. ecological theories (predator–prey, macroecology, and adaptive life Predator functional responses have been developed to describe history strategies; Figure 1) to investigate relationships between individual predators or local populations; we here extend that anal- HANPP, NPP and B (Brown 1995; Grime & Pierce, 2012; Holling, ysis to the global human population. This extension is justifiable 1959; May, 1977; Noy-Meir, 1975). Below is a brief synopsis of each because: (a) humans are dominant, hyperkeystone, super-predators theory as it applies here, and the confluence between them. (Darimont, Fox, Bryan, & Reimchen, 2015; Vitousek, Mooney, Lubchenco, & Melillo, 1997; Worm & Paine, 2016); (b) the human population now outweighs all wild mammals (Bar-On, Phillips, & 1.1 | Graphical predator–prey theory Milo, 2018), consistent with our role as a dominant global popu- lation; (c) the human population acts globally on terrestrial plants, Graphical predator–prey theory evaluates and then compares two as evidenced by > 2 trillion kg of agricultural and forestry products curves. Predator consumption and prey growth are each expressed transported annually (Chatham House, 2018); and (d) HANPP is as a function of prey quantity (Figure 1a,b; May, 1977; Noy-Meir, common to nearly all human socioeconomic and cultural groups. 1975). A predator consumption curve is also known as a functional The other half of predator–prey theory is the prey growth curve response and classically expected to be linear, asymptotic or sig- (Figure 1b; here NPP ~ B). Original models expected a hump-shaped moidal in shape (Types I–III, respectively), where the shape of the curve because only new green growth was considered (May, 1977; curve indicates behaviour of a predator population (Holling, 1959).