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Measuring the value of groundwater and other forms

of natural capital
Eli P. Fenichela,1, Joshua K. Abbottb, Jude Bayhama,c, Whitney Boonea, Erin M. K. Haackerd, and Lisa Pfeiffere
a
School of Forestry and Environmental Studies, Yale University, New Haven, CT 06460; bSchool of Sustainability, Arizona State University, Tempe, AZ 85287;
c
College of Agriculture, California State University, Chico, CA 95929-0310; dDepartment of Geological Sciences, Michigan State University, East Lansing,
MI 48824; and eNorthwest Fisheries Science Center, National Marine Fisheries Service, National Oceanic and Atmospheric Administration, Seattle, WA 98112

Edited by Stephen Polasky, University of Minnesota, St. Paul, MN, and approved December 31, 2015 (received for review July 13, 2015)

Valuing natural capital is fundamental to measuring sustainability. et al. (9) review the theory of natural capital prices in wealth
The United Nations Environment Programme, World Bank, and indices, and note the dearth of theory for measuring them.
other agencies have called for inclusion of the value of natural Moving beyond rhetoric to valuing natural capital is imperative
capital in sustainability metrics, such as inclusive wealth. Much has for reforming national accounting and developing sustainability
been written about the importance of natural capital, but measures (2, 10–13), tracking the sustainable use of specific socio-
consistent, rigorous valuation approaches compatible with the ecological systems (14), and mainstreaming natural capital in a
pricing of traditional forms of capital have remained elusive. We way that makes it comparable to other fiscal goals for policy
present a guiding quantitative framework enabling natural capital analysis (15, 16).
valuation that is fully consistent with capital theory, accounts for We price natural capital in a manner fully consistent with
biophysical and economic feedbacks, and can guide interdisciplin- economic capital theory (17). Our approach reflects real-world,
ary efforts to measure sustainability. We illustrate this framework “kakatopic” (18) institutional and management arrangements

SUSTAINABILITY
with an application to groundwater in the Kansas High Plains and encompasses the dynamics of the coupled socioecological
Aquifer, a rapidly depleting asset supporting significant food system (19). This approach directly addresses the Achilles’ heel

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production. We develop a 10-y time series (1996−2005) of natural of the inclusive wealth metric by responding to Smulders’ (8) call
capital asset prices that accounts for technological, institutional, for “good theorists and clever empiricists to . . . close the gap
and physical changes. Kansas lost approximately $110 million per between theory and practice.” Our framework generalizes the
year (2005 US dollars) of capital value through groundwater with- applicability of Jorgenson’s (17) classic capital asset pricing
drawal and changes in aquifer management during the decade approach—a pillar of economic capital theory—beyond marketed
spanning 1996–2005. This annual loss in wealth is approximately assets, thereby providing natural capital prices for rigorous policy
equal to the state’s 2005 budget surplus, and is substantially more analysis and enabling “apples to apples” comparisons with tradi-
than investments in schools over this period. Furthermore, real tional capital assets (e.g., real estate, machinery, or financial as-
investment in agricultural capital also declined over this period. sets). Specifically, we show that Jorgenson’s asset pricing
Although Kansas’ depletion of water wealth is substantial, it equation does not rely on the often questionable assumption of
may be tractably managed through careful groundwater manage- an optimizing economy, even for nonmarket natural capital. The
ment and compensating investments in other natural and tradi- pricing equation continues to provide the revealed marginal social
tional assets. Measurement of natural capital value is required to benefit from holding an additional unit of natural capital. The
inform management and ongoing investments in natural assets. framework is applicable to the full range of natural capital stocks
and is scalable to capture greater socioeconomic and biophysical
|
groundwater sustainability | inclusive wealth | comprehensive wealth | detail when data permit. Thus, we provide a pathway for resolving
genuine investment Polasky et al.’s (7) concerns about the omission of natural capital
that does not form market commodities. Moreover, we bridge

S ustainability scholars and advocates are abuzz about natural


capital (1). Natural capital is a powerful metaphor conveying
the importance of Earth’s biotic and abiotic natural resources as
Significance

society’s productivity base, capable of providing ongoing flows of Economists have long argued, with recent acceptance from the
socially valuable services. This rhetoric places natural capital on science and policy community, that natural resources are capital
common conceptual terms with “traditional” produced assets, assets. Pricing of natural capital has remained elusive, with the
enabling policy makers to frame resource management and sus- result that its value is often ignored, and expenditures on con-
tainability challenges as a form of “portfolio management.” How- servation are treated as costs rather than investments. This ne-
ever, operationalizing natural capital requires that decision makers glect stems from a lack of a valuation framework to enable apples
evaluate tradeoffs across capital stocks using a common currency. to apples comparisons with traditional forms of capital. We de-
A lack of prices for valuing natural capital stocks continues to velop such an approach and demonstrate it on Kansas’ ground-
hamper progress toward including natural capital in social benefit−cost water stock. Between 1996 and 2005, groundwater withdrawal
analyses or the accounts used to measure social progress (2–4). A reduced Kansas’ wealth approximately $110 million per year.
major barrier for implementing “inclusive or comprehensive wealth,” Wealth lost through groundwater depletion in Kansas is large,
the United Nation’s and World Bank’s advocated sustainability metric, but in a range where offsetting investments may be feasible.
is measuring the value of natural capital (2, 5–7). Smulders (8) writes,
“The Achilles’ heel of the method [Inclusive or Comprehensive Author contributions: E.P.F. and J.K.A. designed research; E.P.F., J.K.A., J.B., W.B., E.M.K.H.,
and L.P. performed research; E.P.F. and J.K.A. contributed analytic tools; E.P.F., J.B., W.B.,
Wealth or Genuine Savings] is the determination of the shadow E.M.K.H., and L.P. analyzed data; and E.P.F., J.K.A., J.B., W.B., E.M.K.H., and L.P. wrote
prices,” where shadow prices refer to the appropriate natural the paper.
capital prices. Polasky et al. (7) emphasize that in current at- The authors declare no conflict of interest.
tempts to measure inclusive wealth, “all measures included in This article is a PNAS Direct Submission.
natural capital were values for market commodities,” but 1
To whom correspondence should be addressed. Email: Eli.Fenichel@yale.edu.
“evaluating sustainability via inclusive wealth . . . requires an This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.
assessment of the changes in value of all types of capital.” Hanley 1073/pnas.1513779113/-/DCSupplemental.

www.pnas.org/cgi/doi/10.1073/pnas.1513779113 PNAS Early Edition | 1 of 6


literature using ecological−economic production functions (20) which measure the social worth of an additional unit of the asset
and the macroeconomic literature on inclusive wealth by providing (18). Wealth is “inclusive” if all assets, including nature, enter
a framework for providing the necessary capital prices. this sum (2, 12, 13). If a country, region, or project’s ability to
We link economic measurement of ecosystem service flows, a generate future well-being is stable or increasing over time, then
form of income dependent on nature (21), with models of bio- it can be said to be sustainable, and nondiminishing inclusive
physical dynamics and adaptive human behavior (shaped by wealth is a necessary condition for this sustainability (37). Thus,
policy and other institutional constraints), to value quantities of what matters for sustainability are changes in wealth, or net in-
natural stocks, which are capital (21). This focus on natural vestment, across all capital stocks. Measuring changes in natural
capital’s durable value in place differentiates our approach from capital value is not sufficient for assessing sustainability, but
the considerable progress made in valuing short-run flows of ben- it is necessary. For example, measuring the sustainability of a
efits deriving from natural capital—termed “ecosystem services” groundwater-dependent agricultural system requires measuring
(20, 22–24). The value of ecosystem service flows affects the value the wealth held in the aquifer. If the value of groundwater capital
of natural capital; however, they are not equivalent. Ecosystem
is declining, then substantial investment in other capital stocks
service values must be integrated with other important social,
may be required to achieve sustainability. Our goal is to dem-
economic, and biophysical data (25) to produce estimates of the
onstrate how natural capital asset prices for use in inclusive
long-run value of natural stocks as durable assets (i.e., natural
capital). This integration has lagged considerably. Moreover, our wealth accounting can be estimated—using a single, but impor-
work contrasts with approaches borne from accounting and mass tant, asset as an example. Developing complete inclusive wealth
balance (26) that suffer from a lack of a consistent framework for accounts is beyond the scope of this paper; instead, we show how
integrating environmental and social data to evaluate tradeoffs to compute natural capital prices to fill an important void in
and that confuse stocks and flows. existing wealth accounts, including in the 2014 IWR.
We illustrate our approach with an application to the High To operationalize the inclusive wealth framework for a spe-
Plains Aquifer in the American Great Plains. This aquifer is of cific natural asset, it is essential to know the accounting price for
critical importance to American agricultural production (27). valuing the stock. For these prices to guide real-world resource
Groundwater is a crucial asset globally, supporting 40% of the management decisions, they must be marginal prices grounded
world’s food production (28). However, its value as natural in capital theory (17), reflecting the change in current and future
capital has not been credibly estimated (29–31). Indeed, the lack well-being from an incremental increase of the stock (38). These
of values for water is lamented in the 2014 Inclusive Wealth conditions require that accounting prices (Fig. 1, gray circle)
Report (IWR) (2). Previous attempts have inventoried ground- connect human investment/consumption behavior (Fig. 1, yellow
water stocks, providing physical measurements of quantities box), including use of natural resources with biophysical dy-
without values (27, 32), or have valued the flows of ecosystem namics of the resource stock (Fig. 1, orange box and arrows), in
services associated with water use (30, 33, 34), which provides a an internally consistent fashion (14). Moreover, accounting prices
poor approximation to the wealth contained in water. To value need to account for projected feedbacks from natural assets
the groundwater capital stock, we focus on the regions of the to human behavior (Fig. 1, maroon circle and arrow to human
aquifer associated with crop agriculture, which received 99% of behavior), which are often adaptive and always policy or in-
the groundwater pumped (35) in western Kansas over a period of stitutionally conditioned. These behavioral rules are often called
a major technological change, 1996–2005. We find that the in- the “economic program” (12, 14). For accounting prices to be
cremental value of an extra acre-foot of water (1 acre-foot = relevant to management, the economic program must reflect real-
1,233.5 m3), the present value of foregone future production by world institutions, technology, and management (Fig. 1, black
pumping an extra acre-foot at present, for the average farmland box) rather than idealized, optimized policies.
acre was between $7 and $17 (7% or 3% discount rate). The Fig. 1 is more than a conceptual framework; it illustrates a
present value of profits attributable to the Kansas portion of the
concrete formula for computing the price of natural capital
aquifer, the value as natural capital, declined from $2.3 billion in
(large light gray box). The natural capital accounting price
1996 to $1.2 billion in 2005—a loss of $110 million per year
(2005 US dollars, 3% discount rate). To achieve sustainability, at function, p(s(t)), can be expressed as a function of the stock of
minimum, Kansas would need to have offset these losses with natural capital, s(t), at an instant in time, t, and parameters
equivalent investments in other forms of capital, i.e., used the characterizing biophysical dynamics, human behavioral feed-
Hartwick Rule (36). The annualized losses are more than twice backs and the value of ecosystem service flows (14). (We sup-
the state’s annual investments in school infrastructure (a pre- press t in our notation when doing so does not cause confusion.)
cursor to human capital) (budget.ks.gov/publications/FY2005/ Importantly, the economic program, x(s(t)), is embedded within
FY2005_Governors_Budget_Report–Volume_1(rev2-09-2004). the price function, thus embracing the role of institutions and
pdf). Moreover, Kansas’s net investment in traditional forms of human behavior as well as ecological and economic dynamics.
agricultural capital (e.g., farm machinery and financial assets)
appears to have declined in real terms [www.ers.usda.gov/data- _
MDðs, xðsÞÞ + pðsÞ
pðsÞ = [1]
products/farm-income-and-wealth-statistics/us-and-state-level- δ − ½MGðsÞ − MHIðs, xðsÞÞ
farm-income-and-wealth-statistics-(includes-the-us-farm-income-
forecast-for-2015).aspx]. These comparisons raise questions The equation for pðsÞ is derived directly from economic capital
about the magnitudes and types of investment that can maintain theory following refs. 14, 17, and 39, and provided in Supporting
public wealth and the sustainability of Kansas agriculture. Information. However, our derivation enables us to connect mea-
surement of natural capital with applications in wealth account-
Valuing Natural Capital ing in a theoretically consistent way. Jorgenson (17) assumes a
Capital assets are stocks with the potential to generate flows of market allocation mechanism to derive Eq. 1, making Jorgenson’s
current and future well-being through production of goods and formulation inappropriate for natural capital not traded in mar-
services (21). All economies rely on a combination of produced, kets. Arrow et al. (39) develop theory for the nonoptimizing
human, and natural capital for production (2). In the case of economy to derive the inclusive wealth measure of sustainability
agriculture, soil, water reserves, machines, and human know-how from first principles, but do not use this theory to inform the
are all capital assets. Wealth is the summed value of productive measurement of accounting prices. We contribute to the litera-
assets, valued at appropriate accounting prices (shadow prices), ture by providing the general link between Jorgenson’s capital

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Fig. 1. Conceptual model and equation for valuing natural capital.

asset price theory and the theory of inclusive wealth accounting from an additional unit of stock. This is the gross physical return to
for economies that don’t optimally allocate all resources. conserving the stock. This term can be positive, negative, or zero,
Eq. 1 corresponds to Jorgenson’s foundational equation (ref. depending on the characteristics of the resource (for example,
17, p. 249) for the value of invested capital. MD stands for whether the stock influences the rate of growth), its abundance,
marginal dividends and is the incremental flow benefit from a and its influence on its own regeneration. MHI is the marginal
small increase in the natural capital stock—the value of the human impact on the capital stock (e.g., drawdown) resulting from
ecosystem services produced with slightly more natural capital an additional unit of stock. This term reflects the feedbacks
(Fig. 1, brown box). The stock of natural capital can directly between changes in resource stocks and human investment/
influence these dividends, or they can arise as a function of consumption decisions. Together, these terms constitute the
natural capital’s influence on human behavior via the economic net physical appreciation or depreciation resulting from an addi-
program. Ecosystem service values have received extensive at- tional investment in natural capital. This “socioecological rate of
tention in the literature but are often poorly distinguished from interest” (Fig. 1, orange box) is subtracted from the baseline dis-
the value of natural capital (20, 22, 40, 41). Fig. 1 makes clear count rate, so that a stock with a positive net appreciation faces a
that, although related, natural capital and ecosystem services are lower effective discount rate (45), increasing its accounting price.
not equivalent. There are several mitigating factors that prevent Neglecting the ecological adjustment to the effective discount rate
a one-to-one mapping. Ignoring these factors can be highly can result in dramatic errors in estimating the accounting price of
misleading, potentially suggesting value is increasing when it is in natural capital, especially for self-renewing resources, e.g., fish or
fact declining, and vice versa (14). naturally regenerating forests (14).
The denominator of Eq. 1 functions as the effective discount The term p_ = dp=dt represents changes in price, reflecting
rate—translating the flows of benefits in the numerator to a capital gains or losses from the overall flux (net of human con-
forward-looking value for an increment of the stock. Here δ is sumption driven through the economic program) in natural
the “raw” discount rate before any adjustments for biophysical or capital stocks (Fig. 1, dark gray box). This rate of price appre-
anthropogenic processes of depreciation or appreciation. Dis- ciation is often observed or forecasted for traditional capital.
count rates (Fig. 1, blue box) for natural capital are controversial This is rarely possible for natural capital, because natural capital
(42, 43). We follow refs. 14 and 43 and use two rates suggested by prices are seldom observable. However, the time path of natural
the US Office of Management and Budget (OMB) (44). OMB capital prices, and hence p,_ is implicit in a fully specified socio-
recommends a constant, conservative, consumption rate of 3% ecological systems model (19), defining the remaining elements
when a “social rate of time preference” is required and a higher in Fig. 1. Function approximation methods (see Supporting In-
rate of 7% when policy affects shifts in private capital. We treat formation) are used to recover values for p_ consistent with the
the 3% rate as the base case and 7% as sensitivity analysis be- maintained model of biophysical and economic dynamics and the
cause the social rate of time preference is arguably most ap- service flow valuation assumptions. The accounting prices com-
propriate for accounting prices for public inclusive wealth accounts. puted using Eq. 1 are conceptually the same as traditional capital
MG is the marginal growth, appreciation, or depreciation arising asset prices (14).

Fenichel et al. PNAS Early Edition | 3 of 6


22 0.9

FracƟon using drop nozzles


mean water in aquifer

Mean water in the acƟve


21.8 0.8

aquifer, acre feet


0.7
21.6
FracƟon using 0.6
21.4 0.5
drop nozzle
21.2 0.4
0.3
21
0.2
20.8 0.1
20.6 0
1996 1997 1998 1999 2000 2001 2002 2003 2004 2005
Year
Fig. 2. The mean acre-feet of water per acre of land and fraction of fields using drop nozzles over time.

The unit price of natural capital and quantity of natural capital function shows that the marginal value of an additional acre-foot
stock are multiplied to ascertain the stock’s contribution to in- of water declines with increases in the stock of water. The av-
clusive wealth (Fig. 1, green box). For small changes in the capital erage acre overlies 21.5 acre-feet of water, and has an accounting
stock, it is acceptable to multiply changes in quantity by a constant price of $17 per acre-foot using a 3% discount rate. This implies
price to calculate changes in inclusive wealth (2, 12, 13, 39). that water underlying the average Kansas acre with access to the
However, Eq. 1 dictates that, for large stock changes, allowances aquifer contributes $396 in value, approximately a third of 2005
must be made for stock-driven changes in price (39, 46). price of irrigated farmland in western Kansas (49). Using a 7%
A unique strength of the framework in Fig. 1 is that it can be discount rate, the accounting price falls to $7 per acre-foot, im-
applied to natural, produced, or human capital—inviting “apples plying an average value contribution of $173 per acre. OMB’s
to apples” comparisons. Consider housing as a stock of produced upper discount rate is 2.33 times greater than its lower rate. Using
capital. MD (brown box) is the net value of housing services arising the lower rate yields an accounting price at mean water quan-
from the “marginal” house—effectively, the rent the house could tity 2.4 times the accounting price associated with the higher
generate after deductions for costs, e.g., for maintenance and in- rate—highlighting the importance of discount rate choice.
surance. The economic program in this case includes the owner’s
decisions regarding insurance, maintenance, and general use of Changes in Water Wealth. Changes in wealth, rather than absolute
the house. The discount rate is the owner’s rate of return on other wealth, matter for measuring sustainability. The changes in ground-
investments. Wear and tear, which is partially a function of living water levels that occurred in Kansas between 1996 and 2005 are of
decisions, made as part of the economic program, physically de- significant magnitude, involving nontrivial changes in the quantity
preciates the house over time. This marginal depreciation is par- and scarcity of the stocks. Small changes in quantities have minimal
tially offset through maintenance, also a function of the economic impact on marginal values, and can be valued at a constant price
program, but likely yields a net positive MHIðsÞ, which results in times the change in quantity. In such a case, the accounting price,
an increase in the effective discount rate. Houses do not renew pðsÞ, function can be approximated as a horizontal line. For
themselves, so MGðsÞ = 0. This is a key difference between most
produced and some natural capital. Capital gains, p,_ are forecasted
based on observed price data. Estimates of these terms can all be 30
found on housing price websites (e.g., Zillow), and the interested mean 3%
reader will see that combining them will typically yield a reason- mean 7%
able estimate for the market price of a house. Although capable of 25 1996, 3%
explaining prices for produced capital, the real power of our
framework is that it can impute prices for natural capital in 2005, 3%
Accounting price

a symmetric fashion. 20
Case Study: Groundwater
To illustrate the power of the framework for natural capital, we ex- 15
amine an application to groundwater. We focus on first-order hy-
drological concerns to illustrate how groundwater and other natural
capital assets can be valued. We define the stock of groundwater as 10
the thickness of the saturated zone, which is mostly rock, multiplied
by an estimate of specific yield to convert the saturated thickness of
rock to the water held in the aquifer (47, 48). This allows us to value a 5
marginal increase in the water volume contained under one surface
acre. We provide an initial estimate for the Kansas High Plains for
1996–2005. There were many changes in agriculture over this decade. 0
For example, farmers adopted high-efficiency center-pivot drop 0 20 40 60 80 100 120
nozzles (Fig. 2). Over this same period, the stock of water contained Acre feet of water in aquifer per square acre
under the average acre fell ∼1 foot, a rate of 0.4% annually. Fig. 3. Accounting price function for 1996, 2005, and the mean of those
and all years in between using a 3% discount rate. The accounting price
The Value of Water. Using Eq. 1, we estimate an accounting price function for the mean of all years using a 7% discount rate is also shown.
for an additional acre-foot of water, pðsÞ (Fig. 3). This price Circles show the price at the mean water stock.

4 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1513779113 Fenichel et al.


example, firm market capitalization is computed as price times including groundwater. For at least the last 20 years, ad hoc em-
the number of shares. However, this is a poor approximation of the pirical approaches attempting to relax the efficient allocation as-
change in value arising from significant changes in natural capital sumption have been suggested (26) and discredited (25). Others
stocks, which are often localized in nature with few readily available have made valiant efforts to measure stocks of resources and track
substitutes. In these cases, the scarcity of the natural capital stock ecosystem dividends (1, 22, 23). By returning to the first principles
dictates its marginal value. To account for scarcity effects, the of capital theory, we present a bottom-up approach that explicitly
change in value of the aquifer is best measured as the area under shows how to integrate social, economic, and biophysical modeling
the accounting price curve between any two quantities of water in and data to yield appropriate forward-looking accounting prices
the aquifer (Fig. 3). Fig. 3 makes it clear that, in the neighborhood rooted in the particular biophysical, social, economic, and in-
of the mean amount of water held in the aquifer (indicated by stitutional conditions of specific natural capital stocks. Such a
circles), the accounting price functions are not horizontal lines. bottom-up approach is necessary in many cases, given the wide-
We compute year-specific accounting price functions (examples spread lack of asset markets for natural capital, formidable physical
in Fig. 3) for water by adjusting year-specific means for all variables, barriers to arbitrage, and the inherently specialized and local na-
including nozzle adoption. Combining the changes in the account- ture of many of the services provided by natural capital.
ing prices with the changes in water volume, we create a time series Beyond providing an equation for valuing natural capital, Fig. 1
for the value of the Kansas High Plains Aquifer. Between 1996 and presents a framework to guide interdisciplinary efforts for gener-
2005, the value of the Kansas portion of the aquifer fell at an an- ating natural capital prices for measuring and monitoring sustain-
nualized rate of 6.5% or approximately $110 million per year, using ability. Economists must continue to improve methodologies for
the preferred 3% discount rate, and 6.5% or approximately $31 valuing ecosystem services, but these efforts must be coordinated
million per year, using the 7% discount rate. Kansas had a pro- with measurements of other parts of the system. Biophysical sci-
jected budget surplus of $113 million in 2005. If this level is rep- entists already play an important role in quantifying natural capital
resentative, then Kansas appears to have had the financial means to and understanding the production functions for ecosystem services;
have invested in a “sovereign wealth fund” to enable offsetting biophysical science also needs to contribute to establishing the ef-
capital investment—a sort of “Hartwick fund.” It is an empirical fective discount rate for natural capital pricing. Measurements of

SUSTAINABILITY
question beyond the scope of this paper as to whether Kansas made marginal human impact will likely require collaboration among

SCIENCE
such durable investments or consumed its surplus. biophysical scientists, economists, and other social scientists, and
Fig. 3 illustrates that price curves shifted downward through broad collaboration across the social sciences (including economics)
time (only results for the 3% discount rate are shown). This is needed to understand the economic program that links institu-
downward shift corresponded with a rise in water-efficient drop tions and natural capital states to human behavior. The strength of
nozzle technology, substantially accelerated by state subsidies for Fig. 1 is that it shows exactly what interdisciplinary teams need to
irrigation upgrades. Prior research (50) found that the adoption measure and how to integrate those measurements.
of this technology actually increased water use, as increased ir- Kansas experienced a nontrivial loss in water wealth from 1996
rigation efficiency reduced the cost of an effective unit of water, to 2005. The annualized rate of loss of physical water stock, 0.4%,
inducing farmers to irrigate a greater proportion of their acreage severely underestimates the rate at which wealth was lost, 6.5%.
and plant more water-intensive crops. The downward shift of the However, these losses are not so great as to be insurmountable, if
price function reflects the fact that the technological shift actu- care is taken to balance resource depletion with sufficient com-
ally made water appear less scarce, reducing the value of a pensatory investments. Our framework for measuring the value of
greater water stock (or the apparent cost of depleting water today). groundwater specifically, and natural capital generally, is readily
The adoption of new technologies often requires institutional ad- transferable to other systems. However, our prices are not. Mea-
aptation to maintain wealth—adaptations that Kansas failed to suring the value of groundwater in more-complex settings may
make as the new nozzles emerged. The shift in price curves il- require different approaches to measuring the marginal dividends,
lustrates the joint importance of institutions and technology in recharge, and marginal human impact. Indeed, not all ground-
determining the value of natural capital. As a thought experiment, water is as valuable as our estimates, but much groundwater flows
imagine that the same amount of water had been withdrawn be- to high-valued residential use and is likely considerably more
tween 1996 and 2005, but there had been no technological or other valuable, as in California and the Desert Southwest.
shift to move the price curve downward from the 1996 level (i.e., Our example illustrates the importance of how institutions,
the realized drop nozzle adoption did not occur). In this case, technology, and other factors shape the dividend flows from
Kansas would have lost $10 million ($4 million) per year using the natural capital and the economic program. Natural capital prices
3% (7%) discount rate, and the rate of decline in value would have are not immutable natural parameters to be discovered. They are
been close to the rate of physical withdrawal. functions of the way society interacts with the resource as mapped
through the economic program and marginal dividends. The large
Discussion loss in aquifer-associated wealth that Kansas actually experi-
The phrase “natural capital” permeates current sustainability enced resulted from a combination of the physical drawdown of
discussions. US federal agencies were recently instructed to ac- the stock and a state-subsidized shift toward an economic pro-
count for ecosystem services and natural capital in policy plan- gram that was less conservation oriented, even though it, para-
ning (51). However, the idea of treating nature as capital is old doxically, involved a shift toward “highly efficient” nozzles. By
(52) and well accepted. Fisher (21) clearly classifies fish stocks failing to anticipate and mitigate the perverse consequences of
and public lands as capital in his foundational 1906 book on the technological transition, statewide “investments” in improved
income and capital. The conceptual problem has been how to technology actually destroyed wealth.
value nature as capital such that the prices used are comparable Measuring the value of natural capital and assessing whether or
to capital prices observed in markets when few natural assets are not resource use is sustainable go hand in hand. However, while
allocated through efficient market mechanisms. Prior efforts to measuring the value of natural capital is necessary, it is not suf-
measure the value of natural capital stocks in situ have required ficient for measuring sustainability. In addition to changes in
assuming a competitive market for the stock and efficient allo- wealth, changes in population can also be important; under certain
cation or that the value of the asset is zero (53, 54). However, circumstances, sustainability can be assessed via changes in per
public policy and informational institutions have led to nonmarket capita (or perhaps median) wealth (39). However, care may need
allocation mechanisms that make the Pareto efficient allocation to be taken in measuring the value of services flows and MHI,
assumption untenable for many important natural resource stocks, which may be a function of population size. Coordinated efforts to

Fenichel et al. PNAS Early Edition | 5 of 6


generate credible accounting prices are needed if the concept of revenues. MHIðs,xðsÞÞ is influenced indirectly through the economic program
natural capital is to be actionable, rather than a trite reminder to by the irrigation requirements of various crop choices. MHIðsÞ was computed
decision makers to “please remember the environment.” Our as the change in water in the aquifer resulting from water withdrawal. We
assume the aquifer recharges at a rate of 1.25 inches per year, and that the
framework provides a rigorous, integrative, and scalable approach
only losses are through agricultural withdrawals—a reasonable assumption for
to transition natural capital from a rhetorical device to a practical the High Plains Aquifer. We abstract from lateral flow and neighbor effects on
tool for fostering the sustainable management of our planet. pumping, which are, at most, secondary drivers of groundwater volume (55).
We find that groundwater withdrawal increases at an increasing rate with
Methods more water in the West Kansas High Plains Aquifer. MGðsÞ = 0 because hy-
We parameterize the case study for the period 1996–2005 (see Supporting drologic theory does not support stock-dependent recharge rates.
Information) using a 10-y data series from the Kansas Water Information In our base analysis, we model the average acre in Kansas. However, when
Management and Analysis System, the United States Department of Agri- we compute aggregate values for Kansas, we use the changes of all fields in
culture Economic Research Service, and the Kansas State University Agri- the dataset rather than the change in the average field. This provides the
cultural Extension, in conjunction with an annual time series of saturated most accurate measurement.
thickness (32).
We estimate MDðs,xðsÞÞ and MHIðs,xðsÞÞ based on the market conditions,
ACKNOWLEDGMENTS. D. Skelly and S. Yun provided helpful comments.
technology, and management in place at the time (35), where the economic K. Krause assisted with graphic design of Fig 1. E. Addicott provided research
program xðsÞ reflects decisions of crop choice and irrigation pumping as a assistance. R. Llewlyn helped us access historic crop budgets. The Knobloch
function of the groundwater stock under an acre (Tables S1–S4). MDðs,xðsÞÞ Family Foundation supported this research. E.M.K.H. was supported by
links the change in the water in the aquifer with the change in field-level net National Science Foundation WSC 1039180.

1. Guerry AD, et al. (2015) Natural capital and ecosystem services informing decisions: 31. Brauman KA, Daily GC, Duarte TK, Mooney HA (2007) The nature and value of eco-
From promise to practice. Proc Natl Acad Sci USA 112(24):7348–7355. system services: An overview highlighting hydrologic services. Annu Rev Environ Resour
2. United Nations (2014) Inclusive Wealth Report 2014, Measuring Progress Toward 32:67–98.
Sustainability (Cambridge Univ Press, Cambridge). 32. Haacker EMK, Kendall AD, Hyndman DW (May 26, 2015) Water level declines in the High
3. Hamilton K, Clemens M (1999) Genuine savings rates in developing countries. World Plains Aquifer: Predevelopment to resource senescence. Ground Water, 10.1111/gwat.12350.
Bank Econ Rev 13(2):333–356. 33. Simonit S, Perrings C (2013) Bundling ecosystem services in the Panama Canal wa-
4. Arrow KJ, Dasgupta P, Goulder LH, Mumford KJ, Oleson K (2012) Sustainability and tershed. Proc Natl Acad Sci USA 110(23):9326–9331.
the measurement of wealth. Environ Dev Econ 17:317–353. 34. Abbott JK, Klaiber HA (2013) The value of water as an urban club good: A matching
5. Pearson LJ, Biggs R, Harris M, Walker B (2013) Measuring sustainable development: approach to community-provided lakes. J Environ Econ Manage 65(2):208–224.
the promise and difficulties of implementing Inclusive Wealth in the Goulburn-Bro- 35. Pfeiffer L, Lin C-YC (2014) The effects of energy prices on agricultural groundwater
ken Catchment, Australia. Sustainability Sci Pract. Policy 9(1):16–27. extraction from the High Plains Aquifer. Am J Agric Econ 96(5):1349–1362.
6. World Bank (2011) The Changing Wealth of Nations (World Bank, Washington, DC). 36. Hartwick JM (1977) Intergenerational equity and the investing of rents from ex-
7. Polasky S, et al. (2015) Inclusive wealth as a metric of sustainable development. Annu haustible resources. Am Econ Rev 67(5):972–974.
Rev Environ Res 40:445–466. 37. Arrow K, et al. (2004) Are we consuming too much? J Econ Perspect 18(3):147–172.
8. Smulders S (2012) An arrow in the Achilles’ heel of sustainability and wealth ac- 38. Nordhaus WD (2006) Principles of national accounting for nonmarket accounts. A
counting. Environ Dev Econ 17(3):368–372. New Architecture for the U.S. National Accounts, eds Jorgenson DW, Landefeld JS,
9. Hanley N, Dupuy L, McLaughlin E (2015) Genuine savings and sustainability. J Econ Nordhaus WD (Univ Chicago Press, Chicago), pp 143–160.
Surv 29(4):779–806. 39. Arrow KJ, Dasgupta P, Maler K-G (2003) Evaluating projects and assessing sustainable
10. Barbier EB (2014) Economics: Account for depreciation of natural capital. Nature development in imperfect economies. Environ Resour Econ 26(4):647–685.
515(7525):32–33. 40. Boyd J, Banzhaf S (2007) What are ecosystem services? The need for standardized
11. Stiglitz JE, Sen A, Fitoussi J-P (2010) Mis-measuring Our Lives: Why GDP Doesn’t Add environmental accounting units. Ecol Econ 63(2-3):616–626.
Up—The Report by the Commission on the Measurement of Economic Performance 41. Daily GC, et al. (2009) Ecosystem services in decision making. Front Ecol Environ 7(1):21–28.
and Social Progress (New Press, New York). 42. Newell RG, Pizer WA (2003) Discounting the distant future: How much do uncertain
12. Dasgupta P (2009) The welfare economic theory of green national accounts. Environ
rates increase valuations? J Environ Econ Manage 46(1):52–71.
Resour Econ 42(1):3–38.
43. Arrow KJ, et al. (2014) Should governments use a declining discount rate in project
13. Dasgupta P (2014) Measuring the wealth of nations. Annu Rev Resour Econ 6:17–31.
analysis? Rev Environ Econ Policy 8(2):145–163.
14. Fenichel EP, Abbott JK (2014) Natural capital from metaphor to measurement. J Assoc
44. Office of Management and Budget (2003) Circular A-4: Regulatory Analysis (Exec Off
Environ Res Econ 1(1):1–27.
President, Washington, DC).
15. Polasky S, Tallis H, Reyers B (2015) Setting the bar: Standards for ecosystem services.
45. Carpenter SR, Brock WA, Ludwig D (2007) Appropriate discounting leads to foward-
Proc Natl Acad Sci USA 112(24):7356–7361.
looking ecosystem management. Ecol Res 22(1):10–11.
16. Schaefer M, Goldman E, Bartuska AM, Sutton-Grier A, Lubchenco J (2015) Nature as
46. Fenichel EP, et al. (2015) Wealth reallocation and sustainability under climate change.
capital: Advancing and incorporating ecosystem services in United States federal
Nat Clim Chang, in press.
policies and programs. Proc Natl Acad Sci USA 112(24):7383–7389.
47. Freeze RA, Cherry JA (1979) Groundwater (Prentice Hall, Englewood Cliffs, NJ).
17. Jorgenson DW (1963) Capital theory and investment behavior. Am Econ Rev 53(2):247–259.
48. Cederstrand JR, Becker MF (1998) Digital Map of Specific Yield for the High Plains
18. Dasgupta P (2007) Human Well-Being and the Natural Environment (Oxford Univ
Aquifer in Parts of Colorado, Kansas, Nebraska, New Mexico, Oklahoma, South Da-
Press, New York).
kota, Texas, and Wyoming (US Geol Surv, Boulder, CO), Open File Rep 98-414.
19. Liu J, et al. (2015) Sustainability. Systems integration for global sustainability. Science
49. Dhuyvetter KC, Taylor M (2014) Kansas Land Prices and Cash Rental Rates (Kansas
347(6225):1258832.
20. Barbier EB (2007) Valuing ecosystem services as productive inputs. Econ Policy 22(49): State Univ, Manhattan, KS).
177–229. 50. Pfeiffer L, Lin C-YC (2014) Does efficient irrigation technology lead to reduced
21. Fisher I (1906) The Nature of Capital and Income (Norwood, Norwood, MA). groundwater extraction? Empirical evidence. J Environ Econ Manage 67(2):189–208.
22. Polasky S, Segerson K (2009) Integrating ecology and economics in the study of 51. Donovan S, Goldfuss C, John H (2015) Incorporating Ecosystem Services into Federal
ecosystem services: Some lessons learned. Annu Rev Resour Econ 1:409–434. Decision Making (Exec Off President, Washington, DC).
23. Naidoo R, et al. (2006) Integrating economic costs into conservation planning. Trends 52. Gaffney M (2008) Keeping land in capital theory: Ricardo, Faustmann, Wicksell, and
Ecol Evol 21(12):681–687. George. Am J Econ Sociol 67(1):119–141.
24. Barbier EB (2011) Pricing nature. Annu Rev Resour Econ 3:337–353. 53. Halvorsen R, Smith TR (1984) On measuring natural resource scarcity. J Polit Econ
25. Bockstael NE, Freeman AMI, Korp RJ, Portney PR, Smith VK (2000) On measuring 92(5):954–964.
economic values for nature. Environ Sci Technol 34(8):1384–1389. 54. Maler K-G, Aniyar S, Jansson A (2009) Accounting for ecosystems. Environ Resour Econ
26. Costanza R, et al. (1997) The value of the world’s ecosystem services and natural 42(1):39–51.
capital. Nature 387(6230):253–260. 55. Pfeiffer L, Lin C-YC (2012) Ground water pumping and spatial externalities in agri-
27. Steward DR, et al. (2013) Tapping unsustainable groundwater stores for agricultural culture. J Environ Econ Manage 64:16–30.
production in the High Plains Aquifer of Kansas, projections to 2110. Proc Natl Acad 56. Dasgupta P, Maler K-G (2000) Net national product, wealth, and social well-being.
Sci USA 110(37):E3477–E3486. Environ Dev Econ 5(1):69–93.
28. Morris BL, et al. (2003) Groundwater and Its Susceptibility to Degradation: A Global As- 57. Dasgupta P, Maler K-G, Barrett S (1999) Intergenerational equity, social discount rates
sessment of the Problem and Options for Management (United Nations Environ Pro- and global warming. Discounting and Intergenerational Equity, eds Portney P,
gramme, Nairobi). Weyant J (Resour Future, Washington, DC), pp 51–77.
29. Daily GC, et al. (2000) Ecology. The value of nature and the nature of value. Science 58. Miranda MJ, Fackler PL (2002) Applied Computational Economics and Finance (MIT
289(5478):395–396. Press, Cambridge, MA).
30. Keeler BL, et al. (2012) Linking water quality and well-being for improved assessment 59. Press WH, Teukolsky SA, Vetterling WT, Flannery BP (2007) Numerical Recipes: The Art
and valuation of ecosystem services. Proc Natl Acad Sci USA 109(45):18619–18624. of Scientific Computing (Cambridge Univ Press, New York), 3rd Ed.

6 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1513779113 Fenichel et al.


Supporting Information
Fenichel et al. 10.1073/pnas.1513779113
Derivation of Eq. 1 For expositional clarity, we replace the subscripts that indicated
The derivation of Eq. 1, which appears in Fig. 1, follows refs. 14, marginal impacts of the stock with M to get
39, and 56. We first define how the resource changes over time,
_
MDðs, xðsÞÞ + pðsÞ
pðsÞ = , [S9]
ds=dt = s_ = GðsðtÞÞ − HIðsðtÞ, xðsðtÞÞÞ. [S1] δ − ½MGðsÞ − MHIðs, xðsÞÞ

G is a growth or recharge function, which is fundamentally an which is Eq. 1.


ecological or biophysical equation. HI is the human impact func-
tion, illustrated by the yellow arrow from human behavior to Function Approximation
growth rate of capital stock (Fig. 1). Next, define DðsðtÞ, xðsðtÞÞÞ Approximating Eq. 1 enables us to develop a price function over
as an index of the net benefits, or dividends, flowing to soci- all possible values of the state variable conditional on the cur-
ety as “income” at instant t. We assume that DðsðtÞ, xðsðtÞÞÞ is rent economic program in a single step. The economic program
measured in stable monetary terms so that discounting at is assumed to continue indefinitely. Any changes in the economic
a constant exponential rate is justified (57). The net present program (as through institutional change) must be embedded
value of benefits generated by natural capital, evaluated at within the model. The embodiment of existing institutions in
time t, is
the economic program often makes these prices the most ap-
propriate for inclusive wealth accounting. However, valuation
Z∞
under alternative assumptions about management institutions,
V ðsðtÞÞ = e−δðτ−tÞ DðsðτÞ, xðsðτÞÞÞdτ. [S2] ecosystem service, etc., can be incorporated through different
t functional assumptions in Eq. 1 (14).
Eq. 1 contains two unknowns, p and p._ If either were known,
V can be expressed solely as a function of sðtÞ by substituting the then computing the accounting price of natural capital would be
economic program, xðsÞ, into D, assuming an infinite planning trivial. Fenichel and Abbott (14) approximate p._ However, this
horizon and that calendar date does not matter explicitly.
requires the HI function to be twice differentiable, a condition
Slightly abusing notation, we define the partial derivative of D
that will not hold if the economic program ultimately leads to
to include the feedback through the economic program,
resource exhaustion, as is the current case (between 1996 and 2005,
water was always withdrawn faster than recharge in the Kansas
∂D ∂D dx
DS ðsðtÞ, xðsðtÞÞÞ = + = DpS = MD, [S3] system). Therefore, the approach in ref. 14 is not fully general.
∂s ∂x ds To provide a general approach, we approximate p ≈ μðsðtÞÞ =
P k−1
where the asterisk indicates the elimination of x, and M simplifies n=0 βn ϕn ðsðtÞÞ, where μðsðtÞÞ is a Chebyshev polynomial in s,
notation and indicates “marginal.” The accounting price for cap- with k basis functions that span the state space (58). Here
ital, conditional on the existing institutions, is ϕn ðsðtÞÞ are the basis functions, and the βn are a set of param-
eters determining the weighting of these basis functions (58).
pðsðtÞÞ ≡ ∂V ðsðtÞÞ=∂sðtÞ. [S4] The other missing value, p,_ is approximated _
P as p_ ≈ μs ðsðtÞÞsðtÞ,
assuming time autonomy, where μs ðsðtÞÞ = k−1 n=0 β n Δϕn ðsðtÞÞ and
The accounting price is how much society gains today and in per- Δϕn ðsðtÞÞ indicates the derivative of the basis function with
petuity from a marginal increase in the stock of capital. Follow- respect to sðtÞ. Approximating p only requires the information
ing refs. 56 and 39, we differentiate V ðsðtÞÞ with respect to t and in Eq. 1, whereas approximating p_ requires higher-order
express it as derivatives.
The approximation problem is a matter of finding the βn co-
dV ∂V ds ds efficients that minimize the error introduced by the approxima-
= δV − Dðs, xðsÞÞ = = pðsðtÞÞ . [S5] tion. We approximate the polynomial μðsðtÞÞ at N nodes. These
dt ∂s dt dt
nodes are distributed as the roots of a Chebyshev polynomial
It follows that over the state space. The combination of Chebyshev nodes and
polynomials distributes the error between the approximating
δV = Dðs, xðsÞÞ + pðsðtÞÞs_ = H p ðs, pÞ, [S6] and unknown true function evenly, resulting in the best poly-
nomial specification for functional approximation (59).
where H p ðs, pÞ is the current value Hamiltonian evaluated with Fenichel and Abbott (14) used numerical solvers to conduct the
the economic program xðsÞ, composed of the flow of current approximation. However, this is unnecessarily complicated. Let
benefits (dividends), Dðs, xðsÞÞ, and the value of increments to the vector of stock-specific approximations of the accounting
the stock, ps._ Rearranging Eq. S6 yields price at each approximation node (while suppressing the de-
pendence on the stock) be written as
Dðs, xðsÞÞ + pðsðtÞÞs_
V ðsÞ = . [S7]
δ p ≈ ϕn β = diagðδ − s_s Þ−1 ðW s + diagðsÞΔϕ
_ n βÞ [S10]
Differentiating V ðsÞ with respect to s and rearranging yields
where ϕn and Δϕn are N × k matrices, and the other factors are
N × 1 vectors (except for the scalar discount rate). This expres-
_
Ds ðs, xðsÞÞ + pðsÞ sion is linear in β. Assuming N > k, the least-squares solution
pðsÞ = . [S8]
δ − ðGs ðsÞ − HIs ðs, xðsÞÞÞ for β is

Fenichel et al. www.pnas.org/cgi/content/short/1513779113 1 of 2


 Projection and Calibration
T
β = ðdiagðδ − s_s Þϕn − diagðsÞΔϕ
_ nÞ The regression models were used to project MHIðs, xðsÞÞ and
−1 MDðs, xðsÞÞ, conditional on the economic program for water
3ðdiagðδ − s_s Þϕn − diagðsÞΔϕ
_ nÞ withdrawal, xðsÞ for the representative parcel. Water withdrawal
  is modeled as a function of crop choice. Following estimation of
3 ðdiagðδ − s_s Þϕn − diagðsÞΔϕ
_ T the multinomial logit model of crop choice, the probability that
nÞ W s [S11]
crop mix j was selected conditional on water in the aquifer s is
 
In the current application, we find that a 10th-order polynomial   exp aj + bj s
together with 100 nodes is sufficient to recover accounting prices P CSj = P   [S12]
1 + j≠1 exp aj + bj s
such that adding basis functions or nodes does not influence the
accounting price. We recover year-specific accounting prices
by approximating using year-specific means for the explanatory where field type j = 1 is the index field type. The constant aj
variables other than volume of groundwater. includes all control variables evaluated at their means, including
technological adjustments for drop nozzles, multiplied by their
Parameter Estimation respective crop mix-specific coefficients. Fields with less water
Prior research suggests that saturated thickness or water in the tend to have more unirrigated land. Large unirrigated fields were
used as the omitted baseline case (with a numerator of 1). Eq.
aquifer influences crop choice and water withdrawal, and hence
S12 can be expressed as a 25 × 1 vector PðCSÞ of shares for each
MHI and MD (35, 50). Estimating MDðs, xðsÞÞ and MHIðs, xðsÞÞ
crop mix. Let κ be a 25 × 6 matrix of acreage allocations by field
requires internally consistent models of planting decisions, type. The first five columns are the dominant irrigated crops:
water withdrawal, xðsÞ, and net revenue. Water withdrawal, or alfalfa, corn, sorghum, soybean, and wheat. The sixth column is
the derived demand for water, is a direct function of water in unirrigated. APFðsÞ = ðPðCSÞÞT κ, gives a 1 × 6 row vector of the
the aquifer and indirectly a function of water in the aquifer expected acres per field of each crop (T indicates transpose).
through its influence on crop choice. We closely follow Pfeiffer Next, we use the acres planted on the representative field of a
and Lin’s (35) linear regression approach by estimating ground- given crop system type to model the representative water with-
water withdrawal as a function of crop choice and water in the drawal per acre (the economic program) as
aquifer, as well as other control variables. Data come from the
2
Kansas Water Information Management and Analysis System eα+βs+APFðsÞγ1 +APFðsÞ γ2

and the US Department of Agriculture Economic Research xðsÞ = P​ [S13]


APFj
Service spanning 1996 through 2005. However, we modify
Pfeiffer and Lin’s early work in three ways. First, base mea- where the numerator is the prediction from a water withdrawal
surements of water in the aquifer use new, detailed estimates in regression on a log scale. The squared operation is element by
ref. 32 and location-dependent specific yield measures (48) element, and α includes all other explanatory variables evaluated
rather than saturated thickness alone. Second, we account for at their means from the water withdrawal regression and a back
pumping constraints using a dummy variable for saturated transformation adjustment for Jensen’s inequality when estimat-
thickness less than 29.5 feet (9 m), which generally means the ing on the log scale; β is the estimated coefficient for water in the
rate of groundwater flow to wells is too low to support pump- aquifer, and γ 1 and γ 2 are column vectors of estimated coeffi-
ing. Third, we estimated the groundwater withdrawal function cients for crop-specific acres planted and acres planted squared.
on a log scale. Other variables in the regression are included Eq. S13 shows that water withdrawal depends on water in the
in Table S3. aquifer both directly and indirectly through links to decisions
Water withdrawal and marginal net revenue depend on crop about crop mix. Although the regression fits the data well, we
choice and production. Our goal is to model a representative acre, do not force predicted withdrawals to be less than the water
so we assume that, conditional on crop choice, farmers realize stock. Therefore, we impose a constraint that xðsÞ ≤ s.
that crop’s expected yield. However, groundwater availability MDðs, xðsÞÞ is the change in net revenue for a marginal change
influences crop choice. We model crop mix by field as a function in groundwater. We model revenue per acre for each crop as the
of groundwater availability using a multinomial logit model using median crop-specific yield per acre multiplied by crop prices,
the top 25 field types, which accounts for more than 98% of the adjusted for direct subsidies in 2005 dollars and less variable costs
acres in the data. A field type is defined as a crop mix and the based on crop budgets. This provides the social value of crops
average areas covered by the crops in that mix. Crop mixes in- produced with groundwater. Price and cost data were taken from
clude fields of alfalfa, corn, soybean, sorghum, wheat, and un- crop budgets from Kansas State University Cooperative Exten-
irrigated land. A field type can be 100% of any of these cover sion. All prices and costs were adjusted to 2005 dollars using crop
types or combinations thereof. An unirrigated field type was specific producer price indices from the US Bureau of Labor
used as the index field. Other variables included in the re- Statistics (www.bls.gov/ppi/data.htm). We assume that all acres
gression are included in Table S4. produce the median yield of a given crop per acre.

Other Supporting Information Files

Table S1 (DOCX)
Table S2 (DOCX)
Table S3 (DOCX)
Table S4 (DOCX)

Fenichel et al. www.pnas.org/cgi/content/short/1513779113 2 of 2

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