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FUNDAMENTALS
OF ECOSYSTEM SCIENCE
FUNDAMENTALS
OF ECOSYSTEM
SCIENCE
SECOND EDITION
Edited by
KATHLEEN C. WEATHERS
DAVID L. STRAYER
GENE E. LIKENS
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than as may be noted herein).
Notices
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our
understanding, changes in research methods, professional practices, or medical treatment may become
necessary.
Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using
any information, methods, compounds, or experiments described herein. In using such information or methods
they should be mindful of their own safety and the safety of others, including parties for whom they have a
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Library of Congress Cataloging-in-Publication Data
A catalog record for this book is available from the Library of Congress
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
ISBN 978-0-12-812762-9
Elena M. Bennett Bieler School of Environment Gary M. Lovett Cary Institute of Ecosystem
and Department of Natural Resource Sciences, Studies, Millbrook, NY, United States
McGill University, Montreal, QC, Canada Pamela A. Matson Stanford University,
Mary L. Cadenasso Department of Plant Stanford, CA, United States
Sciences, University of California, Davis, Judy L. Meyer Odum School of Ecology,
United States University of Georgia, Athens, GA, United
Cayelan C. Carey Virginia Tech, Blacksburg, States
VA, United States Richard S. Ostfeld Cary Institute of Ecosystem
Jonathan J. Cole Cary Institute of Ecosystem Studies, Millbrook, NY, United States
Studies, Millbrook, NY, United States Michael L. Pace Department of Environmental
Holly A. Ewing Bates College, Lewiston, ME, Sciences, University of Virginia, Charlottesville,
United States VA, United States
Stuart E.G. Findlay Cary Institute of Ecosystem Steward T.A. Pickett Cary Institute of
Studies, Millbrook, NY, United States Ecosystem Studies, Millbrook, NY, United
Robinson W. Fulweiler Department of Earth States
and Environment, Department of Biology, Emma J. Rosi Cary Institute of Ecosystem
Boston University, Boston, MA, United States Studies, Millbrook, NY, United States
Peter M. Groffman Cary Institute of Ecosystem Meagan E. Schipanski Department of Soil and
Studies, Millbrook; City University of New Crop Sciences, Colorado State University, Fort
York, Advanced Science Research Center at Collins, CO, United States
the Graduate Center and Brooklyn College Christopher T. Solomon Cary Institute of
Department of Earth and Environmental Ecosystem Studies, Millbrook, NY, United
Science, New York, NY, United States States
Stephen K. Hamilton Cary Institute of Emily H. Stanley University of Wisconsin-
Ecosystem Studies, Millbrook; Kellogg Madison, WI, United States
Biological Station and Department of
David L. Strayer Cary Institute of Ecosystem
Integrative Biology, Michigan State
Studies, Millbrook, NY; Graham Sustainability
University, Hickory Corners, MI, United States
Institute, University of Michigan, Ann Arbor,
Oleksandra Hararuk University of Central MI, United States
Florida, Orlando, FL, United States
R. Quinn Thomas Virginia Tech, Blacksburg,
Clive G. Jones Cary Institute of Ecosystem VA, United States
Studies, Millbrook, NY, United States
Kathleen C. Weathers Cary Institute of
Gene E. Likens Cary Institute of Ecosystem Ecosystem Studies, Millbrook, NY, United
Studies, Millbrook, NY; University of States
Connecticut, Storrs, CT, United States
ix
Preamble to the 2nd Edition
As we write this, the world is gripped by a ecosystems in the form of changes in eco-
devastating pandemic (COVID-19). The nomic demand, trade patterns, pollutant re-
socioecological causes, responses to, leases, and so on. Thus, the COVID-19
and ramifications of COVID-19 have been pandemic reminds us that diseases are an
and will be profound, and they will play ecological problem as well as a medical or
out over short- and long-term scales. Because veterinary problem, and their prevention
we are writing in the midst of this pandemic, and solution will require understanding
the specifics about its ultimate resolution and application of ecological ideas
and effects are far from clear. What is clear and principles. Although pandemics such
from the COVID-19 and previous pan- as COVID-19 have had analogs in the
demics, though, is that ecosystems can affect past (e.g., Black Death in the 1300s, influenza
the emergence and spread of disease (as in 1918), conditions are far different now
discussed by Rick Ostfeld in his essay in this than then—from the number
book), and infectious diseases can have enor- and distribution of people on the planet,
mous effects on ecosystems. Human impacts to the degradation and change of ecosys-
such as habitat alteration, rapid global move- tems worldwide from human activities
ment of plants, animals, people, and the interconnectedness of natural
and pathogens, and the size and density and human systems. These factors may tend
of the human population itself favor the to make future pandemics more frequent
emergence and spread of disease. In turn, and more severe, and lend urgency to the
as we are seeing in this time of COVID-19, effective application of ecosystem science
diseases have such strong effects on human and other sciences to understanding as
activities that they can feed back onto well as the management of our world.
xi
Preface
This book provides an introduction to the “voices” will be evident throughout the
content, ideas, and major findings of contem- book. We believe that this diversity reflects
porary ecosystem science. We wrote the book some of the myriad perspectives and ap-
primarily for beginning graduate students proaches that are fruitfully brought to bear
and advanced undergraduates, but it should on the field of ecosystem science.
also be useful to a broad range of academic The book contains six major sections. The
scientists and resource managers, and even opening chapter introduces the concept of
to dedicated amateurs who seek an intro- the ecosystem, explores some of the conse-
duction to the field. Ecosystem science is a quences of this concept, describes the intel-
rigorous, quantitative science; we assume lectual tools of the science, and briefly
that readers of the book will have had an reviews the history of this young science.
introductory class in ecology and basic Chapters 2–9 lay the foundation for the study
understanding of chemistry and math. The of ecosystems, and cover the two major
book deliberately covers multiple approa- branches of ecosystem science: energetics
ches to understanding ecosystems (e.g., the (Chapters 2–5) and biogeochemistry
use of experiments, theory, cross-system (Chapters 5–9). These chapters present the
comparisons), in multiple environments (ter- core content of ecosystem science—the
restrial, freshwater, and marine; managed, movement and fate of energy and materials
built, and natural ecosystems), across all parts in ecosystems—in some detail. In the syn-
of the world (although many examples come thetic Chapters 10–12, we revisit major
from the authors’ experience in North themes that cut across multiple areas of
America). study in ecosystem science. The authors of
The origins of this book stem from an these chapters review the power and utility
intensive 2-week Fundamentals of Ecosystem of the ecosystem concept, the roles of hetero-
Ecology class (the FEE class) that we have geneity in space and time, and the impor-
taught to graduate students from around tance of various types of controls in
the world every year or two at the Cary Insti- ecosystems. Chapters 13–18 take ecosystem
tute of Ecosystem Studies since 1989. We, and science into the real world by illustrating,
many of the chapter authors, have played cen- through six case studies, the value of ecosys-
tral roles in the development, evolution, and tem science in identifying and solving
running of the FEE class since its origin. a range of environmental problems. The
We decided upon an edited book for sev- book closes with Chapter 19, which lays
eral reasons, not the least of which was its out some challenges and needs for the future.
genesis in this team-taught course. While Today’s ecosystem science is evolving rap-
we shepherded and integrated the chapters idly, with major new discoveries and ideas
and their contents, we also deliberately emerging every year. The ultimate shape
allowed—and even encouraged—multiple and contributions of this science remain to
approaches, and as a result, multiple be discovered.
xiii
xiv Preface
This book benefited from the persistent their scholarship, patience, goodwill, and
and hard work of the Academic Press team, commitment to bringing this project to fru-
especially Jill Cetel, Candice Janco, and ition. The Cary Institute’s assistant, Matt
multiple graphic artists. We were also fortu- Gillespie, was an enormous help as well.
nate to have received helpful and critical Finally, generations of FEE students were
reviews of chapters from colleagues, includ- and continue to be an impetus and inspira-
ing Alexandra Ponette-González, Clifford tion to us and the field of ecosystem science.
Ochs, and several anonymous reviewers
who teach ecosystem science; their com- Kathleen C. Weathers
ments substantially improved the book. We David L. Strayer
thank the authors of various chapters for Gene E. Likens
C H A P T E R
1
Introduction to Ecosystem Science
Kathleen C. Weathersa, David L. Strayera,
and Gene E. Likensa,b
a
Cary Institute of Ecosystem Studies, Millbrook, NY, United States bUniversity of Connecticut,
Storrs, CT, United States
Humans have devised many intellectual systems to understand and manage the
complicated world in which we live, from physics to philosophy to economics. In this book,
we present one such intellectual system, ecosystem science, which tries to make sense of the
complex natural world and helps us manage it better. As we will see, ecosystems can be
highly varied in size and character, from a little pool of water in a tree cavity, to a redwood
forest, to a neighborhood in a city, to a frigid river, to the entire globe (Figure 1.1).
Nevertheless, a common set of tools and ideas can be used to analyze and understand these
varied and complicated systems. The results of these analyses are both intellectually
satisfying and useful in managing our planet for the benefit of nature and humankind.
Indeed, because of the growing demands placed on living and nonliving resources by
humans, it has been argued that ecosystem science is one of the essential core disciplines
needed to understand and manage the modern planet Earth (Weathers et al. 2016).
This book defines the ecosystem, illustrates the ecosystem approach, describes the chief
characteristics of ecosystems and the major tools that scientists use to analyze them, and
presents important discoveries that scientists have made about ecosystems. It also lays out
some critical questions for the future. Although the book is not focused on the management
of ecosystems, several management implications of ecosystem science are described and
illustrated.
What Is an Ecosystem?
An ecosystem is the interacting system made up of all the living and nonliving objects in a
specified volume of space.
This deceptively simple definition both says much and leaves out much. First, as with other
systems (Box 1.1), ecosystems contain more than one object, and those objects interact. More
Fundamentals of Ecosystem Science 3 Copyright # 2021 Elsevier Inc. All rights reserved.
https://doi.org/10.1016/B978-0-12-812762-9.00001-0
4 1. Introduction to Ecosystem Science
FIGURE 1.1 Some examples of ecosystems: (A) the frigid Salmon River, Idaho; (B) a residential neighborhood in
Baltimore, Maryland; (C) a biofilm on a rock in a stream; (D) a section of the southern ocean containing a phytoplank-
ton bloom; (E) a redwood forest in the fog in California; (F) a tree cavity; (G) the Earth. (Photocredits: A—John Davis;
B—Baltimore Ecosystem Study LTER; C—Colden Baxter; D—US government, public domain; E—Samuel M. Simkin;
F—Ian Walker; G—Source: https://www.publicdomainpictures.net/en/view-image.php?image=86448&picture=
planet-earth.)
surprisingly, living and nonliving objects are given equal status in ecosystem science.
A particle of clay and the plant drawing its nutrition from that clay particle are both parts
of an ecosystem, and therefore equally valid objects of study. This viewpoint contrasts with
physiology and population ecology, for example, in which the organism is the object of study,
I. Introduction
What Is an Ecosystem? 5
BOX 1.1
and the nonliving environment is conceived of as an external influence on the object of study.
Finally, the definition implies that ecosystems have definite boundaries, but does not tell us
how we might go about setting or finding the boundaries to an ecosystem.
There are some unexpectedly powerful advantages to this simple definition. First, by in-
cluding all living and nonliving objects in a specified space, it is possible to use the tool of
mass balance to follow the movement and fate of materials (Box 1.2). Material that comes into
an ecosystem must either stay in the ecosystem or leave—there is no other possible fate for the
material. Mass balance offers a convenient quantitative tool for measuring the integrated
activity of entire, complicated systems without having to measure the properties and inter-
actions of each of its parts. It also allows ecosystem scientists to estimate the size of a single
unknown flux by difference. Consequently, it will become evident throughout the book that
ecosystem scientists often use the powerful tool of mass balance.
I. Introduction
6 1. Introduction to Ecosystem Science
BOX 1.2
I. Introduction
What Is an Ecosystem? 7
Modified from Millennium Ecosystem Assessment (2003, 2005) and Costanza et al. 2017.
Second, defining an ecosystem as we have done makes it possible to measure the total ac-
tivity of an ecosystem without having to measure all the parts and exchanges within the eco-
system. This advantage is sometimes referred to as a “black-box” approach, because we can
measure the function (input and output) of a box (the ecosystem) without having to know
what is in the box (Figure 1.2). Sometimes ecologists debate whether it is philosophically pos-
sible to predict the properties of a complex system by studying its parts (reductionism) or
whether it is necessary to study intact systems (holism). It is not necessary to accept the phil-
osophical claims of holism, though, to recognize that studies of whole systems may be a much
more efficient way than reductionism to understand ecosystems. Such a holistic approach to
ecosystems is a powerful tool of ecosystem science, and is often combined with reductionist
approaches to develop insights into the functioning and controls of ecosystems.
ECOSYSTEM BOUNDARY
ORGANIC COMPARTMENT
ATMOSPHERIC Litter
COMPARTMENT Biological uptake
Meteorologic Meteorologic
of gases and aerosols Herbicore Geologic Geologic
Windblown particulates Living Dead
and gases above and
Biological release of Plant
Carnivore
Biomass
Biologic Biologic
below ground
gases and organic aerosols Omnivore
Biomass
INPUT INPUT
Detritivore
W
et
leaching, throughfall,
an
Dry deposition
stemflow, exudation
d
dr
Mineralization,
Inorganic
aerosols
yd
Re ep
Biological
e itio
of n
ga
ses
OUTPUT OUTPUT
AVAILABLE NUTRIENT
SOIL AND ROCK Weathering COMPARTMENT
MINERAL Meteorologic Meteorologic
COMPARTMENT
on in Geologic Geologic
Formation of exchange soil Biologic Biologic
secondary minerals sites solution
INTRASYSTEM CYCLE
FIGURE 1.2 Two views of the same ecosystem. The left side shows some of the parts inside an ecosystem and how
they are connected, as well as the exchanges between the ecosystem and its surroundings, whereas the right side
shows a black-box approach in which the functions of an ecosystem (i.e., its inputs and outputs) can be studied with-
out knowing what is inside the box. (Modified from Likens 1992.)
I. Introduction
8 1. Introduction to Ecosystem Science
Third, the definition gives the investigator complete flexibility in choosing where to set the
boundaries of the ecosystem in time and space. The boundaries of an ecosystem (i.e., size,
location, and timescale) can therefore be chosen to match the question that the scientist is try-
ing to answer. Boundaries often are drawn at places where fluxes are easy to measure (e.g., a
single point on a stream as it leaves a forested, watershed-ecosystem) or so that fluxes across
the boundary are small compared to cycling inside the ecosystem (e.g., a lake shore). Never-
theless, boundaries are required to make quantitative measures of these fluxes. It is true that
ecosystems frequently are defined to be large (e.g., lakes and watersheds that are hectares to
square kilometers in size) and are studied on the scale of days to a few years, but there is noth-
ing in the definition of an ecosystem that requires ecosystems to be defined at this scale. In-
deed, as we will see, an ecosystem may be as small as a single rock or as large as the
entire Earth (see Chapter 7), and can be studied for time periods as long as hundreds of mil-
lions of years.
Fourth, defining an ecosystem to contain both living and nonliving objects recognizes the
importance of both living and nonliving parts of ecosystems in controlling the functions and
responses of these systems. There are examples throughout the book in which living organ-
isms, nonliving objects, or both acting together determine what ecosystems look like (struc-
ture) and how they work (function). Furthermore, the close ties and strong interactions
between the living and nonliving parts of ecosystems are so varied and so strong that it would
be very inconvenient if not misleading to study one without the other. Thus, the inclusion of
living and nonliving objects in ecosystems has practical as well as intellectual advantages.
Finally, we note one further property of ecosystems: they are open to the flow of energy
and materials. It might be theoretically possible to define particular examples of ecosystems
that are closed systems, not exchanging energy or materials with their surroundings, but
nearly all ecosystems as actually defined have important exchanges of energy and materials
with their surroundings. Indeed, such exchanges are one of the central subjects of ecosystem
science. We note in particular that most ecosystems depend on energy inputs from external
sources, either as energy from the sun or as organic matter brought in from neighboring
ecosystems.
Now consider briefly what is missing from the definition. We have already noted that the
definition does not specify the time or space scales over which an ecosystem is defined, or
where exactly the boundaries are placed. Ecosystems are not required to be self-regulating, per-
manent, stable, or sustainable. They are not required to have any particular functional prop-
erties. For example, they need not be in balance or efficient in the way that they process
materials. Our definition does not require ecosystems to have a purpose. Although ecosystems
change over time, the basic definition does not suggest anything about the nature or direction of
that change. It might seem like a shame not to include such interesting attributes in a definition
of ecosystem (O’Neill 2001), and indeed some ecologists have incorporated such attributes
in their definitions, but we think it is neither necessary nor helpful to include them in a basic
definition. They may, however, be useful hypotheses and the subject of fruitful research pro-
jects. For instance, we might hypothesize that as forest ecosystems recover from disturbances
like fire or clear-cutting, they retain a higher proportion of nutrient inputs from precipitation
or release from weathering substrates. This viewpoint is quite different than saying that
ecosystems are systems that tend to maximize efficiency of use of limiting nutrients.
I. Introduction
What Are the Properties of Ecosystems? 9
What Is in an Ecosystem?
We might begin simply by listing the contents of an ecosystem. Plants and animals occur in
most ecosystems. As we will see later in the book, the number and kinds of plants and animals
can have a strong influence on ecosystem function. Many ecosystems also contain people.
Historically, many ecologists treated humans as outside of the ecosystem, or deliberately
studied ecosystems without people, but it is now common to treat people and our institutions
as parts of ecosystems (e.g., Pickett et al. 2001, 2011; McPhearson et al. 2016). Certainly the
structure and function (and change) of many modern ecosystems cannot be understood with-
out considering human activities.
Almost all ecosystems contain microbes (bacteria and fungi); although these are not as con-
spicuous as plants and animals, their activities are vital to ecosystem functioning. Viruses oc-
cur in most ecosystems, and may play important roles as regulators of plant, animal, and
microbial populations. Ecosystems also contain water and air, which are themselves re-
sources for many organisms and also serve as media in which organisms and nonliving ma-
terials can be transported, both within and across the boundaries of ecosystems. Finally,
ecosystems contain an enormous variety of nonliving materials, organic and inorganic, solid
and dissolved. These nonliving materials, including such disparate items as dead wood, clay
particles, bedrock, oxygen, and dissolved nutrients, interact with the living biota and exercise
strong influences on the character and functioning of ecosystems. Thus, the total inventory of
an ecosystem can be very long; it might contain thousands or millions of kinds of items, living
and nonliving, and countless numbers of individuals in these “kinds.”
I. Introduction
10 1. Introduction to Ecosystem Science
often describe ecosystems by their size or location (e.g., latitude, altitude, biogeographic
realm, or distance from the coast).
I. Introduction
What Are the Properties of Ecosystems? 11
the P/R (gross photosynthesis to respiration) ratio for the system. At a steady state, ecosystems
with a P/R ratio less than 1 must import chemical energy (usually organic matter) from neigh-
boring ecosystems and are called heterotrophic; those with a P/R ratio greater than 1 export
chemical energy to neighboring ecosystems and are called autotrophic. Another useful
functional description is the residence time of a given material in an ecosystem—that is, the
average amount of time that a material spends within an ecosystem. Residence time is cal-
culated by dividing the standing stock of the material in the ecosystem by its input rate.
Ecosystem structures and functions can have economic value. For instance, ecosystems
provide lumber, they purify water and air, they regulate the prevalence of human diseases,
and they provide pollination for crop plants. These and many other goods and services
provided by ecosystems are commonly called “ecosystem services”—the benefits that people
derive from ecosystem structures and functions (e.g., Daily 1997; Millennium Ecosystem
Assessment 2005; Kareiva et al. 2011, Box 1.2). Developing ways to estimate quantitatively
the value of ecosystem services is an important and growing field at the intersection of
ecology, sustainability science, and economics.
I. Introduction
12 1. Introduction to Ecosystem Science
Scientists have been motivated to study ecosystems for several reasons. To begin with, if
ecosystems truly are the “basic units of nature” on Earth (Tansley 1935), any attempt to
understand our planet and the products of evolution on it must include ecosystem science
as a central theme. Indeed much study of ecosystem science has been motivated by simple
curiosity about how our world and how systems—whether ecological, social, or socio-
ecological—work. Many salable products such as timber and fish are taken directly from
“wild” ecosystems, so many early ecosystem studies were carried out to try to understand
better the processes that supported these products and ultimately increase their yields. Espe-
cially in the past 30 years, we have come to realize that the valuable products of nature include
far more than obviously salable products like timber and fish. Ecosystems also provide us
with clean air and water, opportunities for recreation and spiritual fulfillment, protection
from diseases, and many more “ecosystem services” (Box 1.2). Human economies and
well-being are wholly embedded in and dependent on wild ecosystems. Thus, many contem-
porary ecosystem studies are concerned with how ecosystems provide this broad array of
services, how human activities reduce or restore the ability of ecosystems to provide these
services, and ultimately how to reconcile the growing demands of human populations with
the needs of both nature and ourselves for functioning ecosystems.
Depending on the problem that they are studying, ecosystem scientists use a wide variety
of approaches and an array of simple to sophisticated tools to measure different aspects of
ecosystem structure and function. We offer a few examples here; however, new approaches
and tools emerge every year, and with them come more ways to open black boxes in ecosys-
tem science (see Chapter 19).
I. Introduction
How Do Ecosystem Scientists Learn about Ecosystems? 13
TABLE 1.2 Strengths and limitations of approaches to understanding ecosystems. Natural history observations
and understanding underpin all of these approaches.
Approach Some Strengths Some Limitations
Theory Flexibility of scale; integration; deduction of Cannot develop without linkage to
testable ideas observation and experiment
Long-term observation Temporal context; detection of trends and Potentially site-specific; difficult to
surprises; test hypotheses about temporal determine cause; costly and difficult to
variation maintain
Comparison Spatial or interecosystem context; detection of Difficult to predict temporal change or
spatial pattern; test hypotheses about spatial response to perturbation
variation
Ecosystem experiment Measure ecosystem response to perturbation; Potentially site-specific; potentially
test hypothesis about controls and difficult to rule out some explanations;
management of ecosystem processes hard to do; expensive
including: (1) natural history or observation; (2) theory and conceptual models; (3) long-term
study; (4) cross-ecosystem comparison; and (5) experiments. These approaches are comple-
mentary to one another (Table 1.2), and are best used in combination. Almost every scientific
question of any complexity or importance in ecosystem science requires the use of two or
more of these approaches to get a robust answer.
Natural History
A good deal can be learned about ecosystems simply from watching them and
documenting what is observed in some fashion. Do fallen leaves decay in place, wash away
into a stream, or burn in episodic fires? Is the soil deep and rich, or shallow and rocky? Does it
freeze in the winter? As a result, our understanding of an ecosystem often is based on simple
observations of its natural history. Indeed, without such careful observations, even the most
sophisticated studies can go astray by formulating nonsensical questions or omitting key ob-
servations or measurements. Not surprisingly, careful natural history studies (such as Forbes’
“The Lake as a Microcosm,” discussed later) were important precursors to modern ecosystem
science. Although these forerunners of ecosystem science often included speculation about
ecosystem processes, they did not have the technical means to measure easily such functions
as net ecosystem productivity or nutrient cycling, or to quantify trophic transfers.
Long-Term Studies
Long-term studies (i.e., those lasting for more than 10 years—longer than the tenure of
most grants or the time it takes to earn a PhD!) are relatively rare in ecology as a whole
(Lindenmayer and Likens 2018). However, long-term studies are especially good at providing
insight into slow processes (e.g., changes associated with forest succession), subtle changes
(e.g., changing chemistry of precipitation), rare events (e.g., the coldest winter, effects of hur-
ricanes or insect outbreaks), or processes controlled by multiple interacting factors (e.g., fish
I. Introduction
14 1. Introduction to Ecosystem Science
recruitment; Likens 1989; Lindenmayer and Likens 2018; and see the Long-Term Ecological
Research Program (LTER) of the National Science Foundation, http://www.lternet.edu).
Sometimes long-term understanding can be obtained by short-term analyses of materials that
record history, such as soil or sediment cores, otoliths (fish ear-stones), or written historical
records. For example, analysis of pollen, diatoms, pigments, or geochemistry in lake sediment
can reveal the millennium-long history of terrestrial vegetation, phytoplankton, soils, and
lake level—in short, the history of the development of the linkages between terrestrial and
aquatic ecosystems. It is from long-term studies or their surrogates that scientists have
documented climatic, atmospheric, geochemical, and organismal changes over decades to
billions of years.
Cross-Ecosystem Comparison
Comparative studies have served two important roles in ecosystem science. Most simply,
scientists often have measured some variable associated with ecosystem structure or function
across a series of ecosystems to identify typical values of that variable, show how it varies
among types of ecosystems, and generate hypotheses about what factors might control that
variable. An example of such an analysis is shown in Table 2.1 in Chapter 2. Alternatively,
scientists often test whether some factor controls an ecosystem by comparing ecosystems that
differ in that factor and not (to the extent possible) in any other relevant characteristic. For
instance, if we wanted to test whether phosphorus inputs control primary production in
lakes, we might try to measure primary production in a series of 10 lakes of similar size,
depth, species composition, and terrain that differ in their phosphorus inputs. In practice,
it often is difficult to find such a perfect series of well-matched ecosystems that could serve
as study sites.
Experiments
Experiments, whether conducted in the laboratory or in the field, are powerful ways to re-
veal controls on ecosystem structure and function (Likens 1985; Weathers et al. 2016). There are
no rules about the size of experimental units: manipulations have been made across hundreds
of square kilometers (e.g., iron fertilization experiments conducted in the ocean) and within
square centimeters. Often, the goal of experiments is to measure an ecosystem’s response to
a change in a single variable while holding all others as constant as possible. For example,
to understand whether phytoplankton in lakes were controlled by phosphorus or by other nu-
trients such as nitrogen and carbon, scientists in the Experimental Lakes Area of Canada
added phosphorus, nitrogen, and carbon to one-half of a lake (cut in two by a massive
curtain) and just nitrogen and carbon to the other half. They then compared responses—such
as the amount of primary production—in each half of the lake to see the effects of the
treatments. This whole-lake experiment helped to demonstrate that phosphorus was a major
factor controlling algal productivity in the lakes being tested (but see Chapter 9).
I. Introduction
How Do Ecosystem Scientists Learn about Ecosystems? 15
Land
management,
natural
disturbance Atmospheric Deposition
Cations and Anions
Atmospheric CO2
SUBLIMATION
TRANSLOCATION Snowpack
CO2
N2O LITTERFALL
NOx ET INFILTRATION
RUNOFF
PHREEQC soil reactions
UPTAKE SOM DECOMP.
MINERALIZATION Cation Exchange
Soil Organic Aqueous reactions
C,N,P,S NO3– NITRIF./DENITRIF.
Mineral denudation
NH4+ CO2 CO2 dissolution
DOC, Cations, Anions, CEC
Aquifer
PHREEQC stream reactions
CO2
FIGURE 1.3 DayCent-Chem model processes. DayCent-Chem was developed to address ecosystem responses to
combined atmospheric nitrogen and sulfur deposition. DayCent-Chem operates on a daily time step and computes
atmospheric deposition, soil water fluxes, snowpack and stream dynamics, plant production and uptake, soil organic
matter decomposition, mineralization, nitrification, and denitrification (left side of figure) while utilizing PHREEQC’s
(an aqueous geochemical equilibrium model) low-temperature aqueous geochemical equilibrium calculations, in-
cluding CO2 dissolution, mineral denudation, and cation exchange, to compute soil water and stream chemistry (right
side of figure), ANC, acid neutralizing capacity; BC, base cations (Ca, Mg, K, Na); CEC, cation exchange capacity; DOC,
dissolved organic carbon; and evapotranspiration; ET, evapotranspiration. The model requires considerable site-
specific environmental data to run. (From Hartman et al. 2009, Figure 1.3.)
simulation model (Figure 1.3). Models are highly flexible, can cover scales of time and space
that are difficult to study using other approaches, and often can provide quick answers at low
cost. They also are very useful as a way to organize facts and ideas, to generate, sharpen, or
narrow hypotheses, and to guide research activities. Scientists often make rapid progress by
tightly coupling theory and models to other approaches.
I. Introduction
16 1. Introduction to Ecosystem Science
of a species, or rates of carbon cycling. They are what is often found on the x or y axes of
graphs, or are used as treatments or are measured as responses in experiments. Ecosystem
structure is sometimes measured by variables such as leaf area index or the number of trophic
levels in a lake (see Chapter 11). Productivity (Chapters 2 and 3), rates of decomposition
(Chapter 4) or mineralization (Chapter 7) over time or space, or the accumulation of some
element of interest can be indicators of ecosystem function. Ecosystem development is often
described by changes in structure, function, and their relationship over time (e.g., linked
changes in soil and vegetation over millennia).
Many, if not most, of our measurements of ecosystem function are indirect. Sometimes we
can measure function directly, such as gas exchange, but these measurements are almost in-
evitably made on a tiny fraction of the ecosystem (e.g., individual leaves within a grassland or
bottle of water from a lake). To estimate a flux over a larger area of a grassland, for example,
an ecosystem scientist might deploy eddy covariance instruments that measure carbon diox-
ide, water, temperature, and wind speed and direction continuously at a place within the
grassland (see Chapter 2). From these measures, a model can be used to infer carbon dioxide
flux into or out of the ecosystem.
Scientists often choose indirect measures because they are easier to obtain across larger
parts of a system or across more systems. As another example, the measurement of
chlorophyll-a is often used as an indicator of primary productivity in aquatic ecosystems.
However, chlorophyll-a is not a direct measure of productivity, but a measure of the presence
of a pigment used in photosynthesis, and the photosynthetic process builds biomass. Like-
wise, the carbon:nitrogen (C:N) ratio in soil is often used as an indicator of litter or soil quality,
and is used to predict decomposition rates, or rates of nitrogen cycling (see Chapters 4 and 8).
To make these indirect measures useful, empirical relationships between direct and
surrogate measures must be established—quantifying these relationships is an active area
of research.
I. Introduction
How Do Ecosystem Scientists Learn about Ecosystems? 17
possible to construct a mass balance and infer processes. For example, a watershed analysis
that does not balance (quantitatively) suggests that either the element of interest is being
retained in (inputs > outputs) or leaking from (outputs > inputs) the ecosystem (see
Chapters 6 and 10). The watershed mass balance approach was pioneered in the 1960s by
BOX 1.3
Mass Balance
To see just how useful the tool of mass bal- that was bounded by the lake shore, the over-
ance can be, suppose we are trying to evalu- lying air, and the bedrock deep beneath the
ate whether a lake ecosystem is taking up or lake sediments. Using mass balance, we note
releasing phosphorus. We could try to mea- that the amount of phosphorus being retained
sure all the exchanges between parts of the by the lake ecosystem is simply the amount of
ecosystem (e.g., the uptake of phosphorus phosphorus going into the lake minus the
by phytoplankton and rooted plants; the con- amount that is leaving the lake. Now we just
sumption and excretion of phosphorus by the have to measure the exchanges across the eco-
animals that eat phytoplankton and plants; system boundary (stream water and ground-
the release of phosphorus during the decay water going into and out of the lake; rain,
of phytoplankton, plants, and animals; and snow, and particles falling on the lake; and
dozens of other exchanges), then simply any animals entering and leaving the lake—
sum up all of these measurements. It would hard enough!) to calculate whether the lake is
take an enormous amount of work to mea- taking up or releasing phosphorus. In the case
sure all the exchanges, and our final answer of Mirror Lake, New Hampshire (Figure 1.4),
would be fraught with large uncertainties. Al- almost 40% of incoming phosphorus is
ternatively, we could define a lake ecosystem retained by the lake.
le
ib 1
y
ar 1
ar
u .
in
ib 0
Tr (Data from Winter and Likens 2009.)
le
t
Tributary inlet
1.4
Outflow
Ground water in 1.7
0.3
I. Introduction
18 1. Introduction to Ecosystem Science
scientists at the Hubbard Brook Experimental Forest, New Hampshire (Bormann and Likens
1967; Likens 2013), and has been used powerfully around the world to understand the abiotic
and biotic movement of a suite of elements through ecosystems.
The other powerful “balance” tool that ecosystem scientists use is charge balance. In water,
the charges held by positive ions (cations) and negative ions (anions) must balance each other.
That is, for every anion (such as chloride) in an aqueous solution, there must be a
corresponding cation (such as sodium). Why is this tool so useful? Charge balance tells us,
for example, that when an anion moves through a forest soil from groundwater into a stream,
it must be accompanied by a corresponding cation (see Chapter 6). The sum of all the negative
charges brought by anions must be balanced by the same number of positive charges from
cations. Charge balance also makes it possible to check whether the major ions in a water sam-
ple have been measured correctly; a charge imbalance tells us that a measurement error has
been made or that we have not quantified all the cations or anions that are important in a
sample.
Tracers
As useful as balances are as tools, they tell us about the bulk (or net) movements of mate-
rials through ecosystems, and rarely allow us to distinguish among different pathways of ma-
terial movement within ecosystems. All nitrogen atoms look alike to a mass balance. Tracers
are tools that allow ecosystem scientists to distinguish among particular pathways of material
movement by labeling just some of the atoms or molecules of interest. Ecosystem scientists
have used several tracer methods, which have been enormously powerful in understanding
how ecosystems work.
Radioisotopes (Box 1.4) were some of the first tracers used in ecosystem science. Radioiso-
topes can be detected and quantified at very low concentrations, so they make excellent
tracers, and have had many applications in ecosystem science. In the mid-20th century, eco-
system scientists added small amounts of radioisotopes to ecosystems to trace the movement
of water and the uptake and movement of carbon and limiting nutrients, such as phosphorus,
through ecosystems. Radioisotopes are no longer added to natural ecosystems as tracers be-
cause of associated health risks, but they continue to be used widely in laboratory studies and
measurements (e.g., to measure microbial production; see Chapter 3). They also are used in
“natural abundance” studies where ecosystem scientists use the very low natural abundance
of radioisotopes to trace the movement of materials through ecosystems, rather than adding
radioisotopes to ecosystems. For example, Caraco and her colleagues (2010) observed that the
concentration of 14C in organic matter that washed into the Hudson River from the soils of its
watershed was very different than that of organic matter produced by photosynthesis within
the river. They could therefore use 14C to trace movement of terrestrial organic matter through
the Hudson River food web, and show that modern zooplankton were being supported in
part by carbon that was captured by primary production thousands of years ago (Figure 1.5).
Stable isotopes have largely taken the place of radioisotopes as tracers outside the labora-
tory (Box 1.4). Although much more difficult to measure and often expensive to use, stable
isotopes do not present a health risk to humans and wildlife. Stable isotopes are available
for many elements of ecological interest, including hydrogen, nitrogen, carbon, oxygen,
and sulfur. Stable isotopes often are added to ecosystems (or to laboratory experiments)
and followed as they move through the system. For example, Templer and her colleagues
I. Introduction
How Do Ecosystem Scientists Learn about Ecosystems? 19
BOX 1.4
(2005) added a stable isotope of nitrogen, 15N, to forest plots in the Catskill Mountains, NY,
and then followed it into soil, microbial biomass, understory plants, tree roots, wood, and
leaves, and found that most of the nitrogen stayed in the soil. Alternatively, ecosystem scien-
tists often use natural abundance studies of stable isotopes to follow the movement of mate-
rials through ecosystems.
Substances other than isotopes can be used as tracers as well. For instance, certain fatty
acids cannot be synthesized by animals and are made only by particular kinds of algae. By
analyzing the fatty acid content of zooplankton and fish, the contribution of different kinds
of algae throughout the food web can be traced. Caffeine, which is not readily degraded in
conventional sewage treatment plants, is sometimes used as a tracer for sewage. The kinds
of substances that can be used as tracers are highly varied, limited only by the ingenuity
and analytical capabilities of the investigator.
Spatial Data
Where are the regions of high and low productivity around the globe? How do they change
over the seasons? These are questions that can now be answered largely as a result of the
availability of remote sensing tools and spatially explicit data. The ability to collect, represent,
and analyze spatially explicit data has risen exponentially over the past decade (Weathers
I. Introduction
20 1. Introduction to Ecosystem Science
100
Modern
Modern FIAV FIAV
terrestrial
terrestrial
0
SAV
SAV
Phytoplankton Phytoplankton
–100
δ14C (‰)
–200
Zooplankton Zooplankton
–300
Aged Aged
terrestrial terrestrial
–400
–35 –30 –25 –20 –250 –200 –150 –100
δ13C (‰) δ2H (‰)
FIGURE 1.5 Use of stable and radioisotopes to determine the source of organic matter supporting zooplankton in
the Hudson River. Isotope bi-plots show 14C vs. 13C (left side) and 14C vs. 2H (right side). Sources of carbon from modern
primary production are shown near the tops of the graphs (FlAV, floating-leaved aquatic vegetation; SAV, submersed
aquatic vegetation). If zooplankton were composed of carbon and hydrogen from these sources, then the data for
isotopic composition of zooplankton would fall in the same region of the graph as the sources. Instead, zooplankton
fall far outside this region of the graph, showing that they must be composed of organic matter from both modern and
“aged” sources (i.e., organic matter thousands of years old from the soils of the Hudson River’s watershed). (From
Caraco et al. 2010.)
et al. 2016). Remote sensing and the georeferencing of basic data on landscape characteristics
such as elevation, water bodies, land cover, and geological materials have opened the door to
a description of ecosystem structure over large areas (see Holmes and Likens 2016; Boucher
et al., 2018). Geographic information systems (GISs) allow analysis of the relationships be-
tween these structures and fluxes in or out of these systems. For example, the variation in
atmospheric deposition across the mountainous terrain of Acadia National Park or Great
Smoky Mountains National Park can be described by a GIS model that links empirical mea-
surements to landscape features that are described in the GIS (Figure 1.6). Such spatially ex-
plicit models greatly enhance our ability to identify places on the landscape and times that
may be subject to particularly high levels of atmospheric deposition (Weathers et al. 2006).
GIS and other technologies are being used creatively and hold tremendous potential for un-
derstanding ecosystem processes across heterogeneous landscapes.
I. Introduction
Another random document with
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faltan allegados que, aunque en segunda fila, toman parte, siquiera
con la atención, en los debates de la primera. Habrá seguramente
entre los allegados un señor muy fino y muy risueño, con bastón y
gafas. No se moverá de la silla, no pedirá un fósforo, no hará una
pregunta, sin despepitarse en excusas y cumplidos. «Usted
dispense», «¿me hace usted el obsequio?» «con permiso de
usted», etc., etc... y no habrá dicho en todo el año cosa más
substanciosa. Pero, en una ocasión, trajo usted á la porfía (y note
que no digo conversación), un apellido que hasta entonces no había
sonado allí. Óyelo el de las gafas, y, clavándolas en usted, le
pregunta, con una voz muy dulce y una cara muy risueña:
—¿Verduguillos ha dicho usted, caballero?
—Verduguillos, sí señor,—responde usted parándose en firme.
—¿Sabe usted—insiste el otro,—(y usted perdone si le interrumpo
un momento), si ese señor de Verduguillos tiene parientes en
Cuzcurrita de Río Tirón?
—¿Por qué he de saber yo eso, si jamás allá estuve, ni conozco á
ese señor más que de vista?—replica usted con el sosiego y la
amabilidad que eran de esperarse.
—Perdone usted, caballero—dice el intruso hecho unas mieles,—y
verá por qué me he tomado la libertad de interrumpirle.
Y en esto, deja la silla, sale al centro, encárase con el grupo
principal, afirma las gafas en el entrecejo, carraspea, sonríese y
dice:
—Pues, señor, verán ustedes por qué me ha interesado tanto el oir
á este caballero nombrar á ese señor de Verduguillos. Por el mes de
septiembre del año treinta y ocho, salí yo de Zamora (donde nací y
me crié y radican los pocos ó muchos bienes que heredé á la
muerte de mis padres, y los que he podido adquirir después acá con
el fruto de mis especulaciones modestas), con el propósito de hacer
un largo viaje, por exigirlo así los asuntos de la familia, y también, si
he de ser franco, el estado de mi salud...
Así comienza este señor la relación de un viaje por media España,
con largas detenciones en todos los puertos y plazas del tránsito, y
minuciosas observaciones estadísticas y climatéricas, sin pizca de
interés, ni método, ni estilo, ni substancia, hasta venir á parar, al
cabo de tres mortales cuartos de hora, á Logroño, en la cual ciudad
conocía al comerciante don Fulano de Tal; y decirnos que, yendo á
visitarle á su escritorio, hallóse allí con un caballero, muy amigo
también del don Fulano, el cual don Fulano le dijo á él al despedirse
el otro:
—Este señor que acaba de salir, es don Pacomio Verduguillos,
natural y vecino de Cuzcurrita de Río Tirón.
Al llegar aquí con el cuento el de las gafas, espera usted el toque de
efecto, el desenlace sorprendente, la gracia del suceso; porque es
de saberse que el narrador se ha quedado en silencio y mirando de
hito en hito á los resignados oyentes. Pero el silencio sigue y la
sorpresa no asoma. Alguien se aventura, y pregunta al del bastón:
—Pero ¿por qué le chocó á usted tanto el oir nombrar á este
Verduguillos?
—Hombre—responde el interpelado, con candidez angelical,—
porque podía muy bien ser pariente del otro Verduguillos que yo
conocí en Logroño.
¡Y para eso interrumpió un animado y sabrosísimo debate; y estuvo,
durante cerca de una hora, ensartando insulsez tras de insulsez,
simpleza tras de simpleza, adormeciendo á unos, quemando la
sangre á otros y aburriéndolos á todos! Y usted llevó la cruz con
paciencia, y yo también; y lo mismo al día siguiente, porque el
bueno del zamorano, desde que pierde la cortedad con el primer
relato, ya no cierra boca en la tertulia, y siempre tan ameno,
divertido y oportuno. Pero nos permitimos los dos un desahoguillo
en un aparte.
—Amigo—dije, ó me dijo usted,—¡este hombre es insufrible:
estando él no se puede venir aquí! Y se oyó el rumor del desahogo,
y ¡qué caras nos pusieron los señores tolerantes, que estaban tan
aburridos como nosotros!
Al día siguiente asoma usted la cabeza á la puerta, ve al de las
gafas en el uso de la palabra, retrocede y no vuelve; ni yo tampoco.
Y porque no volvemos, y además decimos lo que mejor nos parece
del motivo, ¡qué ponernos de intolerantes y hasta de inciviles!...
¡Caramba, protesto contra la enormidad de esta injusticia! En este
caso no hay más intolerantes que el señor de Zamora, que
interrumpe toda conversación racional y obliga á hombres de buen
sentido á que oigan las interminables boberías que él enjareta sin
punto de reposo, y los forzados tolerantes que le escuchan con
paciencia, y no la tienen para oir que otros carecen de ella.
Trátase ahora de un embustero, que un día y otro día le abruma á
usted con narraciones autobiográficas, sin principio ni fin, como la
eternidad de Dios; pero muy punteadas, muy comeadas y con más
espacios que un libro de malos versos. Oye usted una historia, y
dos, y tres, ya con mala cara; pero, al fin, se acaba la paciencia, y
un día interrumpe usted al sujeto de los á propósitos, y le dice:
—Mire usted, hombre: en primer lugar, la mayor parte de lo que
usted me cuenta se lo he contado yo á usted en cuatro palabras; en
segundo lugar, le sucedió á un condiscípulo mío en Oviedo, y no á
un amigo de usted en Zaragoza; en tercer lugar, no pasó como
usted lo refiere, sino del modo contrario: mi condiscípulo no adquirió
una capa aquella noche, sino que perdió la que llevaba, y, además,
el juicio, con costas, á los pocos días...
—Pues lo mismo da...
—Justo: media vuelta á la derecha es lo mismo que media vuelta á
la izquierda, sólo que es todo lo contrario.
—¡Caramba, es usted lo más intolerante!... No se puede hablar con
usted...
¡Todavía le parece poco, al ángel de Dios, la tolerancia que se ha
tenido con él!
Media docena de mujeres, ó menos, si á usted le parecen muchas
seis, se pasan una tarde entera desollando con la lengua al lucero
del alba. ¡Eso sí, con las mejores formas y la intención más santa!
De una dirán que es un dolor que, siendo tan bonita, sea tan charra
en el vestir, tan tosca en el hablar, tan inconsecuente en sus
amistades, tan desleal en sus amores; de otra, que es mordaz y
maldiciente, en lo cual se perjudica mucho, porque teniendo esta
falta, y la otra, y la de más allá, da pie para que cualquiera que se
estime en tan poco como ella, se las saque á relucir; de otra, que es
una desgraciada, porque el marido la ha puesto á ración, así en el
vestir como en el bailar, á causa de que fué algo despilfarrada
siempre en estos dos ramos de buena sociedad; de otra, que ya no
halla modista que la haga un traje si no paga adelantadas las
hechuras, y que no le venden nada en las tiendas, sino con el dinero
en la mano, etc., etc., etc... En esto, entra usted (es un suponer) y,
continuando el desuello, llegan á preguntarle si conoce á cierta
señora de éstas ó las otras señas; y como la tal es mujer de historia,
y usted la sabe de corrido, repítela allí con comentarios, creyendo
hacer á su auditorio un señalado servicio. Yo creo también que
usted se le hace, pues no fué á humo de pajas la preguntita; pero es
lo cierto que todas aquellas señoras, después de oirle á usted,
exclaman, con el más sincero de los asombros:
—¡Jesús!... Con razón dicen que es usted temible.
—¡Yo temible, señoras mías?—responde usted.—¿Y por qué?
—¡Porque es usted lo más intolerante y lo más!...
¡Vaya usted á convencer á aquellas damas de que viven
constantemente encenagadas en el pecado que á usted le cuelgan!
No hay inconveniente en que, abandonando estos tiquis-miquis que
ocurren en el ordinario trato social, dirijamos el anteojo unos grados
más arriba.
Todos los días halla usted en periódicos, en folletos y en libros,
sátiras, burlas y disertaciones en serio contra ideas, sentimientos y
hasta personas muy de la devoción de usted. Ocúrresele mirar al
campo de donde parten tantos proyectiles, y le ve usted sembrado
de ridiculeces, farsas y toda clase de miserias; saca usted al palo
media docena de ellas, por vía de muestra, en un papel, en un
folleto ó en un libro; y ¡Virgen María, cómo le ponen á usted de
intolerante y de mordaz, los mismos que tienen la mordacidad y la
intolerancia por oficio!
Así andan, amigo, las cosas de justicia en el ordinario comercio de
las gentes; así se ataja al más inofensivo en el trayecto social en
que pasea su nombre, y así se pretende conducirle al extremo á que
no llegan en el mundo más que las bestias... y los que tienen la
manía de la tolerancia (siendo lógicos en ella): á ver, oir y callar... es
decir, á matar la sed con petróleo, allí donde haya un extravagante
que tal haga delante de usted.
Usted es hombre de sencillas y ordenadas costumbres (es también
un suponer): ni el mundo le tira, ni sus pompas y algaradas le
seducen. Éstos son gustos lícitos y racionales. Ajustándose á ellos,
en paz y en gracia de Dios, se da usted con un baile en los ojos:
tuerce usted el camino; tropieza usted más allá con una mascarada
de calaveras del gran mundo: echa usted por otro lado; allí topa
usted con la misma gente haciendo cuadros plásticos y animados
acertijos: cambia usted de rumbo; aquí asaltos, en el otro lado
conciertos... pues á la otra acera. Ni usted apedrea á los que bailan,
ni apostrofa á los que jiran, ni se ríe de los que se descoyuntan para
remedar á Cristo en la agonía, ni silba á los que reciben una
sorpresa, anunciada quince días antes, ni influye con el Gobernador
para que meta en la cárcel á toda esa gente: limítase á huir de lo
que le aburre, y á hacer lo que más le divierte ó menos le incomoda.
No haría otra cosa un santo.
Pero es el caso que los señores tolerantes no se conforman con
esto, y quieren que les diga usted por qué no concurre á los bailes, y
á las jiras, y á los cuadros vivos, y á los asaltos... y aquí está el
intríngulis precisamente; y si estos rasguños que trazo no fueran,
como he dicho, un inocente desahogo entre nosotros dos, y en
reserva, me atrevería á llamar la atención del lector hacia el
aparente fenómeno, cuya explicación es sencillísima, por lo cual, no
es fenómeno, aunque por tal le toman algunos.
Cuando á usted se le pregunta por qué no piensa como su vecino
sobre determinados puntos de transcendencia, á buen seguro que
se le ocurra á nadie que oiga la respuesta, agarrarse á ella para
llamarle á usted intolerante; pero que se le pregunte por qué no
baila, por qué no jira, etc., etc... y no bien ha contestado usted, ya
tiene encima el Inri de la intolerancia. Y ¿por qué en este caso y en
el otro no? Porque no está el intríngulis en la persona, ni en sus
razones, ni en el modo de exponerlas, sino en la cosa de que se
trata, que, muy á menudo, es, de por sí, ridícula, ó impertinente, ó
pueril cuando menos, y no resiste, sin deshacerse entre las manos,
el análisis de un hombre de seso; al cual hombre, no pudiendo
replicársele en buena justicia, en venganza se le pone un mote.
Por eso llevan el de intolerantes tantos caracteres dóciles, y creen
poner una pica en Flandes, y hasta se llaman guapos chicos y
excelentes sujetos en la sociedad, los que en ella entran con todas,
como la romana del diablo, menos con el sentido común. Quod erat
demonstrandum.
Á pesar de ello, y aun de la mucha saliva, que al propio asunto
hemos consagrado en nuestras conversaciones verbales, júzgole
apenas desflorado. ¡Cuánto me queda todavía que oir de los
inofensivos labios de usted!
Entre tanto, y dicho lo dicho, despidámonos por hoy, con la íntima
satisfacción, bien añeja en nosotros, de haber pasado juntos, en
espíritu, un agradable rato, sin murmurar de nadie ni ofender al
prójimo con hechos, con dichos ni con deseos.
1880.
EL CERVANTISMO