A teleofunctional account of evolutionary mismatch

Abstract

When the environment in which an organism lives deviates in some essential way from that to which it is adapted, this is described as “evolutionary mismatch,” or “evolutionary novelty.” The notion of mismatch plays an important role, explicitly or implicitly, in evolution-informed cognitive psychology, clinical psychology, and medicine. The evolutionary novelty of our contemporary environment is thought to have significant implications for our health and well-being. However, scientists have generally been working without a clear definition of mismatch. This paper defines mismatch as deviations in the environment that render biological traits unable, or impaired in their ability, to produce their selected effects (i.e., to perform their proper functions in Neander’s sense). The machinery developed by Millikan in connection with her account of proper function, and with her related teleosemantic account of representation, is used to identify four major types, and several subtypes, of evolutionary mismatch. While the taxonomy offered here does not in itself resolve any scientific debates, the hope is that it can be used to better formulate empirical hypotheses concerning the effects of mismatch. To illustrate, it is used to show that the controversial hypothesis that general intelligence evolved as an adaptation to handle evolutionary novelty can, contra some critics, be formulated in a conceptually coherent way.

Full text

A teleofunctional account of evolutionary mismatch
Nathan Cofnas
1
Received: 20 August 2015 / Accepted: 1 May 2016 / Published online: 6 May 2016
The Author(s) 2016. This article is published with open access at Springerlink.com
Abstract When the environment in which an organism lives deviates in some
essential way from that to which it is adapted, this is described as ‘‘evolutionary
mismatch,’’ or ‘‘evolutionary novelty.’’ The notion of mismatch plays an important
role, explicitly or implicitly, in evolution-informed cognitive psychology, clinical
psychology, and medicine. The evolutionary novelty of our contemporary envi-
ronment is thought to have significant implications for our health and well-being.
However, scientists have generally been working without a clear definition of
mismatch. This paper defines mismatch as deviations in the environment that render
biological traits unable, or impaired in their ability, to produce their selected effects
(i.e., to perform their proper functions in Neander’s sense). The machinery devel-
oped by Millikan in connection with her account of proper function, and with her
related teleosemantic account of representation, is used to identify four major types,
and several subtypes, of evolutionary mismatch. While the taxonomy offered here
does not in itself resolve any scientific debates, the hope is that it can be used to
better formulate empirical hypotheses concerning the effects of mismatch. To
illustrate, it is used to show that the controversial hypothesis that general intelli-
gence evolved as an adaptation to handle evolutionary novelty can, contra some
critics, be formulated in a conceptually coherent way.
Keywords Evolutionary mismatch Evolutionary novelty Environment of
evolutionary adaptedness Evolution of intelligence Proper function 
Teleosemantics
&Nathan Cofnas
nbc25@cam.ac.uk
1
Department of History and Philosophy of Science, University of Cambridge, Free School Lane,
Cambridge CB2 3RH, UK
123
Biol Philos (2016) 31:507–525
DOI 10.1007/s10539-016-9527-1
Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Introduction
When the environment in which an organism lives deviates in some essential-but-
not-yet-defined way from that to which it is adapted, this is described as
‘‘evolutionary mismatch,’’ i.e., between the ancestral and the current environment
(Confer et al. 2010; Lloyd et al. 2014). Elements of the current environment that
deviate from the ancestral one are described as ‘‘evolutionarily novel’’ (Irons 1998;
Sterelny 2010).
1
Those that do not, as ‘‘evolutionarily familiar’’ (Kanazawa 2004a).
The environment(s) to which an organism is adapted can be termed its ancestral
environment (AE).
The notion of mismatch plays an important role, explicitly or implicitly, in
evolution-informed cognitive psychology (e.g., Confer et al. 2010; Tooby and
Cosmides 1990), clinical psychology (e.g., Gilbert and Bailey 2000; Nesse 2004),
and medicine (e.g., Eaton et al. 1988; Williams and Nesse 1991). However,
scientists have generally been working without a clear definition of mismatch. This
paper defines mismatch as deviations in the environment that render biological
traits unable, or impaired in their ability, to produce their selected effects (i.e., to
perform their proper functions in Neander’s 1991a,bsense). The machinery
developed by Millikan in connection with her account of proper function, and with
her related teleosemantic account of representation, is used to identify four major
types, and several subtypes, of evolutionary mismatch.
Defining mismatch (or evolutionary novelty) in this way leaves it an open
question whether different types of evolutionary novelty are good or bad for either
the fitness or the general welfare of organisms. Many evolutionarily novel aspects of
our contemporary environment are thought to have negative implications for our
health and well-being (see Barrett 2007,2010; Buss 2000; Lloyd et al. 2014:4;
Nesse 2004; Williams and Nesse 1991: 13–16). However, evolutionarily novel
technology and systems of social organization have allowed our species to greatly
increase its numbers (adaptedness) and have removed many evolutionarily familiar
sources of suffering. Not all deviations from ancestral conditions are bad, and the
neutral definition of mismatch defended in this paper captures that fact.
A previous definition of mismatch
In one of the few previous philosophical analyses of the concept, Lloyd et al. (2014:
4) define mismatch as ‘‘a [detrimental] consequence that results from a trait that
evolved in one environment being placed in another environment.’’ ‘‘‘Detrimen-
tal,’’’ they say, ‘‘will usually be defined in terms of evolutionary fitness, but it can
also refer to welfare in more general terms’’ (6).
This rules out by definition the possibility of mismatch being beneficial. As
Lloyd et al. say: ‘‘Traits that evolved in one environment need not be detrimental in
a second environment; they can be neutral or fortuitously beneficial, but these cases
are excluded by the term ‘mismatch’, which restricts our attention to the detrimental
1
In biology, the term ‘‘evolutionary novelty’’ usually refers to novel phenotypes. Scholars of mismatch
use the term to refer to novel environments.
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cases’’ (5). We need to define mismatch in more neutral terms if we wish to
investigate how deviations from the AE can be bad or good. Take an example of
mismatch considered by Lloyd et al.: A person who is able to tan and ‘‘who spends a
lot of time indoors or covered with clothing will experience sunburn when their skin
is suddenly exposed to the sun, a pattern of variation that seldom, if ever, occurred
in their’’ AE (14). Clearly, sunburns per se are bad from both a fitness (they increase
the risk of cancer) and a general-welfare perspective. But the evolutionarily novel
situation of spending a lot of time indoors and/or wearing clothing, though it may
increase the risk of sunburn in some individuals, may on the whole be a good
thing—both fitness- and welfare-wise. Deviations from AE conditions can have
negative, positive, or mixed effects on fitness and welfare. Furthermore, in actual
scientific debates in which the consequences of evolutionary novelty are at issue, it
is not adequate to define evolutionary novelty as having only negative conse-
quences. One of these debates will be discussed in the following section.
The scientific use of an account of evolutionary mismatch/novelty:
the evolution of general intelligence
Some intelligence researchers hold that general intelligence (g) is an adaptation for
dealing with evolutionary novelty (Chiappe and MacDonald 2005; Kanazawa
2004a). Byrne (1995: 38) argues that intelligence may evolve as an adaptation ‘‘in
generalist animals…to exploit continually changing environments, since they must
daily cope with novelty in order to survive.’’ Sterelny (2012: 3) sees the challenge
posed by ‘‘evolutionarily novel features of the environment’’ as having been an
important driver of hominin cognitive evolution. Critics raise two objections based
on ambiguity in the concept of evolutionary novelty.
First, they say that the proposition that something is an adaptation for
‘‘evolutionary novelty’’ is logically contradictory: Adaptations can only evolve in
response to recurrent features of the environment. If evolutionary novelty is
understood as nonrecurrent features of the environment, it is impossible to evolve an
adaptation to deal with it (Penke et al. 2011).
Second, they say that ‘‘the division between ‘evolutionarily familiar’ and
‘evolutionarily novel’ is so vague that one could legitimately argue that the same
phenomenon was both ‘evolutionarily familiar’ and ‘evolutionarily novel’’’ (Dutton
2013: 612). Kanazawa (2010)—who advocates the hypothesis—claims that
vegetarianism and liberalism are evolutionarily novel. From one perspective these
things surely are evolutionarily novel: Until recently all our ancestors ate meat and
did not go out of their way to help distant, unrelated people (what Kanazawa sees as
the goal of liberalism). But with a little ingenuity we could think up reasons for
regarding vegetarianism and liberalism as evolutionarily familiar. Dietary taboos
are a human universal, and vegetarians can be regarded as members of a moral tribe;
liberalism can be interpreted as an extension of the cooperative ideology that
prevailed among hunter–gatherers. The general-intelligence-as-an-adaptation-to-
evolutionary-novelty hypothesis will not yield any testable predictions if it is
unclear what features of the environment are novel or familiar.
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Notice that Lloyd et al.’s (2014) definition of mismatch as harmful deviations
from the AE cannot be used to resolve this debate. The contention is that general
intelligence evolved to handle novelty, which includes taking advantage of new
opportunities. Organized social codes might be evolutionarily novel, and we might
have evolved greater intelligence in order to deal with them, but the development of
social codes presumably was positive for our species.
The definition of evolutionary novelty defended here could potentially be used to
answer both of the above criticisms. First, it gives sense to the idea of recurrent
evolutionary novelty (Byrne 1995; Kanazawa 2004a; Sterelny 2010). Sterelny
(2010: 59) attributes our ability to ‘‘negotiate evolutionarily novel problems’’ in part
to the fact that ‘‘novelty is not new’’ for our species. ‘‘Our lineage has repeatedly
experienced demanding environmental change.’’ Byrne (1995: 38)—as quoted
above—refers to ‘‘continually changing environments’’ that repeatedly confront
animals with ‘‘novelty.’’ There is a conceptually coherent sense in which the
environment can be recurrently novel over evolutionary time. The environment can
remain evolutionarily novel indefinitely, as long as it continually changes so as to
disrupt the performance of a species’ proper functions. (This idea seems implicit in
the aforecited works.) In regard to the second criticism, the definition of
evolutionary novelty given here can, of course, resolve the tricky cases, such as
those mentioned above or discussed in Dutton (2013). Analysis will likely reveal
that in some cases a feature of the environment may be partly evolutionarily familiar
and partly novel.
The debate about the evolution of intelligence illustrates that ‘‘mismatch’’ may be
more than a verbal category that is useful for scientists. It may potentially be
something out there in the world that has actual consequences for evolution.
Whether this is the case is an empirical, not a philosophical, question. But empirical
hypotheses concerning the effects of mismatch require a coherent concept of
mismatch in order to be formulated. The philosophical analysis given here attempts
to clarify the concept.
Groundwork for cataloging the types of evolutionary novelty
Definition of evolutionary novelty
Neander (1991a,b) defines the ‘‘proper function(s)’’ of a biological trait as the
effect(s) of the trait that it was selected for—the effects that played a causal role in
its selection. She terms these the selected effects. Evolutionary familiarity is defined
here as the environmental conditions necessary for biological traits to perform their
(Neanderian) proper functions, i.e., to produce the effects that are responsible for
their recent (Godfrey-Smith 1994) evolution and proliferation and/or maintenance.
Evolutionary novelty is defined as deviations in the environment that prevent
biological traits from performing, or impair their ability to perform, such functions.
Such impairment is not necessarily bad for organisms from the standpoint of either
fitness or welfare. Mice may possess mechanisms the proper function of which is to
escape from cats. These mechanisms do not perform these functions in cat-free
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environments, but cat-free environments are good for mice. In humans, it is thought
that depression is an adaptation for dealing with certain types of situations, causing
the depressed person to withdraw from hopeless goals (Nesse 2000,2004).
Nevertheless, an environment that never triggered depression would be a good one
(ceteris paribus; cf. Garson 2015: 177).
Millikanian proper functions
In Millikan’s (1984) system, a direct proper function of xis Fif the reason that
xexists with character Cis because, by virtue of C,xcan perform F(26). For
something to have a direct proper function, it must be a member of what she terms a
‘‘reproductively established family.’’ Different items belong to the same reproduc-
tively established family when they resemble each other due to later items being
produced by copying, or reproduction of, earlier ones (or ‘‘models’’; 18–19). The
properties by virtue of which one item is a reproduction of another are
‘‘reproductively established properties’’ (20).
The proper function of a mechanism is relational when it is the function of the
mechanism ‘‘to do or to produce something that bears a specific relation to
something else’’ (39). For example, Smith’s dwarf chameleon (Bradypodion
taeniabronchum) camouflages itself by changing color. The ‘‘pigment-rearranging
device’’ of chameleons has the relational proper function to bring the color of the
chameleon in a certain relation—namely, ‘‘same color as’’—to the surface upon
which it is sitting (39).
When a mechanism with a relational proper function is faced with the type of
entity toward which it is its function to bring about a certain relation, the mechanism
acquires an adapted proper function: the function to produce the specific item or
situation that bears the appropriate relation to the given entity in this particular
instance (40). As noted, the chameleon’s pigment-rearranging mechanism has as a
relational proper function to make the chameleon’s color the ‘‘same color as’’ the
surface upon which the animal finds itself. If the color of the surface is specified as,
say, brown, then the pigment-rearranging mechanism has as its adapted proper
function to turn the chameleon brown. When a device performs its adapted proper
function, it is termed an adapted device. The entity to which the device is adapted is
its adaptor (40).
The proper functions of adapted devices correspond to the proper functions of
their producing devices that ‘‘lie beyond the production of these adapted devices
themselves’’ (41). The proper functions of adapted devices are termed derived
proper functions (41), and are of two types. Adapted derived proper functions are
fixed by the device-producing mechanism and its particular adaptor. Invariant
derived proper functions of an adapted device correspond to what can be described
as the ultimate purpose of the producer. A particular bee dance—the adaptor of
which is a source of nectar sighted by the dancing bee—has as its adapted derived
proper function to induce the dancer’s sisters to fly a certain distance in a specific
direction (say, 100 meters north). Its invariant derived proper function is to guide
them to nectar.
A teleofunctional account of evolutionary mismatch 511
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Proper functions and evolutionary mismatch
Again, this paper defines evolutionary mismatch/novelty as deviations in the
environment that prevent biological traits from producing, or impair their ability to
produce, their selected effects (what Neander defines as ‘‘proper function’’). The
taxonomy of evolutionary mismatch that will be given here is generated by the
following observation: The property of being ‘‘a (Neanderian) proper function’’ is
logically coextensive with that of being ‘‘either a (Millikanian) direct or invariant
derived proper function of a biological trait.’’ If a trait xhas been selected for having
character Cbecause, by virtue of C,xcan perform F,Fis the direct proper function
of x, and also its selected effect—it has been selected because it does F. If a trait
yhas been selected because it produces adapted devices that bring about certain
relationships to things in the world, the invariant derived proper function of the
adapted device is the effect for the sake of which the mechanism that produced it
was selected—invariant derived proper functions are selected effects.
To elaborate on the last point, traits with relational proper functions—which
produce adapted devices with derived proper functions—can be favored by natural
selection when they tend to bring about certain invariant results in the AE: ‘‘getting
honey,’’ ‘‘escaping lions,’’ ‘‘catching mice,’’ and the like. It is these final, invariant
results which are the ultimate evolutionary purposes of such traits—their selected
effects. Obtaining them is the invariant derived proper function of adapted devices.
If environmental conditions change such that traits with relational proper functions
tend to produce adapted devices with very different adapted derived proper
functions (e.g., flying 100 meters south rather than north), while continuing to fulfill
invariant derived proper functions (e.g., getting honey), this change is (ceteris
paribus) irrelevant from an evolutionary perspective: The trait continues to produce
the effects for which it was selected. That is why only environmental change that
prevents an adapted device from fulfilling, or impairs its ability to fulfill, its
invariant derived proper functions is ‘‘evolutionarily novel’’ according to the
definition given here.
Devices with relational proper functions must consume representations
of states of the environment
Devices with relational proper functions (which produce adapted devices with
invariant derived proper functions) ‘‘do or…produce something that bears a specific
relation to something else’’ (Millikan 1984: 39). The entity/world state toward
which the mechanism has the function of bringing about a certain relation is called
its adaptor. Any biological structure with a relational proper function performs its
adapted proper function (in the normal way) in response to correct representations
of the environment, specifically to the adaptor.
To perform relational proper functions, organisms must evolve some way to track
the relevant environmental conditions. For an organism to track some
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environmental condition E, the instantiation of Emust increase the probability of
some type of state M
P
(pfor ‘‘producer’’) occurring in the organism. When M
P
is
tokened, the mechanism M
C
(cfor ‘‘consumer’’) in the organism that uses M
P
as a
sign that Eis instantiated can exhibit a response appropriate for E. When M
P
is in
fact caused by E,M
C
performs its proper function in exhibiting the response
appropriate to Eupon the registration of M
P
. According to Millikan (1989), M
P
tokens represent the conditions that are necessary for the mechanism that consumes
M
P
—namely, some mechanism M
C
—to fulfill its proper function.
To illustrate with camouflaging in Smith’s dwarf chameleon, the chameleon has
to have some way of tracking the color it is sitting on. To do this, it must use some
sort of inner intentional icons to represent different colors. There is no consuming
mechanism that uses the skin color as a sign of anything, so the skin itself has no
representational content (Millikan 1984: 118; 1989: 283). But in order for the
pigment-arranging mechanism to perform its relational proper function (in the
normal way) on (e.g.) a brown surface, a chameleon must have some state—an inner
intentional icon—that covaries with and represents ‘‘brown surface.’’
2
Four types of evolutionary mismatch/novelty
In Millikanian terms, evolutionary mismatch is defined as deviations in the
environment that render biological traits unable, or impaired in their ability, to
perform their direct proper functions or invariant derived proper functions.
Biological traits perform their direct proper functions when the trait itself develops
(unimpaired), and environmental conditions are such that the trait produces the
effect for which it was selected. Environmental change that prevents traits with
direct proper functions from developing, or that prevents fully developed traits with
direct proper functions from producing their selected effects, constitutes two types
of mismatch labeled Un(der)development-inducing and Ineffectiveness-inducing,
respectively. Biological traits with relational proper functions perform their
invariant derived proper functions when the organism correctly represents the
environment (namely, the adaptor), and the adapted device it produces in response
to that representation brings about the right adaptive relationship vis-a
`-vis the
adaptor. Environmental change that causes the organism to misrepresent, or to fail
to represent, the environment constitutes a third type of mismatch labeled
Misrepresentation-inducing. If the environment changes so that the ways in which
the organism uses representations fail to bring about the right invariant end results,
this constitutes a fourth type of mismatch labeled Misresponse-inducing.
The four types of evolutionary mismatch—the four ways in which traits can fail
to produce their selected effects—are discussed in turn in the following four
sections.
2
Such representations are ‘‘pushmi-pullyu representations’’: They are descriptive and directive at the
same time (Millikan 1995).
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Type 1—Un(der)development-inducing: failure of a trait with a direct
proper function to develop
The environment may change so that a biological trait that has a direct proper
function simply fails to develop, or is impaired in its development. For example,
human arms and hands have the direct proper function to be used for grasping (cf.
Neander 1991a: 174). However, if a person develops in a womb that contains
thalidomide, these structures may be greatly impaired in their development.
Thalidomide in the womb is evolutionarily novel because it prevents biological
traits (i.e., arms and hands) with direct proper functions from developing, or from
developing well enough to perform their direct proper functions (i.e., grasping). A
less dramatic example among humans is learning to read as a child, which leads to
myopia among genetically susceptible individuals (Williams and Nesse 1991: 16).
Type 2—Ineffectiveness-inducing: failure of a trait to perform its direct
proper function despite unimpaired development
If a biological trait has a direct proper function, the environment may change so
that, although the trait physically develops unimpaired, it fails to perform its proper
function. Prior to the industrial revolution, peppered moths (Biston betularia)in
England were nearly all of the ‘‘typica’’ form—white with black spots. According to
the well-known story of industrial melanism, this coloration camouflaged the moths
against lichen-covered trees. In the mid-1800s, pollution from factories killed the
lichens, exposing the dark tree bark. This environmental change rendered the moths’
camouflaged structure unable to perform its direct proper function, although the
moths continued to develop unimpaired (Cook et al. 2012).
Learning
Millikan (1984: 24–25) identifies two ways of learning: An organism may repeat
behavior that was previously ‘‘rewarded’’ (or met some other set of criteria), or it
may imitate behavior that it observed in another organism. In the former case, the
direct proper function of the learned behavior is to obtain the reward. In the latter
case, it is to perform the function that the learner associated—by observation—with
the behavior in question (28). In either case, a learned behavior may be as bizarre or
as novel as you wish, but it seems that the conditions that produced it cannot be
evolutionarily novel, according to the definition given here, as long as the reward is
obtained or the (observed) function is performed. For conditions to be evolutionarily
novel, some trait has to fail to perform its direct (or invariant derived) proper
function as a result of environmental change. How can successfully performed
learned behaviors reflect evolutionary novelty?
The answer is that the biological mechanisms that underlie learning perform
their direct proper functions only when they produce (learned) behaviors with
particular characteristics—namely, those characteristics by virtue of which the
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behaviors performed certain functions in the AE, which caused the (past)
proliferation of (tokens of) the underlying learning mechanism (cf. Millikan
1989: 292). Learning mechanisms fail to perform their direct proper functions when
they do not produce learned behaviors with such characteristics. For example, in
their natural environment, cats learn to hunt when they are hungry. A housecat in an
evolutionarily novel environment might learn that, whenever it shakes its owner’s
hand, it gets a treat: It learns to shake hands when it is hungry. Being fed in response
to performing this trick is evolutionarily novel vis-a
`-vis the cat’s mechanism
responsible for learning how to obtain food. The mechanism has as its direct proper
function to establish (by learning) a connection between hunting and eating. Under
novel conditions, the mechanism causes the cat to obtain food in some other way,
but not by performing its direct proper function.
Type 3—Misrepresentation-inducing: misrepresenting or failing
to represent the environment
In order to respond contingently to biologically important objects and situations,
organisms may evolve traits with relational proper functions. In response to correct
representations of their adaptors, these traits produce adapted devices which, under
the right conditions, perform their invariant derived proper functions, thus
accomplishing the ultimate goal for the sake of which the traits were selected.
Environmental change that causes traits to fail to produce adapted devices that
perform their invariant derived proper functions as a consequence of misrepresent-
ing adaptors constitutes the third type of evolutionary mismatch. This section
reviews the basic ways in which environmental change can cause organisms to
misrepresent adaptors.
Identifying objects
Under evolutionarily familiar conditions, organisms tend to correctly represent the
objects in their environment as being of a biologically important type (such as
predator, prey, mother, sibling, potential mate), though sometimes with a bias for
false positives or false negatives (Godfrey-Smith 1991). Objects of certain types
stimulate the production of certain states in the organism, which traits with
relational proper functions use as representations of such objects.
The schema used to recognize object tokens of a particular type—i.e., to produce
tokens of a state that the organism uses to track examples of that object type—can
be acquired in either evolutionary or individual experience. Objects of different
biologically important types have been—when encountered in evolutionary
experience—more or less reliably associated with stimuli (or cues) of certain
intensities in particular configurations (Eibl-Eibesfeldt 1989). The organism may
evolve to use these cues in one of two ways. First, it may use them to pick out
examples of an object type. The organism then learns to recognize tokens of the
object type in question—forming a schema of it—through exposure to the examples.
The cues used to recognize the original example(s) are not necessarily the same cues
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used to recognize the object after the schema has been formed on the basis of the
examples. Second, the organism may, through genetic adaptation, possess innate
sensitivity to the key stimuli (or configurational key stimuli) associated with
biologically important object types. The following two subsections deal, respec-
tively, with these two ways of forming object schemas.
Learning to recognize objects: imprinting
Organisms that rely on learning to form a schema of a biologically important type of
object must be adapted to use (historically reliable) cues to guide them to correct
examples (i.e., tokens) of the object type in question. When a schema formed in this
way is incorrigible, the process is termed imprinting (Lorenz 1970: 124–133). The
time in the organism’s life cycle in which imprinting occurs, or can occur, is termed
the critical period.
If, during the critical period, any of the cues used by an organism to guide it to
biologically correct examples of the object type to imprint on are removed, or are
attached to tokens of other object types, this constitutes evolutionary novelty (cf.
Garson 2015: 180–183). The result will be that the organism fails to imprint on any
object, or imprints on an incorrect object type.
The particular object token(s) that an organism uses to form a schema of a
biologically important type may, paradoxically, not themselves be of that type.
Many birds and mammals use either their ‘‘parents’’ or their ‘‘siblings’’ to form
schemas of ‘‘potential mates,’’ but parents and siblings themselves are (usually) not
potential mates. A mallard drake (Anas platyrhynchos) raised by graylag goose
(Anser anser) foster parents along with mallard siblings will flock with mallards but
court graylags. Thus, it uses its siblings to form a schema of objects that elicit
flocking behavior, and it uses its mother to form a schema of the objects that elicit
courting behavior, although the mother herself will never be courted (Lorenz 1970:
280). A jackdaw (Corvus monedula) that has been reared by a human will, upon
reaching sexual maturity, court Homo sapiens, but not the particular one who raised
it (131). A Muscovy drake (Cairina moschata) raised by a human along with
surrogate mallard siblings will, upon reaching sexual maturity, court mallards,
indicating that it, unlike mallards and jackdaws, uses ‘‘siblings’’ as models to form
the schema of ‘‘potential mates’’ (132).
For ducks and jackdaws, being raised by humans, or by or with birds of another
species, is evolutionarily novel: Cues that, under evolutionarily familiar conditions,
guide them to imprint sexual responses on examples of correct object types are, in
the novel situation, attached to biologically incorrect objects. This causes them to
misrepresent members of other species as potential mates and to produce an adapted
device with the adapted derived proper function of courting these incorrect objects.
Of course, this adapted device fails to perform its invariant derived proper function
of bringing about copulation with (non-relative) conspecifics.
Under evolutionarily familiar conditions, humans imprint sexual disgust
responses on close relatives. Recent research suggests that the sibling-inbreeding-
avoidance mechanism in humans relies on two cues to establish siblinghood:
childhood coresidence and maternal perinatal association (Lieberman et al. 2007).
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The latter cue refers to observing someone as an infant being cared for by one’s own
biological mother—this cue is potentially available only to older siblings. The
former cue is potentially available to older, younger, and same-aged siblings.
In humans, biological siblings who are not raised together often fail to imprint
sexual disgust on each other. Contrary to thousands of years of popular belief—
reflected most notably in the story of Oedipus—explicit knowledge by itself that
someone is a relative appears to have little influence on one’s attitude toward sexual
relations with them. In fact, it is very common for relatives (whether siblings or
parents and children) separated at birth to feel, upon reunion, strongly attracted to
each other, and even to act on these feelings. This phenomenon is termed ‘‘genetic
sexual attraction’’ (Greenberg and Littlewood 1995). People who experience genetic
sexual attraction, however, are typically averse to sexual relations with their
adoptive family members. The situation of being raised without, and reunited in
adulthood with, an opposite-sex sibling is evolutionarily novel: It greatly increases
the likelihood of failure to represent opposite-sex siblings as relatives and thus to
produce an adapted device that fulfills its invariant derived proper function of
preventing sexual attraction to, and relations with, siblings.
Genetic adaptation to recognize objects
Any natural, evolutionarily familiar type of object will (in the AE) more or less
reliably emanate stimuli in particular intensities and in particular configurations.
Organisms can evolve to use a subset of these stimuli to recognize tokens of the
object type. Registration of the ‘‘key’’ stimuli or configurational stimuli produces a
token of a state type used by the organism to represent the object (Eibl-Eibesfeldt
1989: 55; Tinbergen 1951: 25, 27). An object is evolutionarily novel when it
emanates key stimuli associated with an evolutionarily familiar object but in
different intensities or combinations, and/or it emanates key stimuli associated with
an evolutionarily familiar type of object but is not of the same biologically
important type. Visual pornography, for example, consists of images/video
emanating key stimuli associated with ‘‘sexually receptive mate,’’ and these key
stimuli may be arranged in evolutionarily novel combinations. More examples will
be given in the subsection on supernormal stimuli (below).
Consider also Dretske’s (1986) example of marine bacteria that swim toward
oxygen-low water by means of magnetotaxis. These bacteria contain tiny magnets
called magnetosomes. The magnetosomes of species in the northern hemisphere
draw bacteria toward geomagnetic north, which is downwards and away from
highly oxygenated surface water. Species in the southern hemisphere have their
magnetosomes reversed: They are drawn toward geomagnetic south, which is
downwards and away from surface water in that hemisphere. The magnetic pull is
the key stimulus that the bacteria are genetically adapted to use to produce a
representation of the direction of oxygen-low water.
The system that the bacterium uses to orient itself has a relational proper
function. Its function is to bring about a certain relation between the organism and
the oxygen-low water (namely, that the bacterium should swim toward and be in
such water). ‘‘Oxygen-low water’’ is the adaptor—the thing toward which the
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system has the function of bringing about a relation. The bacterium has to represent
the adaptor in order to respond to it. ‘‘Attraction-to-a-direction-at-a-time’’ (cf.
Millikan 1989: 290) does this—it represents the direction of oxygen-low water. The
adapted device produced in response to this representation
3
is a specific orientation
with the flagellum propelling the bacterium forward. The adapted derived proper
function of the adapted device is to swim in the direction of the magnetic pull at that
particular time. The invariant derived proper function is to swim in the direction of
oxygen-low water.
Living in the southern hemisphere would be evolutionarily novel for a northern
hemisphere bacterium and vice versa. In either case the orienting system in the
bacterium would misrepresent its adaptor (low-oxygen water) as being upwards,
because the key stimulus it uses to recognize oxygen-low water is associated with
oxygen-high water. In response to the misrepresentation, the bacterium would
produce an adapted device that fails to perform its invariant derived proper function.
Identifying situations
Under evolutionarily familiar conditions, organisms tend to correctly represent the
situations in their environment as being of a biologically important type, and, just as
with objects, sometimes with a bias for false positives or false negatives. Tokens of
biologically significant situation types stimulate the production of certain states in
the organism, which devices with relational proper functions use as representations.
Environmental deviations that cause situations to remain unrepresented, or to be
incorrectly represented, constitute a subtype of evolutionary novelty.
Mismatch: misrepresenting situations
Consider social exchange (in our species). Rational choice theory often fails to
predict peoples’ behavior in both artificial and natural social-exchange situations.
About half of subjects cooperate in anonymous, one-shot prisoner’s dilemma games
(PDGs), while rational choice theory predicts that they will defect (Kanazawa
2004b: 41). Kanazawa (2004b) hypothesizes that the more a social-exchange
situation corresponds to situations that existed in the AE, the more likely people will
be to behave according to the predictions of rational choice theory. Why do so many
people cooperate when playing anonymous, one-shot PDGs when doing so amounts
to giving money to an anonymous stranger? The answer, Kanazawa proposes, is
that, in the AE, there was no such thing as anonymous social exchange, nor of
guaranteed one-time interactions. The only way to interact with someone in the AE
was face-to-face. And, unless you killed them, there was no such thing as interacting
with someone and knowing that you would never see them again. Situations of
social exchange activate specialized cognitive mechanisms, which do not take cues
of ‘‘anonymity’’ or ‘‘one-time-ness’’ as input. In modern postindustrial society,
anonymous and non-iterated social exchange is common—in some contexts, even
the norm. Our behavior (e.g., cooperating in anonymous, one-shot PDGs and
3
Pushmi-pullyu representation—see note 2 (above).
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contributing to public goods) reflects a representation of our environment (namely,
interactions are not anonymous or one shot) that is now often inaccurate. In
anonymous, one-shot social exchange, our cognitive mechanisms produce an
adapted device with the adapted derived proper function to be generous to our
partner. The device fails to fulfill its invariant derived proper function of
maximizing expected long-run payoff.
In response to cues reflecting specific environmental conditions, organisms can
develop an adaptive phenotype by means of developmental switches. For example,
some species of water flea (crustaceans of the genus Daphnia) develop hard shells if
exposed to chemicals emitted by their predators. The shells are ‘‘metabolically
costly,’’ but worth it if the water flea is in an environment with many predators
(Garson 2015: 178–179). Garson defines ‘‘developmental mismatch’’ as occurring
when a cue triggers the development of a phenotype that turns out not to be
appropriate—for example, if the chemical associated with predators was emitted in
an environment without any predators.
‘‘Developmental mismatches’’ result from misrepresenting the environment. The
developmental switch is part of a system with a relational proper function. The
adaptor is the environmental situation (e.g., many predators), which must be
correctly represented. The phenotype that develops is an adapted device with an
invariant derived proper function to relate to the adaptor in the appropriate way. If
the organism misrepresents the adaptor, it will develop a phenotype that fails to
accomplish its invariant derived proper function.
Supernormal stimuli
The releasing effect of key stimuli (used to identify either objects or situations) can
be increased by intensifying their presentation (Tinbergen 1951: 44–46). Config-
urational key stimuli can also be intensified by exaggerating the stimulus relations
in question (e.g., dolls, teddy bears, and certain cartoon characters; Eibl-Eibesfeldt
1989: 59–63). An object in which several key stimuli with additive releasing effects
are combined may have a releasing effect greater than any, or almost any, natural
object. That is, objects can be represented as being clear examples of a certain
biologically important type, although they may not be of that type at all.
For practically all organisms, habitat selection is an important adaptive problem.
Survival is closely tied to living in the right environment. Organisms evolve to be
attracted to key stimuli associated with their biologically ideal environment, and to
avoid environments that lack these stimuli. Humans seem to have evolved to prefer
environmental conditions associated with the safest part of the Pleistocene African
savannah: verdant, open spaces with scattered trees, especially in close proximity to
clear water. When shown photographs of a variety of environments, children who
have never been to a tropical savanna indicate that it would be the best environment
to live in or to visit. Adults develop a comparable level of attraction to environments
with which they are familiar, but continue to judge the savannah as appealing
regardless of their individual experience (Balling and Falk 1982).
The idyllic savannah environment is recreated in an exaggerated form, albeit on a
smaller scale, in the suburban lawn. The grass of the suburban lawn is greener,
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brighter, and more consistent than that which ever existed in the AE. Ironically,
these lawns attain their supernormally stimulating effects due to the administration
of health-threatening fertilizers and herbicides that sometimes reduce the fitness of
those who come into contact with them (Barrett 2007: 67). The popular herbicide
2,4-dichlorophenoxyacetic acid, for example, threatens fitness in a dramatic way.
Among men, exposure to this chemical is associated with reduced sperm quality;
among pregnant women, with increased risk of miscarriage. Our attraction to key
stimuli associated with the most fitness-promoting environment in the AE leads us
to create an environment that needlessly threatens us, though we are compensated
with a certain amount of aesthetic pleasure. Our representation of certain landscape
features as indicating a vicinity of maximum safety is no longer accurate. Our
habitat-selection mechanism produces an adapted device (namely, a desire to live in
close proximity to supernormally stimulating lawns) that may fail to fulfill its
invariant derived proper function (namely, leading us to an area that is more fitness
promoting than the alternatives).
Type 4—Misresponse-inducing: responding incorrectly
to environmental conditions
The previous section dealt with the principles by which organisms produce states to
track environmental conditions. This section deals with how traits with relational
proper functions use,orconsume, these states in order to bring about adaptive
relationships vis-a
`-vis their adaptors.
Under evolutionarily familiar conditions, particular responses are elicited with
certain frequencies, and are joined together to form behavioral chains to accomplish
biologically important goals (i.e., invariant derived proper functions). When
conditions change such that responses are elicited with different frequencies, or
chains of behavior are not assembled correctly, or responses no longer achieve the
biological purpose for which they evolved, the environment is evolutionarily novel.
Behavioral chains
Biological goals are often accomplished by the successive performance of relational
proper functions, each of which is released by a separate stimulus situation/adaptor
and is subject to its own appetence (cf. Millikan 1984: 41).
In many cases, the completion of one relational proper function in a behavioral
chain brings about the adaptor necessary to release the next relational proper
function, and so on, until a biological goal is accomplished (through performance of
the invariant derived proper function of the adapted device produced by the last
relational proper function in the chain). This is particularly obvious in the case of
reproductive behavior, which, in many animals, begins with courting (itself
composed of a chain of behaviors), the performance of which brings about the
stimulus situation/adaptor releasing copulation/insemination. This produces fertil-
ized eggs or young, which in turn release parental behaviors.
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Successful hunting behavior in cats (and many other predators) requires the
integration of several separate motor patterns: stalking, pouncing, killing, and eating
prey. Each of these behaviors can be released independently, and can be temporarily
exhausted by repeated elicitation without reducing the appetence for the other
behaviors that comprise the chain. Under natural conditions, successful stalking—
released by the sight of a small animal, or signs of such an animal—brings about a
situation that releases pouncing, which, when successful, leads to killing and finally
to eating. Under natural conditions, cats learn to hunt when they are hungry. Under
the evolutionarily novel conditions of apartment living, the elements of the hunting
behavioral chain often become separated: The cat stalks shadows, pounces on balls
of string, directs killing motions at random objects, and eats ‘‘cat food’’ from a dish.
Frequency of elicitation of responses
Objects and situations for which a particular response is adaptive are encountered,
or must be sought or brought about, with a particular frequency in the AE.
Organisms typically evolve so that, under evolutionarily familiar conditions,
particular responses are elicited by the right stimulus situations with the frequency
appropriate to the AE. The drive to perform a behavior (a relational proper function)
is typically matched to the appropriate frequency (Tinbergen 1951: 168). An
environment is evolutionarily novel vis-a
`-vis an evolved response if it presents the
eliciting stimulus situation (which is represented as the adaptor) with a greater or
lesser frequency than it occurred in the organism’s evolutionary experience. In such
environments, evolved responses may be elicited in biologically inappropriate
situations (the organism produces an adapted device in response to the wrong
adaptor) or fail to be elicited in appropriate situations (the organism fails to produce
an adapted device in response to the right adaptor).
As noted in the previous subsection, the elements comprising a behavioral chain
are elicited by different stimulus situations, and are subject to their own appetence.
Such elements will often, under evolutionarily familiar conditions, be elicited with
different frequencies. This occurs when performance of one element of the chain
often does not produce the adaptor for the relational proper function that comes next
in the chain. As discussed, hunting in cats involves the successive performance of
four separate behaviors, viz., stalking, pouncing, killing, and eating prey. Under
evolutionarily familiar conditions, the earlier elements of this behavioral chain are
performed more often than the later, since the prey may escape at any point before it
is successfully killed.
4
Thus, cats possess drives to perform the later elements of this
behavioral chain with successively lesser frequency. If a cat is locked in a room with
live mice introduced in succession—under conditions where the mice may put up a
fight but never escape—the cat responds in the beginning by performing the whole
sequence of hunting behavior to completion. After it has eaten several mice, it will
kill a few more without eating them. Then the cat will stalk and catch mice for a
while, before letting them go. Finally, pouncing behavior is exhausted, and the cat
4
Even killing movements are not always successful.
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will simply stalk mice for a while until this last appetence is exhausted (Lorenz
1966: 92–93).
Summary and application to a scientific debate
This paper has delimited four types of evolutionary mismatch along with several
subtypes (Table 1). This section seeks to illustrate the potential scientific use of this
taxonomy by briefly considering its relevance for the hypothesis about the evolution
of intelligence discussed earlier.
Table 1 Four types of evolutionary mismatch/novelty
Type Subtype Example
Type 1—Un(der)development-
inducing: failure of a trait
with a direct proper function
to develop
Thalidomide in the womb
prevents limbs from
developing; reading from a
young age causes myopia
Type 2—Ineffectiveness-
inducing: failure of a trait to
perform its direct proper
function despite unimpaired
development
Typica peppered moths in a
polluted environment are not
camouflaged
Failure of a learning mechanism
to produce the behavior for the
sake of which it evolved
Apartment-living cats fail to learn
to hunt when they are hungry
Type 3—Misrepresentation-
inducing: misrepresenting or
failing to represent the
environment
Imprinting on the wrong type of
object
Jackdaws raised by humans court
H. sapiens upon reaching sexual
maturity
Failing to imprint Siblings reared apart fail to
imprint sexual disgust on each
other (to avoid inbreeding)
An object emanates key stimuli in
different intensities or
combinations from an
evolutionarily familiar object
Supernormal stimuli (e.g., food
sweetened with refined sugar);
some visual pornography
Key stimuli associated with a
biologically important type of
object are attached to an object
of a different type
Visual pornography; teddy bears;
artificially sweetened, sugar-
free food; Facebook profiles
Misrepresenting situations Many people cooperate in
anonymous, one-shot prisoner’s
dilemma games;
‘‘developmental mismatch’’
Type 4—Misresponse-
inducing: responding
incorrectly to environmental
conditions
Behavioral chains are not
integrated
The four main components of
hunting behavior in apartment-
living cats are not integrated
Responses are elicited with
different frequencies from those
that occurred in evolutionary
experience
Among most inhabitants of
contemporary Western
societies, genuine fear is
elicited much less frequently
than it was in the relatively
violent and dangerous AE
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As noted, some intelligence researchers argue that intelligence in humans, and
potentially other animals, is an adaptation for dealing with evolutionary novelty
(Byrne 1995: 38; Chiappe and MacDonald 2005; Kanazawa 2004a). These
researchers have clearly been referring to Misrepresentation-inducing and Misre-
sponse-inducing evolutionary novelty (Types 3 and 4).
If the cues used by a species to recognize biologically important objects and
situations are rendered unreliable by environmental change (Type 3), and if no
substitute cues are available, natural selection may put a huge premium on general
reasoning ability: Only those organisms capable of interpreting unpredictable,
transient evidence concerning the classification of objects and situations will be able
to navigate the environment successfully. Consider a problem faced by a wide range
of animals: distinguishing ‘‘friends’’ and ‘‘enemies.’’ Most social animals use a
straightforward procedure to solve this problem, relying on key stimuli (e.g., the
individual is/is not a member of my group). When social organization becomes
more complex—as with coalition-forming chimpanzees—relying on simple cues to
represent conspecifics as friends and enemies will not work. Among humans,
enemies may emit every possible sign of friendship until the moment of treachery.
And real friends sometimes act in confusing ways.
Seeing the conditions under which humans distinguish friends and enemies as
evolutionarily novel helps to (partly) unify the social intelligence hypothesis and the
hypothesis that intelligence evolved to deal with evolutionary novelty. Sterelny
(2012: 6) argues that ‘‘[h]ominin cognitive evolution cannot have been driven
mostly by external environmental change, as then we would expect similar
trajectories in other species, and that we do not see.’’ But there may have been
evolutionary novelty in early hominin’s social domain. Changes in social
organization rendered their evolved mechanisms for identifying friends and enemies
unable to perform their proper functions. Since there are no reliable cues that could
be used to determine whether a would-be ally will remain faithful to you over the
course of a long-term political campaign (such as existed in the AE), general
reasoning ability is the only faculty that could help one determine whom to trust and
to what extent.
Conclusion
The method for cataloging evolutionary mismatch given here does not by itself
resolve any scientific debates. But it may allow hypotheses concerning the effects of
mismatch in Darwinian medicine (Williams and Nesse 1991), evolution-informed
clinical psychology (Gilbert and Bailey 2000), and other fields to be formulated
clearly. In regard to intelligence research, it could help make predictions about
which species are likely to have undergone selection for general reasoning ability.
What consequences the different types of mismatch have for species’ evolution and
well-being are empirical questions—which can be investigated in the light of a clear
mismatch concept.
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Acknowledgments I benefited from numerous conversations on this topic with Neven Sesardic, as well
as feedback from him on several drafts of the paper. Thanks to Ruth Millikan, Elliott Sober, Jiji Zhang,
and two reviewers for Biology & Philosophy for helpful comments on earlier drafts.
Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, dis-
tribution, and reproduction in any medium, provided you give appropriate credit to the original
author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were
made.
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