Convergent minds: the evolution of
cognitive complexity in nature
rsfs.royalsocietypublishing.org
Introduction
Cite this article: Powell R, Mikhalevich I,
Logan C, Clayton NS. 2017 Convergent minds:
the evolution of cognitive complexity in nature.
Interface Focus 7: 20170029.
http://dx.doi.org/10.1098/rsfs.2017.0029
One contribution of 12 to a theme issue
‘Convergent minds: the evolution of cognitive
complexity in nature’.
Subject Areas:
biocomplexity, systems biology
Author for correspondence:
Russell Powell
e-mail: rpowellesq@gmail.com
Russell Powell1, Irina Mikhalevich2, Corina Logan3 and Nicola S. Clayton4
1
Department of Philosophy and Center for Philosophy of Science, Boston University, Boston, MA, USA
2
Berlin School of Mind and Brain, Humboldt University, Berlin, Germany
3
Department of Zoology, and 4Department of Psychology, University of Cambridge, Cambridge, UK
RP, 0000-0002-9893-1721; CL, 0000-0002-5944-906X; NSC, 0000-0003-1835-423X
Stephen Jay Gould [1,2] argued that replaying the ‘tape of life’ would result in
radically different evolutionary outcomes. Gould’s ‘radical contingency thesis’
focused primarily on animal morphology, and in particular on the specific par-
ameters of animal body plans, which, he argued, reflect but a small fraction of
the larger set of equally functional morphological possibilities that for historically
contingent reasons were never actualized. Were the tape replayed from different
crucial junctures in the history of life, Gould contended, animal evolution would
be channelled into radically different pathways, its topography would assume a
markedly different shape, and many properties associated with extant animal
life would never have arisen. Imagine how the history of animal life would
have unfolded, for example, were it not for the improbable survival of ancestral
chordates like Pikaia, which gave rise to all modern vertebrates, or for the fortui-
tous persistence of the lobe-finned fish that would eventually give rise to
tetrapods, or for the asteroid-induced extinction of the non-avian dinosaurs
whose emptied niches permitted the unlikely radiation and macro-faunal
dominance of mammals. These examples suggest that the radical contingency
thesis is best interpreted as a ‘modal’ thesis, that is, as a claim about the instabi-
lity of certain evolutionary outcomes across possible evolutionary worlds. More
precisely, it is a claim about the causal dependency of evolutionary outcomes
on small changes in initial conditions, such as quirky survival, extinction or
mutation events, particularly in the early stages of the evolution of higher taxa
like phyla, classes and orders [3,4]. This sensitive dependency explains other
features commonly attributed to the Gouldian view of macroevolution, such as
unpredictability [5,6], stochasticity [7] and path-dependency [8].
Although the radical contingency thesis was aimed at the morphological
features of life, it is particularly provocative for the implications it might have
for the emergence of sophisticated forms of cognition. Is the evolution of mind
a historical accident, unlikely to be replicated across alternative histories of life,
much as Gould argued about animal body plans? Or do we have reason to
think that mind and its associated cognitive competencies is a robustly replicable
feature of the evolutionary process? Given that we are privy to only a single
history of life, how can we even begin to adjudicate such questions? One promis-
ing avenue for investigating evolutionary contingency in general is to examine
patterns of convergent evolution—the independent origination of similar biologi-
cal forms and functions [5,6,9–11]. Under certain conditions, convergence may
constitute natural experimental replications that provide evidence for the robust-
ness, predictability and/or path-independency of the iterated outcomes observed
[4]. Although the term ‘convergence’ is typically reserved for descriptions of mor-
phological character states, the concept can usefully be extended to functional
properties such as cognition [12]. We may therefore investigate the replicability
of cognitive capacities of varying degrees of complexity, as well as the develop-
mental and ecological conditions under which particular cognitive forms
are likely to arise, by establishing the extent and phylogenetic distribution of
cognitive convergence [12,13].
If patterns of cognitive convergence are to serve as evidence for robust evol-
utionary replicability, however, it is critical that the iterated evolution of similar
& 2017 The Author(s) Published by the Royal Society. All rights reserved.
cognitive capacities be demonstrated in taxonomically distant
groups of animals. This is because the evolutionary distance
of converging lineages serves as a natural experimental control
for conserved (homologous) developmental variables that
could potentially undermine the independence of iterations
and thus restrict the generalizability of any regularities
observed. If convergence is phylogenetically deep, then
shared developmental constraints are unlikely to be respon-
sible for the observed repetitions. For instance, if we observe
certain complex cognitive properties in lineages whose
common ancestor in all probability lacked complex brain struc-
tures or even a centralized nervous system—then this indicates
the evolutionary replicability of these properties across tem-
porally deep replays of the tape of life (e.g. across alternative
histories of animal life), and in some cases could indicate
their law-like necessity in any complex multicellular world.
Convergence across great taxonomic distances is also crucial
for establishing the ecological causes and evolutionary conse-
quences of cognitive evolution, as well as the developmental
substrates that facilitate and constrain the emergence of
complex cognitive forms. By controlling for many (though
perhaps not all) phylogenetic constraints, the taxonomic
breadth of convergent iteration increases our confidence that
any regularities observed will hold up across disparate initial
conditions and phylogenetic contexts. In essence, convergence
can indicate that certain cognitive forms are less sensitive
to boundary conditions, less path-dependent, and less unpre-
dictable than the radical contingency thesis might have us
believe. Do patterns of cognitive convergence support a
‘robust replicability thesis’ of mind?
It is crucial that any study of cognitive convergence that
aims to draw broad lessons about the evolution of mind—
not only about its replicability, but also about the specific
developmental and ecological conditions under which it is
likely (and unlikely) to occur—must look to convergence
across higher taxa, both within chordates and between chor-
dates and other phyla in Bilateria, especially arthropods and
molluscs. On prevailing phylogenetic reconstructions, arthro-
pods and molluscs share a common ancestor with vertebrates
that in all probability did not possess a centralized nervous
system or even vision, both of which are thought to have
evolved independently in these groups after their divergence
over 500 million years ago in the Cambrian (notwithstanding
conserved genes and embryonic patterning mechanisms
involved in the ontogeny and evolution of all known nervous
systems) [14]; but see [15]. A growing body of comparative
cognition research on molluscs (in particular, cephalopods;
e.g. [16]) and arthropods (especially insects and arachnids;
e.g. [17–19]) suggests that complex cognition did not arise
solely in the vertebrate clade—and thus, contra Gould,
would not have gone un-actualized had Pikaia and its ilk
been less fortunate. Similarly, certain bird and mammal
clades, which share a common ancestor that lived over
300 million years ago in the Carboniferous, exhibit convergently
enlarged brain structures [20] and a convergent cognitive tool
kit that includes capacities such as causal reasoning, flexible
problem solving and imagination [12,21,22]. Similar patterns
of convergence can be seen at finer grains of taxonomic resolu-
tion, including between numerous mammalian lineages
such as cetaceans, primates, elephants, and carnivores [23],
indicating predictability and path-independency across
shallower—though still quite substantial—replays of the tape
of life.
The independent evolution of complex cognition in the 2
many millions of years since these groups separated from a
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cognitively simpler common ancestor is putative evidence
of the power of natural selection to drive convergent cogni-
tive evolution from highly disparate initial conditions [24],
and thus provides a forceful counterexample to the radical
contingency thesis as it relates to some complex forms of
cognition. Some have argued that certain types of cognition
are uniquely human, such as shared intentionality, moral
norm acquisition, and other cognitive abilities that underpin
ultra-cooperation [25] and cumulative cultural transmission
[26] in humans—which, if truly singular, could indicate the
Interface Focus 7: 20170029
highly contingent origins of these cognitive forms. Gould
himself made a case for the radical contingency of human-
specific cognition, while withholding judgement on the evol-
utionary replicability of more general forms of intelligence
[27]. Yet the fact that there is currently little evidence of
these particular cognitive abilities in nonhuman animals
does not indicate that they are entirely absent in nonhuman
clades, since very little testing has been conducted and the
efficacy of experimental probes has been limited. In addition,
the interplay of biology and culture in the development of
such capacities is unclear, and the extent of culture in nonhu-
man animals is itself the subject of intensive investigation
[28]. We must therefore keep an open mind in asking: What
do patterns of convergence in the history of life on Earth
tell us about the nature of mind, its evolution and its place
in the universe?
This themed issue brings together an international group
of leading researchers and theorists from across the natural
sciences and from the philosophy of biological and cognitive
science to explore the iterated evolution of cognitive complex-
ity. Conceptual problems loom large in this discussion. For
starters, what counts as evidence for the evolutionary replic-
ability of mind will depend on how we define the relevant
explanandum (or ‘reference class’). This poses a particular
challenge in the case of cognition, a phenomenon about
which there is no widespread agreement regarding its defi-
nition, its typology, or in virtue of which properties some
types of cognition may be said to be more complex than
others. On ‘big-tent’ accounts of cognition, such as infor-
mation processing accounts, minimal cognitive processes
are present in nearly all forms of life including prokaryotes
and other unicellular organisms, with cognitive processes
greying into metabolic ones [29]. Other more restrictive but
still biologically broad-based accounts of cognition hold
that minimal cognition consists in sensorimotor mechanisms
that are organizationally decoupled from metabolic processes
and thus support faster sensorimotor information flows—a
function paradigmatically realized by nervous systems. Yet
there are functionally analogous sensorimotor systems in
plants, in unicellular eukaryotes, and even in bacteria—all
of which exhibit functional memory and flexible stimulus –
response pairing mechanisms that are distinct from metabolic
processes [30]. Both bacteria and plants have been shown to
exhibit complex behavioural repertoires with a degree of
flexibility that was long presumed to require a functional ner-
vous system. According to ‘bottom-up’ accounts of cognition,
whether glossed in terms of general information processing
or more specific sensorimotor information flows, cognition
is not restricted to complex nervous systems or even to ner-
vous systems at all—and thus it can be expected to be a
universal feature of life wherever it evolves.
 This bottom-up approach is championed by Keijer [31]
who argues that the nature of cognition is to be discovered
by examining the biological world, rather than decided
a priori by appealing to anthropocentric intuitions. Drawing
on recent findings demonstrating the complex behavioural
repertoires of protists, plants, and fungi, Keijer proposes a
notion of ‘biologically embodied cognition’, which combines
biology’s bottom-up approach to cognition with an embodied,
rather than a computational-representational and brain-bound,
view. Van Duijn [32] does not take a stand on whether cogni-
tion is essentially computational or embodied, but he too
draws on evidence from both unicellular and multicellular
aneural organisms to challenge what he sees as the ‘cerebro-
centrism’ of cognitive science and, by extension, comparative
psychology. Evidence suggests that these aneural organisms
exhibit basic forms of learning and possess complex molecular
mechanisms of information transfer, which undermines the
assumption that cognition requires nervous systems, let alone
brains. Trewavas [33] a leading authority on plant cognition,
offers a wide-ranging exploration of the mechanisms of infor-
mation transfer that underwrite ‘intelligent’ behaviour in
plants. He concludes that all organisms whose survival depends
on coping with a complex environment must evolve ‘intelli-
gence’, by which he means a set of sophisticated cognitive
capacities that underwrite flexible responses to environmental
contingencies. Cognition, on each of these ‘bottom-up’ accounts,
can be expected to be biologically ubiquitous and modally
robust (i.e. stable across alternative evolutionary histories).
Other cognition theorists opt for a ‘top-down’ approach
that conceives of the replicability of mind more narrowly to
refer to the robustness not simply of cognition, but of cognitive
capacities that are canonically deemed complex—such as plan-
ning, insight, imagination, causal reasoning, metacognition,
intentionality, self-recognition, consciousness, and the like—
capacities that are not present, even in rudimentary forms, in
the vast majority of organisms on Earth (only a tiny fraction
of which are multicellular organisms). With a top-down
approach, questions regarding the contingency and predict-
ability of mind become in a sense more interesting and more
pertinent, but also more difficult to evaluate. Research in the
expanding field of comparative cognition purports to identify
numerous instances of convergence on these more demanding
cognitive properties.
A number of researchers from disparate disciplines have con-
tributed to this theme issue on convergent minds, and several of
these authors follow this top-down tradition in probing for the
presence of sophisticated cognitive capacities such as imagin-
ation, proto-linguistic communication, and counting abilities in
lineages as phylogenetically disparate as birds, primates, and
arachnids. Cross & Jackson [34] detail an experimental test for
numerical cognition in the Portia jumping spider, from which
they conclude that these arachnids appear to represent the quan-
tities ‘1’ and ‘2’ as discrete numerical categories; moreover, they
likely discriminate among these through a method other than,
for example, relying on discriminating between the timing of
different sets of events. Their experimental set-up relies on the
spiders’ well known ability to select the most effective path
(which is not necessarily the most direct one) in pursuit of
prey, which is itself suggestive of the possession of a cognitive
map, or an internal representation of the relationship among
objects in an environment. Numeric cognition, cognitive maps
and other complex representations are often thought to be
beyond the reach of tiny-brained animals such as insects and
arachnids [35], despite recent evidence to the contrary 3
[36,37]—though the status of cognitive maps and other forms
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of spatial cognition in arthropods remains unsettled [19].
Moore [38] meanwhile, makes the case that animal com-
munication bears a much stronger resemblance to human
language-mediated communication by reinterpreting the
demands of the prevailing theory of successful communi-
cation. According to Moore, the ability to direct a message to
an intended audience ‘in pursuit of a communicative goal’ is
found in many nonhuman animals and is likely to have
evolved many times in the history of life. Finally, Clayton &
Wilkins [39] draw simultaneously on Clayton’s extensive
Interface Focus 7: 20170029
body of work on episodic-like memory in jays and on Wilkins’s
literary exploration of the relation between memory and iden-
tity to make the case that the ability to remember past events to
plan future events likely evolved to aid in prospection rather
than recall. If they are correct, then evidence of episodic-like
memory in jays may simultaneously constitute evidence of
future-oriented cognitive abilities, such as planning. Moreover,
they suggest that engaging science with narrative art may
present a novel method for exploring questions about the
unobservable phenomena that are the subject of study in
comparative cognition.
A central research question in comparative cognition
concerns the evolutionary and ecological circumstances that
give rise to complex forms of cognition. Under what conditions
does complex cognition tend to evolve and what is it used for?
The prevailing theory is that cognitive evolution is an adaptive
response to environmental complexity. Several contributors to
this issue build on or test elements of this hypothesis in an
effort to explore the ecological and evolutionary correlates of
cognitive evolution. Dunlap et al. [40] investigate whether
bumblebees modify their information-seeking behaviour
in response to a changing and unstable foraging ecology.
Holekamp & Benson-Amram [41] investigate the source
of spotted hyaenas’ social intelligence by evaluating the
‘cognitive buffer hypothesis’. According to this hypothesis,
domain-specific cognition such as social intelligence is a
by-product of the selection for general intelligence, which is
required to cope with a changing environment. The cognitive
buffer hypothesis challenges the social intelligence hypothesis,
according to which gregarious animals acquired their social
intelligence (and correspondingly larger brains) in response
to the demands of social living. If general intelligence has its
casual-etiological roots in the demands of ecological complex-
ity, then we should expect to find elements of intelligence in
all organisms that face such demands, barring constraints or
contingencies to the contrary.
Studies of cognitive convergence, though compelling, are
fraught with conceptual and methodological difficulties that
often make their findings difficult to interpret. For instance,
how precisely should we delineate the cognitive traits that
are putatively converged upon, and at what level of description
are they convergent? Additionally, it is widely acknowledged
that convergence judgements are relative to hierarchical
level—a trait may be convergent at the level of macro-
morphology but homologous at the level of tissues, proteins,
or genes. Likewise, complex cognition can be convergent at
the level of cognitive mechanisms, while arising from diver-
gent neuroanatomical structures (e.g. the cortex in mammals
and the telencephalon in birds) that are comprised of homolo-
gous neural substrates and formed by conserved patterning
mechanisms. McShea [42] aims to pick apart the cognitive
traits that are convergent, making a Humean case for the con- take up the question as to how sources of evidence beyond 4
ceptual decoupling of motivational and affective faculties the behavioural experimental setting—in particular, evolution-

(‘wants-preferences-cares’), on the one hand, and intellectual
faculties (‘logic-calculation-problem-solving’), on the other.
He argues that both are crucial yet distinct causal components
of the behaviour typically associated with cognition, and he
points out that the evolutionary histories of these two distinct
faculties can be dissociated.
Establishing convergent cognitive mechanisms is an
especially challenging task, because unlike other phenotypic
traits such as morphological or even behavioural characters,
cognition implicates unobservable processes and hence must
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ary and ecological theory and data—can inform hypothesis
choice and experimental design in comparative cognition.
Rejecting a blanket preference for simple explanations,
they propose an evolutionary–ecological model according to
which the repeated emergence of trait-clusters including
(i) flexible behaviours, (ii) complex neuroanatomy and
(iii) heterogeneous selective environments can be used to infer
complex cognitive abilities because they comprise a non-acci-
dental convergent regularity. This convergent regularity, in
turn, may be used to support the inference to one trait (e.g. com-

Interface Focus 7: 20170029

be inferred from behavioural observations. Some convergent
behavioural traits, such as the ‘agriculture’ phenotype in ants
and humans, are unlikely to be underpinned by similar cogni-
tive mechanisms, even if some prominent discussions of
convergence have loosely described such outcomes as conver-
gent minds (e.g. [5]). However, many observed animal
behaviours can be explained equally well by simpler and
more complex cognitive mechanisms, and thus the logical
structure of the inference from observed behaviours to the
presence of particular cognitive mechanisms remains under-
specified. In some cases, a preference for simpler cognitive
explanations may be justified by additional sources of evidence
(e.g. neuroanatomy) conjoined with background theoretical
assumptions (e.g. about the relation between neuroanatomy
and cognition). For example, Cartmill [43] argues that
behavioural convergence need not be evidence of cognitive
convergence for animals whose neurophysiology is unlikely
to sustain intentionality and consciousness—two features of
mind that he takes to be of particular interest in the quest to
discover cases of convergent cognition. Thus, the tiny brains
(in terms of total number of neurons) of jumping spiders rule
them out as plausible contenders of cognitive convergence
even though, as Cartmill notes, their predatory behaviour,
which involves alternating among attack strategies such
as stalking, ambushing, and frontal attacks, is as flexible as
those of large cats. For similar reasons, Cartmill urges scepticism
in evaluations of cephalopod behaviour (e.g. puzzle-solving
behaviours of octopuses) given the radically alien organization
and developmental history of cephalopod brains.
Cognitive models, neuroanatomy and broader evolution-
ary–ecological considerations may therefore be drawn upon
to strengthen or weaken inferences of cognitive complexity;
but precisely how these respective lines of evidence should
be weighted and integrated is unclear. Mikhalevich et al. [44]
plex cognition) whenever lineages exhibit two or more of the
other traits in the cluster. On this view, while, for example,
the jumping spider may possess just a few hundred thousand
neurons, its behavioural flexibility, heterogeneous ecology
and comparatively encephalized neuroanatomy collectively
speak in favour of the convergent complexity of its cognitive
toolkit. A direct prediction of this model is that complex cogni-
tive capacities will begin to degrade under relaxed selection in
less informationally demanding environments—which agrees
with Cartmill’s reminder that convergences include convergent
losses, such as decreased encephalization in response to relaxed
predation pressures, and that these, too, deserve our theoretical
and empirical attention.
In sum, the time is ripe for a conceptually rich, multi-
disciplinary investigation into the evolution of cognition in
its multiplicity of forms. It is only by examining cognitive con-
vergence—and the lack thereof—in clades as phylogenetically,
developmentally, and ecologically disparate as corvids,
whales, carnivores, primates, cephalopod molluscs, jumping
spiders, insects, green plants, fungi, and unicellular organisms,
that we can begin to map out the phylogenetic distribution of
cognition as well as its evolutionary drivers, constraints and
consequences. As this broad-based picture begins to resolve,
it will illuminate the interplay of contingency and necessity
in the evolution of cognitive worlds.
Competing interests. We declare we have no competing interests.
Funding. R.P. is grateful to Templeton Foundation grant no. 43160 and
C.L. thanks the Leverhulme Trust and Isaac Newton Trust for a
Leverhulme Early Career Research Fellowship, which supported
this research.
Acknowledgements. We would like to thank the contributors to this
theme issue for their fascinating and provocative papers, as well as
the editors of Interface Focus, especially Tim Holt, for invaluable
editorial support.

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