Ageless Apes:

Life Histories and Evolutionary Histories in the

Evolution of Language
David A. Leavens1, Mahmoud M. Elsherif2, & Hannah Clark1,3
1University of Sussex, 2University of Birmingham, 3University of Portsmouth

The Problem State of the Literature

Ape-human comparisons are poorly matched on many We found 109 ape-human comparisons from 1930 through
variables, including age (Leavens et al., 2019). 2022 that used children (Leavens et al., 2023).

Therefore, alleged ‘species differences’ in cognition Only 11 (10%) reported the life history stage of the apes.
between apes and humans might be due to lurking
variables, rather than differences in evolutionary histories. Only 9 (8%) matched the humans and apes on age.

Table 1: Ape-human comparisons (1930-2022; N = 109)

Human Ape
Matched
Human Ape
Matched
Human Ape
Matched 60
on Life on Life on Life
Year Title Life Life Year Title Life Life Year Title Life Life
History History History
History History History History History History
Stage? Stage? Stage?
Learning from others’ mistakes?
Limits on understanding a trap-tube The origin of representational drawing:
The mentality of a child compared with
1930 Child - Noa 2007 task by young chimpanzees (Pan Children Young Yesm 2014 A comparison of human children and Children - No
that of apes
troglodytes) and children (Homo chimpanzees 50
sapiens)

Form discrimination in chimpanzees Two-year-

Reliance on head versus eyes in the
gaze following of great apes and
Children conform to the behavior of Non-matched (n = 101)
1933 and two-year-old children: II. Form old - No 2007 Infants - No 2014 peers; other great apes stick with what Children - No?q
versus background chldren
human infants: the cooperative eye
hypothesis
they know Matched (n = 8)
Number of Papers

1933
Form discrimination in chimpanzees Two-year-
and two-year-old children: I. Form old - No 2007
Spontaneous sltruism by chimpanzees Young
and young children children
- No
Coordination strategies of
2014 chimpanzees and human children in a Children - No 40
(triangularity) per se chldren Stag Hunt game
Observational learning in Differences in the ability of apes and
The perception of movement by young
1958 Children Young Nob 2008 chimpanzees and children studied Children - No 2015 children to instruct others using Children - No
chimpanzees and human children
through ‘ghost’ conditions gestures
Looking ahead? Computerized maze
Ability of chimpanzees to respond to
Rational tool use and tool choice in
task performance by chimpanzees
(Pan troglodytes), rhesus monkeys
30
1978 symbols of quantity in comparison with Children - Noc 2008 Infants - No 2015 Children - No
human infants and great apes (Macaca mulatta), capuchin monkeys
that of children and of monkeys
(Cebus apella), and human children
(Homo sapiens)

How young children and chimpanzees Chimpanzees (Pan troglodytes) and

Object permanence in child and (Pan troglodytes) perceive objects in a Young human children (Homo sapiens) know
1980
chimpanzee
Child - Yesd 2008
2D display: putting an assumption to children
- No 2015
when they are ignorant about the
Children - No
20
the test location of food

Partitioning the influence of level from Focusing and shifting attention in

Tracking and inferring spatial rotation
1982 rate factors on the performance of Children - Yese 2008 Children - Non 2015 human children (Homo sapiens) and Children - No
by children and great apes
children and apes on a cognitive task chimpanzees (Pan troglodytes)
Prelinguistic infants, but not
1982
Logical and illogical errors made by
apes and children on a cognitive task
Children - No?f 2009 chimpanzees, communicate about
absent entities
Prelinguist
ic infants
- No 2015
Communication about absent entities
in great apes and human infants
Infants - No 10
Spatial construction skills of
The effects of being watched on
Development of manipulations with chimpanzees (Pan troglodytes) and Young . . .
1983 Infants Young Young 2009 - No 2016 resource acquisition in chimpanzees Children - Non
objects in ape and human infants young human children (Homo sapiens children
and human children
sapiens)
The communicative context of
1984
object
manipulation in ape and human
Infant Young Young 2009
A competitive nonverbal false belief
task for children and apes
Children - Non 2016
Comprehension of iconic gestures by
chimpanzees and human children
Children - No 0
adult-infant pairs
1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s 2010s *2020s
The structure of individual differences
Language comprehension in ape and Sensitivity to relational similarity and
1993 Child - No 2010 in the cognitive abilities of children and Children - No 2016 Children - No
child object similarity in apes and children

Processes of social learning in the tool

chimpanzees

Reaching around barriers: the 3-5-year- Children’s and apes’ preparatory

Decade
1993 use of chimpanzees (Pan troglodytes) Children - No 2010 performance of the great apes and 3– old - Noo 2016 responses to two mutually exclusive Children - No
and human children (Homo sapiens) 5-year-old children children possibilities

1993
Imitative learning of actions on objects
by children, chimpanzees, and
enculturated chimpanzees.
Children - No 2010
Two-year-old children copy more
reliably and more often than
nonhuman great apes in multiple
Two-year-
old
chldren
- No?p
Great apes and children infer causal
2016 relations from patterns of variation and Children
covariation
- No
Figure 1: Apes and humans rarely matched on age.
observational learning tasks
Executive function in young children
Young children, but not chimpanzees,
Levels of causal understanding in and chimpanzees (Pan troglodytes): Young Young
1994 Children - Yesg 2011 - No 2017 are averse to disadvantageous and - No
chimpanzees and children Evidence From a nonverbal children children

The Horowitz Validation Protocol

advantageous inequities
dimensional change card sort task

Joint attention and imitative learning in Different social motives in the gestural Prelinguistic human infants and great
Prelinguist
1995 children, chimpanzees, and Children - No 2011 communication of chimpanzees and Children - No 2017 apes show different communicative - No
ic…infants
enculturated chimpanzees human children strategies in a triadic request situation

Nonhuman primates do declare! A Preparatory responses to socially

Imitative learning of artificial fruit
comparison of declarative symbol and determined, mutually exclusive
1996 processing in children (Homo sapiens} Children
and chimpanzees (Pan troglodytes)
- No 2011
gesture use in two children, two
bonobos, and a chimpanzee
Children - No?q 2017
possibilities in chimpanzees and
children
Children - No
Horowitz (2003) devised a procedure to validate
Comparing children’s Homo sapiens
1997
Exploitation of pointing as a referential
gesture in young children, but not
adolescent chimpanzees
Young Adolesc
children ent
No 2011
and chimpanzees’ Pan troglodytes
quantity judgments of sequentially
presented sets of items
Children - No 2017
Chimpanzees, bonobos and children
successfully coordinate in conflict
situations
Children - Nok
experimental protocols when apes and humans have not
1997
Comprehension of novel
communicative signs by apes and
human children.
Children - No
Carryover effect of joint attention to
2011 repeated events in chimpanzees and
young children
Young
children
- No
Children, chimpanzees, and bonobos
2017 adjust the visibility of their actions for
cooperators and competitors
Children - Nok been matched on life history stage:
Distinguishing intentional from Comparing the performances of apes
accidental actions in orangutans (Gorilla gorilla, Pan troglodytes, Pongo Testing differential use of payoff-
1998 (Pongo pygmaeus), chimpanzees
(Pan troglodytes), and human children
(Homo sapiens)
Children - No 2011 pygmaeus) and human children
(Homo sapiens) in the floating peanut
task
Children - Non 2017 biased social learning strategies in
children and chimpanzees
Children - No
1. Assume that human adults have the most
1998
Use of experimenter-given cues
during object-choice tasks by
chimpanzees (Pan troglodytes), an Infants - No 2011
Collaboration encourages equal
sharing in children but not in Children - Nok
Learning the rules of the rock–paper–
2018 scissors game: chimpanzees versus Children - No
sophisticated cognitive processes.
orangutan (Pongo pygmaeus), and chimpanzees children
human infants (Homo sapiens)

1999
A nonverbal false belief task: The
performance of children and great
apes
Children - No 2011
Children, but not chimpanzees, prefer
to collaborate
Children - No 2018
Preschool children and chimpanzees
incur costs to watch punishment of
antisocial others
Children - No
2. Administer protocols previously applied to young
Comprehension of seeing as a
1999 referential act in young children, but
not juvenile chimpanzees
Object permanence in orangutans
Young
children
Juvenil
e
No 2011
A comparison of temperament in
nonhuman apes and human infants

Differences in cognitive processes

Infants - No 2019
Chimpanzees and children avoid
mutual defection in a social dilemma

Human children but not chimpanzees

Children - Nok
children and older apes to young, human adults.
(Pongo pygmaeus), chimpanzees
2001 Children - No 2012 underlying the collaborative activities Children - No 2019 make irrational decisions driven by Children - Non
(Pan troglodytes), and children (Homo
of children and chimpanzees social comparison
sapiens)
Majority-biased transmission in The “sh-ape bias” in non-linguistic

Interpretations
The sources of skill in seriating cups in
2002 Children - No 2012 chimpanzees and human children, but Children - No 2019 categorization: Comparisons between Children - No
children, monkeys and apes
not orangutans children and other apes

Direct and indirect reputation

Referential understanding of videos in
formation in nonhuman great apes Helping in young children and
chimpanzees (Pan troglodytes),
2003 Children - No 2013 (Pan paniscus, Pan troglodytes, Children - No 2019 chimpanzees shows partiality towards Children - No
orangutans (Pongo pygmaeus), and

Interpretation 1: If human adult response profiles match

Gorilla gorilla, Pongo pygmaeus) and friends
children (Homo sapiens)
human children (Homo sapiens)

Can chimpanzee infants (Pan

Understanding communicative Chimpanzees monopolize and
2005
troglodytes ) form categorical
representations in the same manner
as human infants (Homo sapiens )?
Infants Young Young 2013 intentions and semiotic vehicles by
children and chimpanzees
Children - No 2019 children take turns in a limited
resource problem
Children - No
the human children in the original study, then the original
Mental rehearsal in great apes (Pan Remembering in tool-use tasks in Spontaneous categorization of tools
2005 troglodytes and Pongo pygmaeus)
and children
Causal knowledge and
Children - No?h 2014 children and apes: The role of the
information at encoding
Children - No 2019 based on observation in children and
chimpanzees
Chimpanzees help others with what
Children - No
test is validated for discriminating between species, albeit
imitation/emulation switching in Younger apes and human children Younge
2005
chimpanzees (Pan troglodytes) and
children (Homo sapiens)
Copying results and copying actions in
Children - Noi 2014
plan their moves in a maze task
Children
r
No 2019 they want; children help them with
what they need
Children - No
this does not isolate evolutionary history.
the process of social learning: Prospective memory in children and Children, but not great apes, respect
2005 Children - No 2014 Children - No 2019 Children - Nok
chimpanzees (Pan troglodytes) and chimpanzees ownership

Interpretation 2: If human adult response profiles match

human children (Homo sapiens)
Self-awareness in human and
How chimpanzees and children
chimpanzee infants: What is Children, but not chimpanzees, have
2006 Infants Infants Noj 2014 Children - No 2020 perceive other species’ bodies: Children - No
measured and what is meant by the facial correlates of determination

apes in the previously published study, this implies that the

Comparing the expert effect
mark and mirror test?
Differences in the early cognitive Human children, but not great apes,
Altruistic helping in human infants and
2006 Infants Young No 2014 development of children and great Children - Yes 2020 become socially closer by sharing an Children - No
young chimpanzees

Do chimpanzees (Pan troglodytes)

and 2-year-old children (Homo 2-year-old
apes

Cross-species variation in gaze

following and conspecific preference Infants, r
experience in common ground

The strategies used by chimpanzees

(Pan troglodytes) and children (Homo
original study misinterpreted the group difference as
2006 - No 2014 - No 2020 Children - No
sapiens) understand double invisible children
displacement?
among great apes, human infants and
adults
adults sapiens) to solve a simple
coordination problem

The contingency symmetry bias

evidence for human cognitive superiority.
Public information use in chimpanzees (affirming the consequent fallacy) as a Pre-
Cooperative activities in young Young
2006 - No 2014 (Pan troglodytes) and children (Homo Children - No 2021 prerequisite for word learning: A linguistic - No
children and chimpanzees children
sapiens) comparative study of pre-linguistic …infants
human infants and chimpanzees

Tracking the displacement of objects:

A series of tasks with great apes (Pan
Young Differences in the nonverbal requests Great apes and human children
References
2006 troglodytes, Pan paniscus, Gorilla - No 2014 Infants - No 2022 Children - Nos
gorilla, and Pongo pygmaeus) and
young children (Homo sapiens)
children of great apes and human infants rationally monitor their decisions
Horowitz, A. C. (2003). Do humans ape? Or do apes human? Imitation and intention in humans
Apes’ and children’s understanding of
Human children rely more on social
Joint attention in human and
(Homo sapiens) and other animals. Journal of Comparative Psychology, 117, 325–336.
2006 cooperative and competitive motives Children - No 2014 Children - No 2022 chimpanzee infants in varied Infants Young Young
in a communicative situation
information than chimpanzees do
socio‐‐ecological contexts Leavens, D. A., Bard, K. A., & Hopkins, W. D. (2019). The mismeasure of ape social cognition.
Faithful replication of foraging
All great ape species (Gorilla gorilla,
Pan paniscus, Pan troglodytes, Pongo
Two-and-
a-half- The structure of executive functions in Preschool
Animal Cognition, 22, 487-504.
2006 techniques along cultural transmission Children - Nok 2014 abelii) and two-and-a-half-year-old - No 2022 - No
chains by chimpanzees and children children (Homo sapiens) discriminate
appearance from reality
year-old
children
preschool children and chimpanzees children
Leavens, D. A., Elsherif, M., & Clark, H. (2023). What animals can tell us about attentional
Titular Both
prerequisites of language acquisition. [Special issue: N. Gonthier, O. Vasileva, & P.
Matche
2006
Push or pull: Imitation vs. emulation in
great apes and human children
Children - Nol
Notes: See Leavens et al. (2022) for Mention of
notes and references Ape Life
d Life
History
Titular
Mention & Zywiczynsk (Eds.), Eco-evo-devo approaches to language and communication.] Language &
History Matched
Communication, 92, 55-73.