Life History as a Predictor for Language

Severine Hex
Evolution of Language Fall 2016

Introduction
Language has been long considered an evolutionary phenomenon unique to humans, leading to
a torrential body of work surrounding the evolution of this most singular capacity. What has only
exacerbated the apparent mystery surrounding the origins of human language is its seeming disparity
with the existing vocal communication systems of our closest phylogenetic relatives. Such seemingly
unsurmountable differences in the nature of human vocal communication compared with those of
other apes have even led some researchers to hypothesize that the only reasonable explanation would
be if a sudden mutation had occurred relatively recently within the human lineage, enabling languages
emergence (Chomsky, 2010). Such views are formed under the assumption that the human brain
evolved to suit language. If this assumption were taken a step farther, the unspoken bias may be that
language could only have emerged within a brain like that possessed by humans, and given these neural
constraints, language may have only emerged in a form similar, in an abstract sense, to that observed in
natural human language. This paper will base its arguments on the reverse argument, namely that
language adapted to suit the brain, and the implications that may be drawn from this stance from a
comparative standpoint (Christiansen & Chater, 2016).
If language did adapt to suit the pre-existing neural machinery of the brain, it would seem
reasonable to hypothesise that should the same pressure for complex communicative capacities have
arisen in other species, then language would have evolved to suit the neural machinery of the species in
question. However, seeing as language is the system under selection instead of the brain, the language
that manifested in two different species under the same evolutionary pressure would be very different.
Under this hypothesis, I propose an answer to an old question: why do only humans appear to possess

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language? The answer would be simply: because only humans possess human language. While this
response may seem asinine, it is important to consider a differentiation between human language and
Language in an abstract sense. If language truly did evolve to suit the brain, then Language must be
considered as a capacity abstracted from that of merely what manifests within the human system, just
as a wing must be abstracted from those possessed by only birds to encompass those also evolved by
insects, bats, and pterosaurs.
To continue with the metaphor of the wing, consider a definition of a wing based entirely on
that of a birds wing. By forming this definition, we would disqualify the wings possessed by other
flying taxa because theirs do not identically match those of a bird. Further, in the study of the
evolution of the bird wing, we only limited our search to the closest taxonomical relatives of birds.
While there exist some gliding lizards and snakes, more distant relatives, crocodilians, birds closest
living relatives, are decidedly unable to fly. We may then be left bemused as to how the bird evolved
wings and the ability to fly. Of course, the above scenarios we know to be criminally absurd. We know
that the wing evolved independently in four distinct taxa, and while the structure may appear different
in each iteration, the function is the same. Convergent paths to the same solution, presumably
prompted by similar evolutionary and environmental pressures, which lead to four very different taxa
taking flight.
I argue that the same thinking must be applied to the evolution of Language if we are to
understand its origin and its current manifestations. This paper will be taking the viewpoint that
language is a tool that may have arisen as an answer to similar environmental problems in various taxa
via convergent evolution. The environmental pressures explored within this paper will specifically be
life history features, features of the animals natural ecology that may have placed a pressure on the
species to develop a complex communicative system. This paper will be exploring the hypothesis that a
predictive manner of searching for and identifying candidates for non-human languages may be
through identifying species that share similar life history features that could have placed pressure on the
species to develop a linguistic system. First, two categories of life history features will be explored, and
their links to the emergence of complex communication and language argued. Then, a few

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phylogenetically diverge exemplar species will be surveyed, and their possession of lack thereof of the
identified life history traits will be examined. Finally, implications and future directions for research and
theoretical consideration will be proposed.

1 Life history features

The examples of life history being more consequential than phylogenetic relatedness in the
emergence of physical or behavioural similarities between species abound. Life history features may be
anything from dietary patterns and number of offspring produced to seasonal changes in behaviour and
mate choice. Because language serves an inherently social function, this paper will focus on life history
features that would exert pressure on the social capacities of the individual. Under this constraint, this
paper will explore two types of social life history traits, group dynamics and breeding system.

1.1 Group Dynamics

While disputed by some, most would agree that language above all else serves a social function.
Language enables an individual to navigate social bonds, maintain and signal social affiliation, and
negotiate social situations. It may be reasonable to hypothesize, given the purpose of language as a
social tool, that the shape of the social environment of an organism may prove a sufficient pressure to
prompt the evolution of a complex communicative system.

More specifically, group size and

complexity, the presence of a stable hierarchy, and/or the presence of a fission-fusion society may be
predictive of social, ecological pressures for the emergence of a linguistic system.
1.1.1 Group Size and Complexity

Group size and complexity are not immediately interchangeable, but rather linked, perhaps even
as multipliers of one another. Group size alone could not be considered necessarily an impetus for the
evolution of a linguistic system, as it is easy to imagine a system in which large aggregations of
individuals assemble but interactions are limited to one or two individuals exclusively, or there is no
repetition of interaction partner. However, it may be difficult to think of a complex network of
relationships without a certain threshold of size met. Therefore, we find the definition of social

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complexity proposed by Freeburg et al. (2012) to be of satisfactory nuance; namely that complexity is a
combination of many individuals interacting over many different contexts with considerable
consistency. An individual in a more socially complex environment would experience pressure to
convey a greater range of motivational or contextual information, express more intricate social
affiliative relationships, or convey a wider range of meanings (Freeburg, Dunbar, & Ord, 2012;
Freeburg 2006).
There has been a fair amount of work in the social calls of songbirds and the effects of social
complexity on the complexity of the emergent vocal communication system. Freeburg (2008) found
that by manipulating the group size of Carolina chickadees (Poecile carolinensis), there was a positive
correlation between group size and the complexity of the chick-a-dee calls produced by flock members,
with members of the larger flocks displaying greater note type variation and variation in combinations
of notes (Freeburg, 2008). This study demonstrates that there may be an effect of the complexity of the
social group on vocal complexity, and in species that tend to form larger social groups, there may exist
pressure to continue to develop a complex linguistic system.
1.1.2 Stable Hierarchy

The presence of a hierarchy may not be considered in isolation of group size. A group of three
animals may have a stable hierarchy, but without many relationships, there would not be a large
pressure to communicate a range of complex social affiliations. However, a large group with a stable
hierarchy would manifest as complex social contexts to navigate and communicate. Even in the
absence of a particularly complex vocal communication system, the sorts of mental pre-requisites for
navigating a convoluted social hierarchy may serve as the substrate into which language may nestle as it
evolves to suit the brain.
Sayfarth and Cheney (2014) found that while baboons do not have a particularly complex vocal
repertoire, due to the complexity of their social hierarchies, they are able to predict how two individuals
ought to interact in an antagonistic situation based solely upon recognizing an individuals call and
placing it in the context of their knowledge of the senders standing within the social hierarchy. In

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playback studies in which a recording of an individual known to occupy a subordinate social rank
produces an antagonistic, dominant call and an individual in typically occupying a more dominant
position produces the call of submission, the listening baboons display behavioural signs of
dishabituation, indicating they had expectations of how the interaction ought to have occurred and
these expectations were violated (Seyfarth & Cheney, 2014). Furthermore, if a playback of a baboon
producing an antagonistic vocalisation is presented, baboons will respond either with indifference or
alarm depending on the quality of their interaction with the baboon whose call was being presented
prior to hearing the call; that is, if the last interaction had been benign or antagonistic. It is believed,
based on these results, that this level of social cognition may be a crucial precursor for language
(Seyfarth & Cheney, 2014). The ability to extract this level of information depending not only upon the
senders identity, but also in context, may be analogous to pragmatic information of human language.
1.1.3 Fission-Fusion Society

This life history feature is potentially, though not necessarily, one that occurs in mutual
exclusivity with the above discussed feature of stable hierarchy. By definition, a stable hierarchy would
take time and experience with the individuals of the group in order for meaningful information to be
retained about the social standing of any given individual. However, fission-fusion societies are fluid by
nature, with the composition of the group varying from day to day. While certainly dominancesubordinate relationships would still exist, and need to be established, there would be no stable context
onto which to map pragmatic information as was found by Seyfarth & Cheney (2014). This sort of
society would instead place pressure on individuals to be able to interact with and attract the attention
of novel conspecifics, as well as maintain contact with conspecifics that may be spatially separated.
To the former challenge various species of parrotlets have developed what are called signature
calls (Bradbury & Balsby, 2016; Balsby & Bradbury, 2006). A signature call is a vocalization that
uniquely identifies an individual. To gain the attention of an intended receiver, an individual will imitate
their social partners signature call. Similar to the cocktail party effect in humans, in playback studies
where the signature call of a parrotlet is played to a group, the most robust and rapid approach

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response is from the one whose signature call had been played (Bradbury & Balsby, 2016; Balsby &
Bradbury, 2006). This could be seen as a referential system, with the referent being the individual and
the symbol used to represent them being their signature call.
A fission-fusion society may also lead to situations in which conspecifics are separated and must
reunite. Contact calls, widely found among both birds and mammals serve this function, allowing an
individual to signal its location in relation to its intended receiver. A feature of many contact call
systems is antiphonal communication, known otherwise in the human literature as turn taking. The
complexity of vocal processing and response formulation is belied by the relative brevity of pauses
between utterances, and yet turn taking is indisputably universal in human language (Levinson, 2016).
In many social species, too, antiphonal communication is seen in interactions among conspecifics, with
latency between vocalizations varying in terms of species, distance of caller from receiver, social status
or sex of the individual, the function of the call, and the experience of the caller (Henry, Craig,
Lemasson, & Hausberger, 2015). Besides being potentially capable of carrying pragmatic information
such as the social rank of the caller, antiphonal communication would also ensure maximal efficiency of
communication by ensuring that mutual intelligibility is maintained between individuals.

1.2 Breeding System

A powerful driver of evolution is sexual selection and the pressures that arise from being able to
successfully rear offspring. Particularly in social species that engage in parental care, there may be
pressure to communicate more effectively with mates and conspecifics in order to maximize the
chances of offspring survival. More complex vocalizations may be able to coordinate and synchronize
activities, as well as solidify and reinforce social bonds.
1.2.1 Monogamy

In socially monogamous species, reproductive success is almost entirely dictated by the ability
of the pair to effectively communicate and maintain their pair bond. This pressure may lead to the
emergence of increasing vocal complexity in monogamous species in order to facilitate synchronization
of activities, and the solidification of bonds. Zebra finches (Taeniopygia guttate) will display changes in the

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patterns of call communication in relation to their reproductive status, with pair-bonded birds changing
the composition of their call repertoire and fine-tuning their vocal interactions with their social partner
(Gill et al., 2015). Changes in the call type combinations within pairs was associated with successful
egg-laying, which may be support for the hypothesis that vocal complexity may facilitate the success of
socially monogamous relationships (Gill et al., 2015). What is striking about these vocal exchanges is
that, unlike vocal duets, another mechanism used by pair bonded birds to maintain bonds or defend
territories, the calls displayed by the zebra finch pairs appeared antiphonal, with any given call answered
after an average .5 second latency (Gill et al., 2015). As was discussed above, antiphonal
communication may be a necessary feature of a linguistic system. It must be noted that social
monogamy may not be the only breeding system that may benefit from complex communicative
abilities. A stable polygyny or harem based social structure may be an alternative breeding strategy to
monogamy that could potentially place a similar pressure on the members to communicate between
themselves for coordination and affiliative purposes.

2 Application across Taxa

Using the above life history features, I intend to test the predictive quality of these traits by
comparing them to the life history features of several well studied exemplar species. These species will
be representatives of a diverse phylogenetic groups, each with documented vocal communicative
complexity. The purpose of this small-scale literature review is to test the hypothesis that language may
have evolved via convergent evolution as answers to similar environmental and life-history pressures.
The species to be surveyed will represent birds, cetaceans, bats, carnivores, and primates. Humans will
be used as a standard comparison (see table below).
The species representing birds is the spectacled parrotlet (Forpus conspicilltus), known for its
unique and complex social structures that include creches for offspring and powerful sibling bonds
playing a fundamental role in socialization. Spectacled parrotlets are also known for their vocal
complexity, most famously work investigating the presence of vocal labelling of family members
(Wanker, Sugama, & Prinage, 2005). Cetaceans are represented by common bottle-nosed dolphins

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(Tursiops truncates), well studied for their social and cognitive complexity, as well as their intricate vocal
communicative system, a feature of which is the presence of signature whistles used to convey
information of individual identity and to attract conspecifics (Janik, Sayigh, & Wells, 2006). Vampire
bats, while not a single species, represent bats due to their complex social structures consisting of nonrandom associations and altruistic behavior, as well as evidence of antiphonal communication enabling
individual identification and a large vocal repertoire (Carter et al., 2008). Spotted hyenas (Crocuta crocuta)
have been selected to represent carnivores, living in large, stable, matriarchically based social groups
and possessing a rich vocal repertoire (Holekamp, Sakai, & Lundrigan, 2007). Finally, gelada baboons
(Theropithecus gelada) have been selected to represent primates, for while their social structure is similar in
complexity to other primates, their vocal repertoire size is markedly larger and attributed to the stable,
harem social structure (Gustison, le Roux, & Bergman, 2012).
Species

Large/complex Stable
social
social group?
hierarchy?
Spectacled
Yes (Wanker et Somewhat; female
parrotlet (Forpus al., 1998)
mate
choice
conspicilltus)
influenced
by
males social status

Fission-fusion
society?
Yes

Common Bottlenosed
dolphin
(Tursiops truncates)
Vampire bats

Yes (Bruno, Politi, No

Spotted
hyena
(Crocuta crocuta)
Geladas baboon
(Theropithecus gelada)

Yes

(Bruno,
Politi, & Bearzi,
2004)

Yes (Kerth,

Perony, &
Schweitzer, 2011)

Yes

1986)

(Frank,

(GarnetzkeStollmann & Franck,

1991
Yes
(Connor,
Heithaus, & Barre,
2001)

(Krasheninnikova,
Brger, & Wanker,
2013)

Yes; though shifts Yes

(Kerth,
&
with shifts in Perony,
Schweitzer,
2011)
tenure
of
dominant males
(Wilkonson, 1985)
Yes (Frank, 1986)

Yes

(GarnetzkeStollmann & Franck,

1991)

& Bearzi, 2004)

No

No; stable harem

for each breeding
season
(Wilkinson,
1985)

No

Yes (Kawai, et Yes (Kawai, et al., Somewhat; there No

al., 1983)
1983)
are non-random
associations
between groups
(Kawai,
1983)

Human
sapiens)

Monogamous?

(Homo Yes

Yes (though varies Yes

with culture)

et

al.,

Somewhat

From the above survey of a few phylogenetically diverse species, the evidence of some life
history features being predictive are immediately apparent. In all species, there is the presence of a both

Life History as a Predictor for Language

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large and complex social structure which would place pressure on the species to evolve the capacity to
potentially communicate a wider range of meanings. All surveyed species also demonstrates some sort
of stable social hierarchy, though in the cases of the vampire bat and the spectacled parrotlet, this was
less stable and evident than that found in the spotted hyena, geladas monkey, and bottlenose dolphin.
Only the spectacled parrotlet, bottlenose dolphin, and vampire bat show evidence of a true fissionfusion society, though apparent non-random associations between groups of one-male geladas baboon
groups suggest the potential existence of a larger social unit above the harem (Kawai, et al., 1983).
Only the spectacled parrotlet showed social monogamy, though the gelada baboon possesses a stable
harem system that may serve as a functional equivalent life-history feature.

3 Implications and Future Directions

While the sample size of surveyed species was small and largely limited by the availability of
relevant life-history information about the species, some patterns indeed seemed to begin emerging.
Foremost and perhaps the least surprising is the discovery that every species possesses a large and
complex social group. Indeed, while there are many theories about the original purpose of language, it
is arguably undeniably that language serves an inherently social function. It would be difficult to
imagine the use for a full language in an organism that does not engage in consistent interactions with
numerous other individuals. Language serves the purpose of mediating social bonds and
communicating information about the world both internal and external to the sender. Therefore, it
seems a fundamental pre-requisite when searching for non-human language cross-phylogenetically to
first run the species through the sift of whether or not they exist in a sufficiently large and complex
social structure to even warrant the evolution of a linguistic system.
The presence of a stable hierarchy seemed at least partially to interact with the existence of a
fission-fusion dynamic. While bottlenose dolphins and humans seem to display both life features,
spectacled parrotlets do not appear to have an obviously linear and stable social structure, and while a
dominance structure exists between males so long as a resident dominant male presides over a roost of
unrelated females, turnover of dominant male is relatively frequent. These facts bring to question

Life History as a Predictor for Language

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whether the presence of both are necessary, or if one seems to have more predictive power over the
other. A larger sample size of exemplar species would be required to refine the resolution of the data.
One that seemed only weakly predictive cross phylogenetically appeared to be monogamy.
While the spectacled parrotlet was socially monogamous and the gelada baboon a stable harem, the
vampire bat, bottlenose dolphin, and hyena are highly polygynous. It is possible that monogamy or
stable polygyny could simply be seen as slight variants on stable social hierarchy, or maybe more
generally, stable social structure. All would introduce the necessity of communicating with one or more
individual on a consistent basis across a variety of contexts, potentially leading to similar evolutionary
pressures.
If we take the viewpoint that language evolved to suit and was shaped by the brain, then we
must consider the possibility that language could have taken dramatically different forms in different
species even if under the same environmental pressures. The goal of future research must be to
identify these environmental pressures and use them to find candidates for non-human language, all
while maintaining the open-mindedness that language as it may have emerged in other species may be
radically different from human language. The challenge that resides over both tasks is that of
abstracting from our potentially inalienable biases as humans, the rub of differentiating between
features that are both necessary and sufficient from those that are simply additional features. To only
make the task more difficult, we must be cautious to prevent our own hubris from guiding our
conclusions and inquiries.
To resurrect the fully exhausted wing metaphor a final time, if the definition of the wing were
constructed by birds themselves, and they looked to the rest of the natural world in search of another
species with wings and flight, would they not do so with the preconceived notion that theirs is the
pinnacle of perfection? And though bats and insects, too, seemed to flap appendages and elevate from
the ground to locomote through the air, since they lacked feathers, could theirs truly be deemed wings?
And though some philosopher birds may concede that their method of locomotion and their
appendages superficially resembled a birds wings, because of painfully obvious and infinitely
consequential differences, the pseudo-wings of other taxa could only be seen as less complex varieties.

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There is a certain hopeless enormity to the task of casting away the assumptions that have
defined centuries of scientific inquiry. However, it is only by resolving to look beyond our immediate
surroundings into the tree of life that we may be able to gain new insights into the evolution of human
language, as well as perhaps gaining a better understanding of what Language is as a whole. By
identifying similarities in life history traits expressed by species that possess complex communicative
systems, we may gain two-fold in scientific understanding. Not only may we be able to focus our
scientific inquiry on furthering our admittedly superficial understanding of the complexities of these
systems, but we may also gain in flickering fragments a glimpse into the dawn of humanity when the
first homo sapiens began to form complex utterances. We may be able to at once gain a better
understanding of our early history and the circumstances that lead to the evolution of human language,
but also a sense of solidarity with the rest of the world in which we live.

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*reading from class

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