SPECIAL ISSUE: ANIMAL BEHAVIOUR: A HISTORICAL APPROACH

Animal Behaviour 164 (2020) 233e239

Contents lists available at ScienceDirect

Animal Behaviour
journal homepage: www.elsevier.com/locate/anbehav

Special Issue: Animal Behaviour: A Historical Approach

Animal territoriality, property and access: a collaborative exchange

a r t i c l e i n f o
Territoriality is central to animal behaviourists' understanding of many facets of animal behaviour,
Article history: including resource acquisition, space use behaviour, communication and mating systems. However, the
Received 24 March 2019 term itself, how it is conceptualized and defined, has long been nebulous and contentious. Here, we ask
Initial acceptance 22 July 2019 whether juxtaposing debates about territoriality from animal behaviour with parallel discussions of
Final acceptance 14 November 2019 territoriality from the social sciences can offer a historically and sociologically informed path out of the
Available online 14 January 2020 conceptual gridlock in which animal territoriality has been located for decades. We delineate two key
MS. number: SI-19-00217R problems with territoriality identified in the animal behaviour literature: First, that it focuses on how
animals are expected to behave rather than how they actually behave and, second, that it assumes rather
Keywords: than demonstrates the function of, and specific relationships among, individuals. We then link these
access
problems to social scientists' theorizations of the difference between property and access: whereas
animal
property is focused on how people are expected to behave under juridicalelegal rules governing resource
geography
movement
use, access focuses on a wide array of means by which people actually access resources. We thus argue
property that longstanding problems with animal territoriality have arisen due to implicitly embedded notions of
resource property and ownership. Our juxtaposition raises two further problems with territoriality: first, it
space use unwarrantedly serves to attribute authority to individuals described as territory ‘owners’ and casts others
territoriality (‘intruders’, ‘sneakers’) as transgressors and, second, conceiving of ownership is unfeasible in animal
societies lacking the particular juridicalelegal institutions that establish and enforce property rights.
Instead, we advocate for an access-based approach that will obviate these problems. Ultimately, we argue
that the theory of access, as developed in social science literatures on spatial and relational resource use,
will allow for a fuller and more nuanced understanding of variation in animal behaviour than that
afforded by current dominant notions of territoriality.
© 2019 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Territoriality is a powerful framework in animal behaviour, (1979), Kaufmann (1983), Maher and Lott (1995), Stamps (1994),
motivating and lending context to research on myriad aspects of Bush and Simberloff (2018) and Kamath and Losos (2017, 2018a)
animals’ biology, including resource acquisition, space use behav- have all grappled with similar questions of how to conceive of,
iour, communication and mating systems (Noble, 1939; Carpenter, define and demonstrate territoriality in animals even as empirical
1958; Orians, 1969; Emlen & Oring, 1977; Maher & Lott, 1995, research based on territoriality has continued unabated. In this
2000). In spite of, or perhaps because of, this widespread utility, paper, we present a historically and sociologically informed path
territoriality is challenging to define. In fact, from a historical out of the conceptual gridlock underlying research on animal
perspective, the concept of territoriality has been problematic in territoriality. Specifically, in addressing some of the history of de-
part because animal behaviourists have disagreed about territori- bates on territoriality, (1) we discuss previous definitions of terri-
ality for decades; for example, Emlen (1958), Waser and Wiley toriality (Kaufmann, 1983; Maher & Lott, 1995) to illustrate that
animal territoriality has long been both nebulous and contentious.
(2) We delineate two central problems with territoriality: first, it
* Correspondence: A. Kamath, 468 Donner Lab, University of California, Berkeley,
focuses on how animals are expected to behave rather than how
CA, 94720, U.S.A. they actually behave and, second, it assumes rather than
E-mail address: ambikamath@gmail.com (A. Kamath).

https://doi.org/10.1016/j.anbehav.2019.12.009
0003-3472/© 2019 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
 SPECIAL ISSUE: ANIMAL BEHAVIOUR: A HISTORICAL APPROACH

234 A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239

demonstrates the function of particular behaviours and specific SOME DEFINITIONS OF TERRITORY IN ANIMALS AND HUMANS
relationships among individuals (Emlen, 1958; Kaufmann, 1983;
Stamps, 1994). We describe how these problems with territori- Both animal behaviourists and social scientists have long
ality manifest in studying mating systems, an example we return to considered how to define territory and territoriality. In both areas
throughout the paper. (3) We present potential reasons for, and of scholarship, the mainstream view characterizes territory as a
solutions to, the problems of territoriality by juxtaposing literature ‘defended area’, and territoriality as the ‘defence of area’
on animal territoriality with literature on the concepts of property (Kaufmann, 1983; Maher & Lott, 1995; Sack, 1983). These defini-
and access from the fields of critical social theory, political theory tions appear straightforward, but territoriality is, in fact, far from
and geography (Blomley, 2016; Ribot & Peluso, 2003; Rose, 1985, simple.
pp. 73e88; Sack, 1983). Specifically, we argue that animal behav- In a review of how territory is defined in vertebrates, Maher and
iourists have implicitly and explicitly incorporated aspects of Lott (1995) found 48 different definitions of the term, most of these
property and ownership from human societies into animal terri- implicit; only 12% of papers in their sample included an explicit
tory, a conflation that is unwarranted and problematic. We examine definition. The three most common conceptual definitions were
how social scientists have grappled with similar problems of ‘(1) defended area, (2) exclusive area and (3) site-specific domi-
conflating property and resource use, and draw from their theori- nance’ (Maher & Lott, 1995, p. 1583). The authors note that whether
zations to suggest that instead of enrolling a theory of property to or not a species could be classified as territorial depended on the
better define animal territoriality (Strassmann & Queller, 2014), conceptual definition employed; for instance, in a low-density
animal behaviourists might clarify and strengthen studies of animal population, individuals in a species may effectively occupy exclu-
territoriality by moving towards a theory of access (Ribot & Peluso, sive areas without defending them and would thus be territorial
2003). under the second, but not the first, definition.
Our overarching motivation in writing this paper is to recognize Moreover, ‘territoriality is a hypothetical construct, and its only
that, notwithstanding scientists' aspirations towards objectivity, reality lies in the operational definition’ (Maher & Lott, 1995, p.
scientific practices are embedded within a cultural and social 1588). The operationalization of these conceptual definitions can
milieu (Haraway, 1989). Research in animal behaviour is especially reveal further nuances and complexities. For example, attempting
ripe for sociocultural analysis because of the similarities between to quantify territory using the conceptual definition of ‘defended
the concepts being studied and the lives of the people studying area’ raises the following questions. For how long does a particular
them (Fausto-Sterling, 1997; Lawton, Garstka, & Hanks, 1997; Tang- area need to be defended in order to count as a territory? How
Martínez, 1997; Zuk, 1993). In particular, research on animal terri- much of an individual's time must be spent defending this area, or
toriality is influenced not only by ‘the sort of resources that are what proportion of its home range must the area comprise? What
limiting…and the animal's behaviour’ but also by ‘tradition [and] behaviours constitute defence: is it behaviour at the boundaries of
opinion’ (Maher & Lott, 1995, p. 1582), encouraging us to turn to the the territory and, if so, how are those boundaries determined? Is
social sciences to better understand how sociocultural traditions any aggressive or display behaviour indicative of territoriality?
and opinions may have shaped scientific studies of animal Must the defence of a territory be effective, that is, how successful
territoriality. must an individual be at excluding others from an area? How ani-
mal behaviourists choose to answer these questions, and thereby
construct an operational definition of territory in the context of a
METHODOLOGY particular study, depends upon the taxon and research question
being addressed, as well as logistical considerations. This leads to a
This paper is the outcome of a collaboration between a behav- wide variety of phenomena falling under the umbrella of territo-
ioural ecologist (A.K.) and an interdisciplinary social theorist riality, making comparisons across studies, taxa and contexts
(A.B.W.), wherein we initially exchanged and discussed readings challenging (Linklater, 2000). Maher and Lott (1995, p. 1589) thus
from each of our areas of scholarship to find shared language and exhorted animal behaviourists to be ‘consistent, or at least explicit,
ideas on the concepts of territoriality (animal behaviour: in their terminology’.
Kaufmann, 1983, Strassman & Queller, 2014; sociology: Ribot & In response to these definitions focused on an area as the object
Peluso, 2003). In response to questions arising during our initial of defence and exclusion, both natural and social scientists have
conversation, we included further literature from the social sci- advocated for more nuanced definitions that instead focus on how
ences (critical social theory, political theory and geography; Sack, territory is socially constructed through interactions among in-
1983; Rose, 1985, pp. 73e88; Blomley, 2016) in our discussion. dividuals (e.g. ‘site-specific dominance’, the third most common
We also broadened our discussion of animal territoriality to include conceptual definition in Maher & Lott, 1995). Sack (1983, p. 55)
Emlen (1958), Emlen and Oring (1977), Waser and Wiley (1979), defined human territoriality as ‘the attempt to affect, influence, or
Kaufmann (1983), Stamps (1994) and Maher and Lott (1995), six control actions and interactions (of people, things, and relation-
touchstone papers that review and theorize about how animal ships) by asserting and attempting to enforce control over a
behaviourists use the term ‘territoriality.’ These discussions formed geographic area’, or more simply as a ‘strategy for establishing
the basis for writing this text. Our approach was a less-formal differential access to things and people’. Sack’s (1983) character-
version of a methodology for interdisciplinary research between ization resonates with definitions of animal territory that similarly
biologists and social scientists, detailed in Wesner (2019), that focus on the relationships among individuals from which space use
hinges on mutual engagement (Willey, 2016) via exchanged read- or resource use patterns emerge. Kaufmann (1983, p. 2), for
ings followed by focus group discussions. instance, described territoriality as ‘one form of social dominance’,
Our primary focus in this text is not the behaviour of animals but and defined a territory as ‘a fixed portion of an individual's or
the ideas and practices of scientists studying animal behaviour. We group's range in which it has priority of access to one or more
therefore include quotations from the literature more frequently critical resources over others which have priority elsewhere or at
than in a typical scientific paper, because these quotes constitute another time’ and added, ‘this priority of access must be achieved
the basis on which we discuss how scientists have conceived of through social interaction’ (Kaufmann, 1983, p. 9). This distinction
territory. Further, for clarity and convenience, we use the word between defence or exclusion versus social interactions as a basis
‘animals’ to refer to nonhuman animals only.
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A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239 235

for conceptualizing territory provides a route into understanding behavioural observations under a territorial framework (Avise et al.,
two problems central to animal territoriality. 2002; Boomsma, Kronauer, & Pedersen, 2009; Griffith, Owens, &
Thuman, 2002; Uller & Olsson, 2008). Thus, inferring mating pat-
SOME PROBLEMS WITH TERRITORIALITY REVEALED BY terns through a territorially derived lens frequently fails to predict
CONCEPTUAL EXAMINATION or explain animals’ actual mating behaviour; subsequent attempts
to explain these departures from expected behaviours are posed as
A powerful critique of animal (specifically bird) territoriality was exceptions to the norm derived from territoriality (e.g. Cockburn,
developed by Emlen (1958). He made the case that the definition of Brouwer, Double, Margraf, & van de Pol, 2013).
a territory as a ‘defended area’ constrained research on bird The second problem with territoriality is that it leads animal
behaviour, because it was a misleading assumption to describe behaviourists to assume rather than demonstrate the reasons for
birds as defending areas per se. Rather, his observations suggested particular behaviours, constricting these reasons to a small subset
that while some individual birds were aggressive towards and of all possibilities (Stamps, 1994; Tang-Martínez, 2014). Consider,
perhaps dominant over other individuals in some areas (consistent for example, Maher and Lott’s (1995, p. 1587) description of the
with Kaufmann’s (1983) definition of territory described above), function of communication in relation to territoriality in birds:
there was little ‘justification for including a purely speculative ‘Territory in birds was most commonly defined as defended area…
assumption that the area carries special significance to the bird as Most bird studies involved passerines that advertise their terri-
an object to be defended’ (Emlen, 1958, p. 352). In other words, tories by singing. Singing provides a relatively easy way to quantify
Emlen (1958) argued against automatically inferring the defence of and identify territoriality; in low visibility habitats, measuring [the]
an area (or other resources) from aggressive behaviour, as many amount of singing is easier than counting numbers of intruders
studies of territoriality do. moving through vegetation’. This example shows that alternative
Moreover, Emlen (1958) suggested that most ornithologists' explanations for what information individuals might communicate
conceptions of territory conflate definition with hypothesis. It is when singing (e.g. simply communicating identity), or why they
possible to hypothesize, and then test the hypothesis, that an ani- might behave aggressively (e.g. to establish rank in a dominance
mal defends an area and is thus territorial. However, when defined hierarchy), are rarely considered once a species is believed to be
as ‘the defence of an area’, the concept of territoriality is con- territorial (Emlen, 1958; Linklater, 2011). This means that, under the
structed such that applying the concept to a particular taxon de- paradigm of territoriality, communication is assumed to have
pends upon assuming that the hypothesis of defence is true. As specific territorial function, be it advertising territory ownership
Emlen (1958, p. 352) put it, ‘assumptions are natural features of (Rose, 1985, pp. 73e88) or communicating imminent aggression
hypotheses proposed for experimental testing, but have no place as towards would-be intruders; similarly, aggression is assumed to
integral parts of a functioning concept’. In his view, a useful concept serve the function of defence.
of territory would focus on ‘doings and happenings rather than A particularly salient way in which territoriality prompts animal
objects and entities’. Thus, the first problem with common scien- behaviourists to integrate assumptions into the conceptual foun-
tific usage of territoriality is that it perpetuates a disconnect be- dations of their work while also restricting possible interpretations
tween what is versus what should be: Rather than focusing of animals' behaviours is in the classification of individuals into
attention on how social interactions actually unfold in spatial ways particular roles within a territorial framework (Stamps, 1994).
(what is), territoriality is often operationalized in prescriptive ways Consider, for example, the ubiquitous use of the term ‘intruder’ to
(what should be) that rely on assumptions of how animals relate to describe an individual that moves into an area that has been
and occupy a predetermined type of space. assigned as the territory of another (Kaufmann, 1983; Maher & Lott,
Emlen’s (1958) critique resonates deeply with an observation 1995). Describing that individual as an intruder (as opposed to, say,
made by Sack (1983) in his foundational characterization of human a visitor) leads us to expect an antagonistic interaction between the
territoriality. Sack (1983, p. 59) stated that the notion of territory two individuals. The two individuals' relationship to one another
shifts attention away ‘from the relationship between controller and and to the space they occupy are attributed to the centrality of a
controlled’ and towards the space itself, ‘as when we say ‘it is the possessive figure (the ‘territory owner’) rather than necessarily
law of the land’ or ‘you may not do this here’ [emphasis in original]’. derived from observed behaviours.
This shift is precisely what Emlen (1958) cautioned against. To illustrate how territoriality assumes relationships between
This difference between what is and what should be is especially individuals in the context of mating systems, consider another
salient in considering territoriality in relation to mating systems. In term, ‘sneaker’. Sneakers are males that enter dominant males'
the classic paper linking territory to polygyny, Emlen and Oring territories to covertly attempt to mate with females within the
(1977) stated that whether a species can be polygamous depends territory (or otherwise seek to covertly mate with females thought
on whether multiple mates are economically monopolizable; to be in the ‘possession’ of a dominant male; Taborsky, 1994). An
whether the species is in fact polygamous depends on its ability to animal behaviourist mapping the territories of individual males is
capitalize on this potential for monopolizing mates. They linked especially likely to assign the label of ‘sneaker’ to a comparatively
this potential for, and realization of, polygamy to various ecological, smaller male that enters an area assigned as the territory of another
social and physiological factors. In so doing, they build a unified (usually larger) male, even if the two males do not interact with one
theory of animal mating systems based on a territorial notion of another. Indeed, the term ‘sneaker’ suggests that the first male will
monopolization by defence, not necessarily of a delimited area, but behave surreptitiously to avoid interacting with the second male. A
of mates themselves. Thus, even though most animals are not ‘sneaker’ thus can be seen as not only a transgressor of territorial
classified as territorial, territoriality is deeply embedded in our boundaries but also a perpetrator of infidelity (e.g. Jones, Walker,
understanding of animal mating systems writ large. Within this Kvarnemo, Lindstro € m, & Avise, 2001), reinforcing an underlying
framework, behavioural descriptions of mating systems are assumption that only the dominant, territorial figure has legitimate
essentially predictions of what mating patterns should be if animals claim to a space or the contents of that space.
adhere to rules of defence for the purpose of mate monopolization. When nonterritorial or nondominant individuals are defined
However, the advent of genetic tools to determine paternity primarily in terms of their purported transgressions against
revealed that actual patterns of animal mating (what is) were dominant, territorial males, it can become difficult to interpret their
starkly inconsistent with expected mating patterns derived from behaviour in any other light (Ah-King, 2007). In the coho salmon,
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236 A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239

Oncorhynchus kisutch, where males belong to either the ‘jack’ or within this space. It is also presumed that these rights attach to an
‘hooknose’ morphs, the mating strategy of the smaller jack males individual owner who therefore is assumed to command all the
has traditionally been described as ‘sneaking’ or ‘parasitic’ and their resources within this designated space. The owner is also assumed
continued existence described as ‘problematical’ (Gross, 1985, p. to have the right to govern the access of others…to this space,
47). As is common in species with comparable male phenotypic allowing conditional access or denying it entirely’. Yet social sci-
variation, animal behaviourists had assumed that dominant hook- entists have long documented that juridicalelegal rights, or even
nose males were females' preferred mates. However, in a radical informal social agreements, do not in fact determine how humans
reframing of coho salmon mating strategies, Watters (2005) hy- benefit from objects, resources, spaces or relationships. A property
pothesized that females in fact prefer jack males and cooperate in framework, for example, does not best enable social scientists to
mating with them, whereas less-preferred hooknose males coerce understand how communities collectively manage commons
females into mating. In this case, Watters (2005) opted not to (Ostrom, 1990), prioritize cultural norms of resource sharing over
delimit female individuals as objects of defensive territoriality, nor state-sanctioned legal titles (Walker & Peters, 2001) or whether the
assume the centrality of a dominant ‘territorial’ male individual. named property-holder on a land title is actually the same person
Rather, he tested a hypothesis that opened a broader range of farming the material plot (Peters, 2010). Theories of access, on the
possibilities for how social relations between individuals are spa- other hand, enable researchers to turn to the informal agreements,
tialized, thus contributing novel findings to the study of his system. the grounded practices and the actual use or occupation of different
Watters’s (2005) observations of coho salmon behaviours are spaces to understand how, when and where different individuals
consistent with his hypothesis: females spent more time preparing benefit from resources across time and space.
their nests and spawned for longer in the presence of jacks, and in We suggest that these parallel arguments in the natural and
situations where a jack was chased away from a nest by a hooknose, social sciencesdabout what should be and what is, and the undue
the female spawned soon after the jack's return. focus on possessive functions and interactions in both animal ter-
ritory and human propertydare more than just similar. Rather, we
PROPERTY AND ACCESS IN HUMAN AND ANIMAL CONTEXTS contend that notions of animal territoriality have incorporated
aspects of classically defined property (see Blomley, 2013 for more
Animal Territory as Property? capacious cultural engagements with the idea of ‘property’), and
that these incorporations contribute to the problems with territo-
Just as animal behaviourists have contended with a disconnect riality described above. We show that by paying close attention to
between assumptions about animal behaviour borne of sociocul- social scientists' differentiation of property and access, animal
turally imbued concepts (what should be) and animal behaviours behaviourists will be better able to disambiguate and resolve the
actually observed (what is), so, too, have social scientists grappled most challenging confusions of territoriality.
with the distinction between the concepts of property (what should Burt (1943), in a key early paper on territoriality that has been
be) and access (what is). This theoretical conversation among social cited over 2500 times and represents a foundational paper in the
scientists can illuminate, even in the study of animals, the limita- study of animal territoriality, overtly conflated animal territory and
tions of conceiving territory as a fixed area that is defended by human property. The paper opens with the following conceptual
individual(s) and bounded by rights or rightful relations (property). assertion about territoriality:
New analytical possibilities emerge when animal behaviours are
The behavioristic trait manifested by a display of property own-
instead conceived of through a rubric of access to other individuals,
ershipda defense of certain positions or thingsdreaches its high-
spaces and resources.
est development in the human species. Man considers it his
Ribot and Peluso (2003, p. 153) first defined access ‘as the ability
inherent right to own property either as an individual or as a
to benefit from thingsdincluding material objects, persons, in-
member of society or both. Further, he is ever ready to protect that
stitutions, and symbols. By focusing on ability rather than rights as
property against aggressors, even to the extent at times of sacri-
in property theory, this formulation brings attention to a wider
ficing his own life if necessary. That this behavioristic pattern is not
range of social relationships that can constrain or enable people to
peculiar to man, but is a fundamental characteristic of animals in
benefit from resources without focusing on property relations
general, has been shown for diverse animal groups (Burt, 1943, p.
alone…The concept of access…aims to facilitate grounded analyses
346).
of who actually benefits from things and through what processes
they are able to do so [emphasis in original]’. Thus, the theoretical
framework of access insists that a researcher investigate processes, Two subsequent papers, Stamps (1994) and Strassman and
relationships and behaviours that determine how resources are Queller (2014), continued to centre notions of property and
used. The call for social scientists to focus on how people interact to ownership in discussions of territoriality, and thereby highlight the
benefit from resources is very similar to calls by Emlen (1958) and problems of animal territoriality that we describe above. In a tour-
Kaufmann (1983), described above, to focus on ‘doings and hap- de-force critique of the many untested assumptions of animal
penings’ rather than ‘objects and entities’ in the study of animal territoriality, Stamps (1994, p. 176) described the ‘economic
behaviour. approach to the problem of territory function’, which focuses on
Social science scholars have turned to the theory of access in costs and benefits of territory defence and thus readily lends itself
response to the limitations of property frameworks for really un- to a framework of ownership. She suggested that this economic
derstanding how people benefit from, or are excluded from, ma- approach allowed for generalized discussions of territoriality across
terial and social resources. Property, they have demonstrated, many different taxa but also that it ‘led to a reduction in the number
maintains a focus on the objects or spaces that are owned and, and type of environmental and social factors thought to affect the
specifically, on the formal rules surrounding who is and is not benefits or costs of territorial behavior’. In particular, ‘the ascen-
allowed to use the owned entities. These rules are expressed in dency of the economic approach…affected assumptions about the
language on property through notions of the owners' rights. effects of conspecifics on one another’ (Stamps, 1994, p. 178). Earlier
Consider, as an illustration, Blomley’s (2016, p. 594) description of animal behaviourists ‘were willing to entertain the possibility that
land as property in human society: ‘the rights of the owner (to use, conspecifics might have a positive as well as a negative effect on
occupy, alienate, and so on) apply uniformly across and exclusively one another in territorial species’. These positive effects could
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A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239 237

include the formation of social groups that aid in collective defence, communication, resource acquisition, social interactions, survival
synchronized reproduction and the attraction of mates. But ‘with and reproduction.
the advent of the economic approach, the emphasis shifted to the First, notions of property embedded within the concept of ani-
competitive effects of conspecifics on one another. Neighbours, mal territoriality serve to naturalize both authority and discipline.
floaters, intruders etc., all entered the equation in the same place, as In other words, rather than authority and discipline being recog-
factors contributing to the cost of territorial defense’. Stamps (1994) nized as power-laden social relationships emerging from human
seemed to imply, but did not state outright, that territoriality's societies, they are envisioned as innate qualities of animal behav-
problem of assuming and constricting function and interactions iour emerging directly from ‘nature's order’ (Haraway, 1991). Think
arises because of its incorporation of economic notions of costs and again of ‘intruders’: it seems clear that when we think of ‘intruders’
benefits. While a straightforward focus on costs and benefits does as crossing the boundaries of another animal's territory, then we
not necessarily preclude consideration of positive interactions, may, however unintentionally, conceive of the intruder as trans-
Stamps’s (1994) critique resonates with an ownership-based un- gressing the rights of a property owner. The term ‘intruder’ evokes
derstanding of social interactions that prompts disproportionate the idea of trespass, or the ‘act of unlawful entry upon land in an-
attention to antagonistic interactions motivated by, and costs other's possession’, an act that, in many current conceptions of
incurred due to, the maintenance or transgression of possession. property law ‘is actionable even though no actual damage is done:
The lens of property, along with that of privatization, has been the mere act of boundary transgression is sufficient’ (Blomley, 2016,
applied explicitly to animal territories by Strassman and Queller p. 597). In this way, the intruder's actions take on a sense of being
(2014, p. 306), who suggested that territory ‘is perhaps the best- ‘unlawful’, in opposition to the actions of a territory-holder which
recognized form of privatization’ in animals, and ‘can be defined are configured as the ‘norm’. Similarly, ‘sneaker’ males are
as property in the form of an area that an organism controls, uses, conceived of as subverting the rules surrounding more dominant
and defends’. This explication is consistent with Emlen and Oring’s males' naturalized possession of females. As we saw above in the
(1977) sense of territory as a means for monopolization. By making example of coho salmon, framings of animal behaviour that focus
these linkages between the concepts of animal territory and human on possessive maleefemale interactions potentially led to animal
property explicit, and by considering this connection obvious even behaviourists ignoring, for decades, the possibility that females
as they acknowledge its limits, Strassman and Queller (2014) might benefit more from mating with the subversive ‘sneaker’ male
demonstrate both the prevalence of and the problems with than with dominant males whose actions are considered the norm
property-based notions of animal territoriality. (Watters, 2005). Such framings project onto animals the notion of
In particular, their discussion of mating systems clearly dem- an individual's inherent right to exclude another from a particular
onstrates how territory shifts the discourse surrounding animal space or particular interactions: the sociocultural valuation of pri-
behaviour towards what should be and away from what is. As vate property is naturalized. A conception of territoriality that
Strassman and Queller (2014, p. 306) put it, when describing how implicitly incorporates notions of property can lead animal
territorial male birds' behaviour is inferred to serve the purpose of behaviourists to interpret animal behaviours as obviously indica-
‘privatizing’ reproductive access to females, ‘such privatization is tive of ownership, without necessarily considering alternative in-
imperfect, as shown for example, by the often high frequencies of terpretations and frameworks that do not hinge on ownership.
young in the nests that are not the progeny of the attending male… The non-neutral subversive and transgressive connotations of
a large part of the challenge for territorial males is that they are terms such as ‘intruder’ or ‘sneaker’ are rarely explicit and probably
attempting to privatize a living conspecific that may have con- not intended, but they are nevertheless consequential. As Gowaty
flicting interests [emphasis added]’. In this context, territoriality (1982) illustrated through her analysis of the term ‘rape’ in socio-
has served to reinforce the notion that individuals take possession biology, using words that have emotional resonance in human
of other individuals (and specifically, that males take possession of discourse as technical terms in research on animals comes with
females). Moreover, the discrepancy between expected and multiple downsides, including imprecision, the introduction of bias
observed mating patterns is seen by Strassman and Queller (2014) and unwarranted sensationalism (see also Haraway, 1989, Mesnick,
as animals executing territoriality imperfectly, and not as a short- 1997, Lawton et al., 1997, Tang-Martínez, 1997, and Elgar, Jones, &
coming of territoriality itself as a means of understanding animal McNamara, 2013 for discussions of the consequences of anthro-
behaviour. In other words, attention remains focused on what an- pomorphic terminology in animal behaviour).
imals are expected to be doing under the rules of territory and how A second important problem with incorporating notions of
they are or are not conforming to these expectations, at the cost of property into animal territory is that the existence of property,
developing novel nonterritorial frameworks to understand how which is often conceptualized as a ‘bundle of rights’ (Maine, 1917),
animals actually behave (e.g. Kamath & Losos, 2018b; Kokko & depends upon the existence of historically specific juridicalelegal
Mappes, 2013). Exactly as Emlen (1958) warned, territoriality institutions that enshrine rights and impose sanctions when
leads ‘the study of natural phenomena [to be] subordinated to the rights are transgressed (Blomley, 2016; Rose, 1994). Yet when
study of intellectual creations’. studying the spatial elements of animal behaviour and communi-
cation, scientists do not purport to identify or examine animals'
Problems Revealed by the Parallels Between Animal Territory and formal institutions, mechanisms of government or legal enforce-
Human Property ments and protections of rights. Strassman and Queller (2014, p.
308) acknowledged that ‘nonhumans do not generally have the
Pondering the parallels between animal territory and classic higher-level institutions and each individual must therefore rely on
notions of property in humans reveals two other potential prob- itself or its kin group’. However, they maintain that property rights
lems that may have contributed to the challenge of conceptualizing, in animals exist none the less, suggesting that ‘although only
defining and effectively using territoriality in the study of animals. humans have higher-level social institutions that protect property,
We pose these problems here as starting points from which animal other organisms may sometimes have evolved recognition of
behaviourists could embark on further critical inquiry, asking ownership and hesitation to challenge it’. Strassman and Queller
whether and how these problems have influenced the ways in (2014, p. 307) proposed that ‘physical force, or the threat of it, is
which animal behaviourists construct research questions and perhaps the primary method of privatization’, and therefore of
interpret data on animals’ space use, movement behaviour, property maintenance in animals. This reveals the problems with
 SPECIAL ISSUE: ANIMAL BEHAVIOUR: A HISTORICAL APPROACH

238 A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239

embedded notions of property in territory: while the threat of force could be integrated with much of the existing literature on animal
could be a (potentially costly) strategy for maintaining access to territoriality as long as animal behaviourists are concurrently
resources (Ribot & Peluso, 2003), on its own it is insufficient for careful to examine and clarify the ways in which implicit notions of
establishing, enshrining and enforcing rights. The example cited by property have influenced research questions and hypotheses. An
Strassman and Queller (2014) to illustrate putative ownership is access-based lens could also readily integrate research on animal
that territory ‘owners’ fight more readily than ‘intruders.’ But such contests, which focuses on individual interactions in the light of
behaviour is explicable without invoking ownership, simply by access to resources (Arnott & Elwood, 2009) but may also include
considering access. For example, an individual that is familiar with embedded notions of ownership (e.g. Hammerstein, 1981).
an area may benefit more from remaining in it than would an un- Existing research on mating systems can also be incorporated
familiar individual, and thus may be more inclined to fight for ac- into an access-based framework, which will allow animal behav-
cess to its resources. The theory of access (Ribot & Peluso, 2003) iourists to be unsurprised by natural variation in mating patterns
offers animal behaviourists a way forward for understanding ani- and will facilitate an important shift away from describing certain
mal behaviour that is not constrained by notions of property. mating strategies, such as ‘sneaker’ or ‘satellite’ strategies, as al-
ternatives to a territorial or dominant norm. More radical reconfi-
gurations of mating systems theory, such as Watters's (2005)
A Mode of Animal Behaviour Analysis Informed by Theory of Access
cooperator and coercer model, or Kokko and Mappes’s (2013)
encounter-based approach to understanding female multiple
According to the theory of access (Ribot & Peluso, 2003),
mating as a null expectation and not an anomaly demanding
different individuals and groups are understood to benefit from
explanation, fit within an access-based approach as well.
access to resources in different ways for different reasons; property
ownership is conceived of as one of many avenues or strategies for
accessing resources, as opposed to being the fundamental deter- CONCLUSION
minant. Thus, when viewed through an access-based lens, animal
behaviours are no longer coupled a priori with legal or moral ‘Territoriality, after all, is only an abstraction that describes more or
connotations. For example, animals need no longer be categorized less accurately a certain range of behaviours that occurs often
as territory ‘owners,’ which are implicitly ascribed property rights, enough to merit a name’. (Kaufmann, 1983, p. 10).
or as ‘intruders’ or ‘sneakers’ that transgress those rights. The
notion of rights would only be pertinent in those species that are
Throughout the history of research on territoriality, animal
demonstrated to have juridicalelegal institutions capable of
behaviourists have contended that territoriality is an accurate-
enforcing rights: access allows for but does not demand ownership
enough abstraction for understanding space use, resource use
or rights-based exclusion. Thus, the lens of access immediately
and mating patterns but can be resistant to acknowledging its
addresses the two problems of territoriality that this paper has
limitations as an explanatory framework. Concurrently, periodic
illuminated through the juxtaposition of the natural and social
critiques have not yielded a shift away from the hegemonic para-
sciences literatures on territoriality.
digm of territoriality, possibly because these critiques live squarely
Shifting away from implicit linkages to property towards an
within the fields of animal behaviour and behavioural ecology and
explicit framework of access also allows animal behaviourists to
are thus seen as equivalently valid alternatives to a conception of
address the two problems of territoriality identified from the ani-
animal behaviour that centres possession and ownership. By taking
mal behaviour literature. First, the theory of access is designed to
past critiques of animal territoriality seriously and juxtaposing
focus on what is and not what should be. As Ribot and Peluso (2003,
these critiques with literature from the social sciences, we have
p. 154) explained by quoting Neale (1998, p. 48, italics in original), a
attempted to bring about such a paradigm shift. Specifically, we
theory of access ‘retains an empirical “… focus on the issues of who
argue that many past and current conceptions of animal territori-
does (and who does not) get to use what, in what ways, and when
ality frequently contain deeply embedded but fundamentally
(that is, in what circumstances)”’. Second, the theory of access is
incoherent notions of property, ownership and authority. We sug-
expansive and therefore avoids constraining the possible functions
gest that spatial analyses of social interactions and resource use
of behaviours and relationships among individuals: ‘access is about
informed by social scientists’ conceptions of access will allow us to
all possible means by which a person is able to benefit from things’
reframe how we conceive of animal behaviour in a fuller, more
(Ribot & Peluso, 2003, p. 156) and ‘access analysis is, thus, the
nuanced and more complex way. We contend that comparable
process of identifying and mapping the mechanisms by which ac-
interdisciplinary examinations that meld the natural and social
cess is gained, maintained, and controlled’ (Ribot & Peluso, 2003, p.
sciences should be a critical part of evaluating whether and how to
160).
develop frameworks for understanding any dimension of animal
An access-based approach to animal behaviour already has
behaviour.
parallels in many conceptions of animal territoriality: see
Kaufmann's (1983) definition of territory above, which focuses on
socially determined access, or Waser and Wiley’s (1979, p. 173) Acknowledgments
proposal ‘to abandon the search for a unitary definition of territo-
riality and any simple dichotomy between territorial and non- We thank Todd Freeburg for pointing us to papers in the animal
territorial species’ and instead ‘address the variation among species behaviour literature that sparked this endeavour and Zuleyma
in relationships of aggression, isolation, and activity fields, first by Tang-Martínez for encouraging and facilitating the publication of
considering the behavioural mechanisms that control these re- this paper, as well as for her insightful comments on the manu-
lationships…then by considering the selection pressures that can script. For feedback on this paper and the ideas herein, we thank
explain their evolution’. But these parallels have thus far been Max Lambert, Ignacio Escalante, Trinity Walls, Maggie Raboin,
implicit. Without explicitly disambiguating access from property in Malcolm Rosenthal, Damian Elias, Erin Brandt, Yoel Stuart, Nick
this context, conceptions of animal territoriality will remain Herrmann and two anonymous referees. A.K. is supported by the
confusing and challenging. An explicitly access-based approach Miller Institute for Basic Research in Science.
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A. Kamath, A. B. Wesner / Animal Behaviour 164 (2020) 233e239 239

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