Biol. Rev. (2019), 94, pp. 1430–1442.

1430
doi: 10.1111/brv.12509

All by myself? Meta-analysis of animal

contests shows stronger support for self than
for mutual assessment models
Nelson S. Pinto1 , Alexandre V. Palaoro2 and Paulo E. C. Peixoto3,∗
1
Graduate Program in Ecology, Universidade Federal da Bahia, Salvador, BA 40110-909, Brazil
2
LAGE do Departamento de Ecologia, Universidade de São Paulo, São Paulo, SP 05508-090, Brazil
3
Instituto de Ciências Biológicas, Departamento de Biologia Geral, Universidade Federal de Minas Gerais, Belo Horizonte, MG 31270-901,
Brazil

ABSTRACT

Since the 1970’s, models based on evolutionary game theory, such as war of attrition (WOA), energetic war of attrition
(E-WOA), cumulative assessment model (CAM) and sequential assessment model (SAM), have been widely applied to
understand how animals settle contests. Despite the important theoretical advances provided by these models, empirical
evidence indicates that rules adopted by animals to settle contests vary among species. This stimulated recent discussions
about the generality and applicability of models of contest. A meta-analysis may be helpful to answer questions such
as: (i) is there a common contest rule to settle contests; (ii) do contest characteristics, such as the occurrence of physical
contact during the fight, influence the use of specific contest rules; and (iii) is there a phylogenetic signal behind contest
rules? To answer these questions, we gathered information on the relationship between contest duration and traits
linked to contestants’ resource holding potential (RHP) for randomly paired rivals and RHP-matched rivals. We also
gathered behavioural data about contest escalation and RHP asymmetry. In contests between randomly paired rivals,
we found a positive relationship between contest duration and loser RHP but did not find any pattern for winners. We
also found a low phylogenetic signal and a similar response for species that fight with and without physical contact. In
RHP-matched rivals, we found a positive relationship between contest duration and the mean RHP of the pair. Finally,
we found a negative relation between contest escalation and RHP asymmetry, even though it was more variable than
the other results. Our results thus indicate that rivals settle contests following the rules predicted by WOA and E-WOA
in most species. However, we also found inconsistencies between the behaviours exhibited during contests and the
assumptions of WOA models in most species. We discuss additional (and relatively untested) theoretical possibilities that
may be explored to resolve the existing inconsistencies.

Key words: war of attrition, sequential assessment model, cumulative assessment model, meta-regression, resource-holding
potential, territoriality, agonistic interaction.

CONTENTS
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1431
II. Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1434
(1) Eligibility criteria and information sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1434
(2) Search, study selection and data collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1434
(3) Data items and summary measures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1435
(4) Data analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1435
(a) Relationship between contest duration and traits linked to RHP in randomly paired rivals . . . . . . . 1435
(b) Relationship between contest duration and mean trait values in closely matched rivals . . . . . . . . . . 1435

* Address for correspondence (Tel: +55 (31) 99919-0958; Fax: +55 031 3409-2567; E-mail: pauloenrique@gmail.com).

Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society

Meta-analysis of contest resolution rules 1431

(c) Relationship between contest escalation and asymmetry in traits linked to RHP in randomly paired
rivals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1435
(5) Risk of bias across studies and synthesis of results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1436
III. Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1436
(1) Relationship between contest duration and traits linked to RHP in randomly paired rivals . . . . . . . . . . 1436
(a) Analyses of the data set with all variables related to RHP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1436
(b) Analyses of the data set with the best predictors of RHP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1437
(2) Relationship between contest duration and mean trait values in RHP-matched rivals . . . . . . . . . . . . . . . 1437
(3) Relationship between contest escalation and RHP asymmetry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1437
IV. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1438
V. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1439
VI. Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1440
VII. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1440
VIII. Supporting Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1442

I. INTRODUCTION
Animal contests over limited resources are ubiquitous in
nature (Hardy & Briffa, 2013). Such contests are frequently
initiated when an intruder approaches a resource that is
defended by another individual, and end when one of the
individuals flees. Thus, it is fairly easy to determine when
contests begin or end. However, what happens between
the beginning and ending of contests, namely the contest
dynamics, is highly variable among species. For example,
animal contests may consist of stereotyped behaviours
with distinct behavioural phases (Green & Patek, 2018),
endurance displays without physical contact (Peixoto &
Benson, 2011) or even disputes characterized by continuous
pushing and biting between rivals (Painting & Holwell, 2014;
Palaoro et al., 2014). Interestingly, despite these frequent tugs
of war, contests rarely involve the death of an individual.
Rivals typically terminate the contest before any damage is
done (Lane & Briffa, 2017; Palaoro & Briffa, 2017). Since
individuals are contesting valuable resources and exposing
themselves to damage, the rarity of death indicates that
individuals adopt some sort of criterion to decide when
to give up a contest before reaching extreme costs. The
decision of when to give up on a contest has been modelled
using evolutionary game theory since the 1970’s (Briffa
& Elwood, 2009). However, despite the large number of
models (Mesterton-Gibbons, Marden & Dugatkin, 1996;
Briffa & Sneddon, 2010; Kokko, 2013; Mesterton-Gibbons &
Heap, 2014), only four of them have been thoroughly tested
empirically: war of attrition (WOA) (Mesterton-Gibbons
et al., 1996), energetic war of attrition (E-WOA) (Payne &
Pagel, 1996), cumulative assessment model (CAM) (Payne,
1998) and sequential assessment model (SAM) (Enquist &
Leimar, 1983).
One of the central concepts of all assessment models (i.e.
WOA, E-WOA, CAM and SAM) is the resource holding
potential (or resource holding power, RHP sensu Parker,
1974). The RHP is related to the persistence capacity during
a contest and/or the ability to inflict costs on rivals (Vieira
& Peixoto, 2013), and is often correlated with morphological
and physiological attributes of the individual (Hardy &
Briffa, 2013; Palaoro & Briffa, 2017). The four assessment
models assume that assymetries in RHP are important in
determining who wins the contest (Briffa & Hardy, 2013).
However, they also assume differences on how animals gather
information about RHP during contests.
According to WOA, E-WOA and CAM, individuals follow
a self-assessment strategy that assumes that individuals do not
obtain information about the rival’s RHP (Arnott & Elwood,
2009). Consequently, each individual should give up when
its own cost accrual reaches a threshold determined by its
own RHP (Arnott & Elwood, 2009). According to WOA, the
main cost should be the time spent in the interaction, while
according to E-WOA, energy expenditure should be the main
cost. However, for most situations WOA and E-WOA are
empirically indistinguishable (e.g. when individual size is used
as a proxy of RHP), and for this reason, we will refer to both
models as (E)WOA. The last self-assessment strategy, CAM,
assumes that costs also accrue from injuries caused by the
rival during physical contact (Payne, 1998). Hence, the key
difference between (E)WOA and CAM is that, in (E)WOA
cost accrual is unaffected by the rival’s actions, while in CAM
injuries inflicted by rivals affect individual persistence in the
contest (Payne, 1998). For this reason, according to (E)WOA,
the loser’s persistence should be strongly correlated with its
own RHP (such as size or energy reserves) and be unaffected
by the winners’ actions. Under CAM, however, the loser’s
persistence is again strongly correlated with its own RHP,
but also negatively correlated to the winner’s RHP because
stronger rivals cause losers to reach their cost threshold faster
and, therefore, persist for less time in a fight.
In contrast to the self-assessment strategies, SAM assumes
that rivals are capable of assessing their rivals’ RHP, making
a decision based on relative RHP, rather than absolute RHP
(Enquist & Leimar, 1983). Thus, each individual decides to
give up when it considers itself to be the weaker rival (and not
when reaching a specific cost threshold). If both individuals
have similar RHPs, they should escalate the contest to
costlier behaviours that provide more accurate information
on the rival’s RHP, until one individual identifies itself
as the weaker rival and gives up. Under SAM then, the
loser’s persistence will depend on the winner’s RHP and

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1432
its own RHP. This entails a positive relationship between
loser persistence and loser RHP because, as the loser’s RHP
increases, it becomes more difficult for the loser to detect
that it is weaker than the winner – increasing the length of
the contest. Furthermore, there is also a negative relation
between persistence and the winner’s RHP because, as the
winner’s RHP increases, the easier it will be for the loser
to detect that it is weaker than the winner. In summary,
theory expects that under (E)WOA contest duration should
be mainly related to traits linked to the losers RHP. Under
CAM and SAM, however, information about winners (SAM)
or damage received from them (CAM) also play a role
in determining when losers should give up. Therefore,
according to CAM and SAM contest duration should be
related to both loser and winner RHP.
Empirical tests of (E)WOA, CAM and SAM have been
developed since 1990 (e.g. Marden & Waage, 1990; Leimar,
Austad & Enquist, 1991). However, detailed evaluations
of these models were carried out only after the analytical
framework proposed by Taylor & Elwood (2003) (e.g. Kelly,
2006; Stuart-Fox, 2006; Peixoto & Benson, 2011). Before this
analytical framework, most studies correlated fight duration
with the difference in trait values between winners and
losers. A negative relationship between fight duration and
the difference in trait values was considered evidence in
support of SAM, and, in fact, many tests based on such
correlations found support for SAM (e.g. Enquist et al., 1990;
Leimar et al., 1991; Hack, 1997). However, Taylor & Elwood
(2003) showed that this approach led to biased conclusions.
When individuals are randomly paired, losers with low RHP
have a high chance of fighting against much stronger winners.
This occurs because there is a large set of the population
with individuals with greater RHP values than losers with
low RHP. Losers with high RHP, on the other hand, would
come from a small set of the population that is constituted by
large RHP individuals. To lose, these high-RHP individuals
would have fought against individuals with even higher
RHP. Since there are few individuals at this extreme of the
population, losers with high RHP have a high probability of
fighting against similar-RHP rivals. In this situation, even
if individuals follow a self-assessment strategy, negative
correlations should be obtained when regressing contest
duration against trait differences between winners and losers.
To avoid finding negative correlations between contest
duration and trait differences under self-assessment strate-
gies, Taylor & Elwood (2003) suggested that assessment
models should be tested using linear regressions between
contest duration and winner and loser RHP separately
(Taylor & Elwood, 2003; but see Briffa & Elwood, 2009).
According to their framework, under (E)WOA, there should
be a positive relationship between contest duration and
traits linked to RHP for losers and no relationship (or
a weak positive one) between contest duration and traits
linked to RHP for winners. According to CAM and SAM,
on the other hand, there should be a positive relationship
between contest duration and traits linked to RHP for losers
and a negative relationship of similar magnitude between
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
Nelson S. Pinto and others
the same variables for winners (Fig. 1). It is important to
note, however, that for SAM, if individuals have better
information about their own fighting ability than the fighting
ability of their rivals, the relationship between fight duration
and individual RHP should be stronger for the losers than
for the winners (Prenter, Elwood & Taylor, 2006).
The distinction between CAM and SAM is more difficult
to make and cannot be accomplished using the correlations
between contest duration and traits linked to RHP (Briffa &
Elwood, 2009). At least two additional sources of information
are needed to distinguish between CAM and SAM. First,
it is necessary to use information about contest duration
and traits linked to RHP for rivals with a high degree of
similarity between these traits (i.e. RHP-matched rivals).
In a hypothetical scenario where it is possible to arrange
contests between RHP-matched rivals across all ranges of
RHPs in the population, different relations between contest
duration and the mean RHP of the pair would emerge
if animals follow CAM or SAM. Since the relative RHP
remains the same, under CAM (and also under (E)WOA),
we expect a positive relation between contest duration and
the mean RHP of the pair, while under SAM, there should
be no relationship between contest duration and the mean
RHP of the pair (Arnott & Elwood, 2009, Fig. 1). Second,
the assessment models were built based on differences in
escalation patterns, and for this reason, it is necessary to
evaluate how individuals are escalating and de-escalating
their contests (Briffa, 2015). According to SAM, the most
escalated interactions (e.g. change from displays to physical
contact during the contest, Painting & Holwell, 2014) should
occur only between individuals with similar RHP (Enquist
& Leimar, 1983). Therefore, if animals behave according
to SAM, there should be a negative relationship between
escalation probability and the difference in trait values for the
pair. However, under CAM, there should be no relationship
between contest escalation and the difference in trait values
between rivals (Payne, 1998, Fig. 1).
Despite methodological advances in how predictions from
contest models are tested, few studies have found unequivocal
support for (E)WOA, CAM or SAM (e.g. Prenter et al.,
2006; Briffa, 2008; Junior & Peixoto, 2013; Schnell et al.,
2015). Most have found partial support for some of the
models, support for more than one model, or patterns that
were not predicted by any model (e.g. Palaoro et al., 2014;
Guillermo-Ferreira et al., 2015). These findings have sparked
discussions about the applicability and generality of the
assessment models (Briffa & Sneddon, 2010; Lane & Briffa,
2017; Palaoro & Briffa, 2017). One largely ignored aspect,
for instance, is that some assessement rules might be more
likely to evolve in some groups than in others. For example,
in species that fight with physical contact, individuals may
be able to cause injuries during the fight. Consequently,
fights settled according to CAM or SAM may be more likely
to evolve in species that fight with physical contact, while
(E)WOA may be more common in species that fight without
physical contact (although SAM may also explain assessment
rules in some species that fight without physical contact – e.g.
Meta-analysis of contest resolution rules 1433

Fig. 1. Legend on next Page.

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1434 Nelson S. Pinto and others

Vilela et al., 2017). Phylogenetic history may also be
important in explaining how individuals within a given clade
decide when to give up. In particular, closely related species
might make contest decisions similarly due to common
ancestry. Hence, a systematic evaluation of the empirical tests
of these models is essential to: (i) show the level of empirical
support for the different assumptions and predictions of the
assessment models; and (ii) determine if most species tend
to fight according to a specific model or whether different
groups of species follow the rules of different models.
Here, we conducted a meta-analysis to evaluate the
empirical support for (E)WOA, CAM and SAM. We first
evaluated the empirical support distinguishing (E)WOA from
CAM and SAM. For this, we investigated the relationship
between contest duration and traits linked to RHP for
winners and losers in randomly matched pairs (Fig. 1).
We then investigated the empirical support for CAM
and SAM by evaluating the relationship between contest
duration and traits linked to RHP for RHP-matched rivals
(Fig. 1). Additionally, we investigated the support for SAM
through the relationship between escalation patterns and
RHP asymmetry (Fig. 1). For all tests we also measured
the phylogenetic signal in the observed patterns and, when
possible, tested if the observed patterns differed between
species that fight with and without physical contact.
II. METHODS
(1) Eligibility criteria and information sources
We followed the PRISMA protocol because it provides a
standardized method for reporting meta-analyses (Moher
et al., 2009). Our eligibility criteria for study inclusion
required that the study presented information on the
relationships between contest duration and morphological,
physiological or behavioural traits linked to RHP for winners
and losers in dyadic contests (Taylor & Elwood, 2003). These
comparisons could involve contests between randomly paired
individuals or individuals systematically paired to minimize
differences in traits linked to RHP (i.e. RHP-matched rivals).
We also included papers that provided information about the
probability of escalation (i.e. change from less costly to costlier
behaviours) during the contest in relation to differences in
traits linked to RHP between rivals. We conducted the
literature search in ISI Web of Knowledge, Scopus and Google
Scholar between July 2014 and January 2019.
(2) Search, study selection and data collection
We performed two distinct types of searches. First, we
conducted a search for articles that provided information
on the relationship between contest duration and traits
linked to RHP for losers and winners using the following
key words: ‘animal contests’, ‘RHP’, ‘resource holding
potential’, ‘contest resolution’, ‘SAM’, ‘sequential assessment
model’, ‘WOA’, ‘war of attrition’, ‘E-WOA’, ‘energetic war
of attrition’, ‘CAM’, ‘cumulative assessment model’ and
‘contest duration’. We performed the search using each key
word separately.
For each selected paper, we first read the title and the
abstract searching for information indicating that the study
investigated contests between pairs of rivals. For studies that
provided this information, we searched the methods for
evidence that the study evaluated contests according to the
protocol of Taylor & Elwood (2003). If the study used this
protocol, we continued to the results to collect information on
the statistics for the relationships between traits linked to RHP
and contest duration for winners and losers (we considered
as a trait linked to RHP any morphological, physiological or
behavioural trait that was stated in the original article as an
attribute that indicated individual RHP). We also recorded
if the study used randomly paired rivals or RHP-matched
rivals and if physical contact occurs during the contests for
each species.
Second, we conducted a search for articles that provided
results regarding relationships between differences in traits
linked to RHP of pairs of rivals and the probability of
escalation during the contest. To perform this search,
we used the following key words separately: ‘animal
contests’, ‘RHP’, ‘resource holding potential’, ‘contest
resolution’, ‘SAM’, ‘sequential assessment model’, ‘CAM’,
‘cumulative assessment model’ and ‘contest escalation’.
Contest escalation may involve escalation both between
(i.e. a progressive use of costlier behaviours) and within

Fig. 1. Summary of the responses expected under different assessment models for the relationships between: (i) contest duration and
traits linked to resource holding potential (RHP) for winners and losers (often tested through linear regressions – top row); (ii) contest
duration and mean RHP of the fighting pair (often tested through linear regressions – middle row); and (iii) escalation patterns (i.e.
changes from less costly to costlier behaviours) and trait difference between rivals (often tested through logistic regressions – bottom
row). For each predicted relationship, we depict the expected pattern in the original articles (top figures in each row) and the
expected pattern in our meta-analyses (bottom figure in each row). The expected pattern in our meta-analysis is based on mean
values calculated from data recorded in the original studies and converted to Fisher’s z values (Zr), which represents the effect size
we used. A mean Zr value of zero (represented by dashed lines) indicates that there is no consistent relationship among species
between contest duration or escalation and winners/losers’ traits. (E)WOA represents the expectation for both War of attrition and
Energetic War of Attrition models, CAM represents the expectations for the Cumulative Assessment Model and SAM represent
the expectation for the Sequential Assessment Model. Randomly paired rivals occurred in experiments in which individuals were
randomly paired to induce contests, while RHP-matched pairs occurred in experiments in which rivals were paired to minimize the
difference between the traits linked to RHP.

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Meta-analysis of contest resolution rules
(increase in the number or frequency of the same behaviour)
phases. However, because SAM assumes escalation between
phases, we only included articles with suitable data to test
this assumption of SAM. In general, escalation between
phases was represented by the transition from non-contact
behaviours to physical contact and grappling.
As in the search for studies involving the relationship
between traits linked to RHP and contest duration, we read
the title and the abstract searching for information indicating
that the study investigated dyadic agonistic interactions. For
studies that provided this information, we proceeded to the
methods section to search for information indicating that the
study measured the relationship between contest escalation
and traits linked to RHP for rivals. We then continued to
the results section to collect information about statistics for
the relationships between the probability of escalation and
differences in values of traits linked to RHP for rivals.
(3) Data items and summary measures
We used Fisher’s z scale (Zr) as a measure of effect size for
the relationship between contest duration and traits linked
to RHP. We also used the same effect size for the relation
between contest escalation and traits linked to RHP. Fisher’s
z is a normalization of the Pearson correlation coefficient
(r) and allows comparisons to be made among studies with
different sample efforts (Borenstein et al., 2009). To obtain Zr
values, we first obtained the Pearson correlation coefficients
and then transformed them to Zr (Borenstein et al., 2009).
When only the coefficient of determination was reported,
we transformed it to r using the square root of the observed
value (Nakagawa & Cuthill, 2007). When only F , t, χ 2
values and degrees of freedom were reported, we performed
transformations to Zr using the package compute.es (Del Re,
2013). When a study provided information in graphs, we
used the webplotdigitizer software (Rohatgi, 2012) to estimate
raw values and then calculated the required information.
When some information was unavailable in a paper, either
explicitly or in graphs, we requested it from the authors. We
retained the signs of all original statistics (indicating positive
or negative relationships) after all the transformations.
(4) Data analyses
(a) Relationship between contest duration and traits linked to RHP in
randomly paired rivals
To evaluate if contest duration is related to traits linked to
RHP of winners and losers, we performed a mixed-model
meta-regression (Koricheva, Gurevitch & Mengersen, 2013).
We performed the meta-regressions for two different data
sets. The first data set contained all effect sizes provided by
each study. However, since some studies measured more
than one trait related to RHP, we scanned the papers for
any analysis or author commentaries stating which was the
most important RHP trait for that study. We then pruned
our first data set and left only the effect sizes related to the
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
1435
most important determinant of RHP. Our second data set,
hence, only included one trait related to RHP per species.
For each data set, we built a model considering the Zr
values as the response variable, and status (winner or loser),
occurrence of physical contact and the interaction between
status and occurrence of physical contact as moderator
explanatory variables. We included study identity and the
type of experiment (laboratory or field) as random variables
to account for variation among multiple effect sizes obtained
for the same study and to account for variation among
studies conducted in laboratory and field conditions. We
also included a variance/covariance matrix based on the
phylogenetic relationships among species as a random
variable to account for variation among groups of related
species (Chamberlain et al., 2012). We weighted each Zr
value by the inverse of the variance of Zr (Borenstein et al.,
2009). Therefore, in these models we tested if the mean Zr
differed between winners and losers for species that fight
with or without physical contact (Fig. 1). We calculated
the significance tests for moderator variables using the QM
statistic (Borenstein et al., 2009). If (E)WOA was the best
model to explain animal contests, we expected that mean
Zr values would be greater than zero for losers, but not for
winners. If SAM or CAM is the best explanation for animal
contests, we expected that mean Zr values would be greater
than zero for losers, but smaller than zero for winners (Fig. 1).
(b) Relationship between contest duration and mean trait values in
closely matched rivals
To test if there was a relationship between contest duration
and mean RHP for RHP-matched rivals, we used a
random-effects meta-analysis (Koricheva et al., 2013). In this
model, we considered Zr values as the response variable and
inserted the inverse of the variance of each Zr value as the
weight – no moderator (explanatory) variables were used. In
this data set, all species had fights that can involve physical
contact. Therefore, for this analysis we only tested if the
mean Zr value differed from zero (Fig. 1). As in the previous
analysis, we included study identity, type of experiment
(laboratory or field) and a variance/covariance matrix based
on the phylogenetic relationships among species as random
variables. Since we did not have moderator explanatory
variables, we tested if the mean Zr value differed from zero
using a z-test (Viechtbauer, 2010). According to CAM,
the mean Zr value should be greater than zero, while
according to SAM this value should not differ from zero
(Fig. 1).
(c) Relationship between contest escalation and asymmetry in traits
linked to RHP in randomly paired rivals
To evaluate if the probability of escalation is related to
differences in traits linked to RHP between rivals, we used
a random effects meta-analysis (Koricheva et al., 2013).
We considered Zr values as the response variable and
inserted the inverse of the variance of each Zr value as
the weight – again no moderators were used. We tested
1436 Nelson S. Pinto and others

if the mean Zr values differed from zero because all Table 1. Heterogeneity I 2 and variance (σ 2 ; V ) measures for
species in this data set also fight with physical contact. the random variables included in each meta-analytical model
As random variables, we included the type of experiment used herein. I 2 values represents the percentage of variability
(laboratory or field) and a variance/covariance matrix based in estimates of effect sizes due to data heterogeneity rather than
on the phylogenetic relationships among species (we did sampling error. I 2 = 25, 50 and 75% indicates low, moderate
not include study identity because only one study provided and high heterogeneity, respectively (Higgins & Thompson,
2002; Higgins et al., 2003)
more than one datum). If rivals fight according to SAM,
we expected the mean Zr value to be smaller than zero
(a) Model 1.1 – randomly matched pairs: duration versus RHP
(Fig. 1). traits
I 2 (95% CI) V (95% CI)
(5) Risk of bias across studies and synthesis of
results Study ID 22.48 (20.56–24.4) 0.05 (0.03–0.09)
Experiment <0.001 (0.0–1.9) <0.001 (0.0–0.02)
To evaluate if there was publication bias in our data set, Phylogeny <0.001 (0.0–1.9) <0.001 (0.0–0.02)
we conducted a modified version of Egger’s test (Egger et al., Total 22.48 (20.56–24.4) —
1997), in which we regressed the residuals of each model
(b) Model 1.2 – randomly matched pairs: duration versus RHP
against the variance of the Zr values (Nakagawa & Santos, traits considering only the most important measure for each
2012). A significant intercept suggests that the publications species
used in our meta-analyses are concentrated toward positive
or negative effect sizes. We estimated the heterogeneity I 2 (95% CI) V (95% CI)
among effect sizes using I 2 and its corresponding 95% Study ID 27.24 (25.32–29.16) 0.06 (0.02–0.12)
confidence intervals for multivariate studies. The I 2 statistic Experiment <0.001 (0.0–1.9) <0.001 (0.0–0.34)
varies between 0 and 100% and can be interpreted as the Phylogeny <0.001 (0.0–1.9) <0.001 (0.0–0.13)
percentage of variability in estimates of effect sizes due to Total 27.24 (25.32–29.16) —
data heterogeneity rather than sampling error (Higgins & (c) Model 2 – RHP-matched rivals: duration versus mean RHP
Thompson, 2002; Higgins et al., 2003). We also calculated
the H 2 metric that estimates the phylogenetic signal in each I 2 (95% CI) V (95% CI)
test (Nakagawa & Santos, 2012). Values of H 2 near 100% Study ID 10.97 (9.05–12.89) 0.02 (0.0–0.09)
indicate a strong phylogenetic signal in the responses we Experiment <0.001 (0.0–1.9) <0.001 (0.0–0.43)
measured. Phylogeny <0.001 (0.0–1.9) <0.001 (0.0–0.10)
We conducted all analyses in R software (R Development Total 10.97 (9.05–12.89) —
Core Team, 2015). We used the metafor package (d) Model 3 – Escalation versus RHP difference including the
(Viechtbauer, 2010) to build and test all models. We used outlier study
the rotl package (Michonneau, Brown & Winter, 2016) to
I 2 (95% CI) V (95% CI)
build a phylogenetic tree with the species present in our
data set [the information to build this tree was obtained Experiment <0.001 (0.0–1.9) <0.001 (0.0–1.03)
from the Tree of Life Project (http://tolweb.org/tree/)]. We Phylogeny 25.83 (23.91–27.75) 0.05 (0.01–0.22)
then used the ape package to convert the topological tree to Total 25.83 (23.91–27.75) —
an ultrametric tree (see online Supporting information, Figs (e) Model 3 – Escalation versus RHP difference excluding one
S1–S3) with simulated branch lengths (Paradis, Claude & outlier study
Strimmer, 2004) in order to include a matrix of phylogenetic
I 2 (95% CI) V (95% CI)
correlations in our models. We present the results as mean
Zr values and 95% confidence intervals. Experiment 8.30 (6.38–10.22) 0.01 (0.0–2.23)
Phylogeny 0.38 (0.0–2.30) <0.001 (0.0–0.07)
Total 8.68 (6.76–10.60) —

III. RESULTS RHP, resource holding potential.

(1) Relationship between contest duration and
traits linked to RHP in randomly paired rivals
(a) Analyses of the data set with all variables related to RHP
For the data set with all variables related to RHP, we
obtained 174 effect sizes (88 for winners and 86 for losers) for
36 species (Table S1 and Fig. S1, Supporting information).
There was no evidence for publication bias (Egger’s test:
estimate = −0.06; 95% CI = −0.15 to 0.02; P = 0.15)
and a low level of heterogeneity (Table 1a). The variance
components and heterogeneity due to type of experiment
and to phylogeny were low; most of the variance was due
to study ID (Table 1a). The phylogenetic signal was also low
(H 2 < 0.001%).
The relationship between contest duration and traits
linked to RHP was not influenced by the interaction
between status and the occurrence of physical contact

Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society

Meta-analysis of contest resolution rules
Fig. 2. Mean Fisher’s z (Zr) values and their 95% confidence intervals for the relationship between contest duration and traits linked
to resource holding potential (RHP) for losers and winners of all contests (with and without physical contact). (A) Results from the
data set with all trait values used to estimate individual RHP in the original articles. (B) Results from the data set including only
the original authors’ best predictor of RHP for each species. A Zr value of zero (dashed line) indicates that contest duration was
unrelated to trait values.
(QM status*physical contact = 0.60; df = 1; P = 0.44) and did
not differ between species that fight with and without
physical contact (QM physical contact = 1.30; df = 1; P = 0.25).
However, the relationship between contest duration and
traits linked to RHP differed between winners and losers
(QM status = 187.5; df = 1; P < 0.001). Contest duration
increased with loser RHP, but there was no relationship
between contest duration and winner RHP (Fig. 2A). These
results thus provide support for the (E)WOA model regardless
of the presence of physical contact.
(b) Analyses of the data set with the best predictors of RHP
For the second data set with only one trait linked to RHP
per species, we obtained 68 effect sizes (34 for winners
and 34 for losers) for 36 species (Table S1, Supporting
information). There was no evidence for publication bias
(Egger’s test: estimate = 0.03; 95% CI = −0.13 to 0.18;
P = 0.74) and a low level of heterogeneity (Table 1b). The
variance components and heterogeneity due to type of
experiment and to phylogeny were low; most of the variance
was due to study ID (Table 1b). The phylogenetic signal was
also low (H 2 < 0.001%).
Similar to the results considering all traits linked to
RHP, the relationship between contest duration and traits
linked to RHP was also not influenced by the interaction
between status and the occurrence of physical contact
(QM status*physical contact = 0.67; df = 1; P = 0.41) and did
not differ between species that fight with and without
physical contact (QM physical contact = 2.72; df = 1; P = 0.10).
We also found a positive relation between contest duration
and loser RHP, while winner RHP was not correlated
to contest duration (QM status = 170.7; df = 1; P < 0.001;
Fig. 2B). Thus, regardless of the number of measured
traits linked to RHP, the data suggest (E)WOA models
as the best explanation for how individuals give up on
contests.
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
1437
(2) Relationship between contest duration and
mean trait values in RHP-matched rivals
We obtained 21 effect sizes from 16 species (Table S2 and
Fig. S2, Supporting information). There was no evidence for
publication bias (Egger’s test: estimate = 0.04; CI = −0.11
to 0.18; P = 0.59) and a low level of heterogeneity (Table 1c).
The variance components and heterogeneity due to type of
experiment and phylogeny were low. Most of the variance
was concentrated on variation among studies (Table 1c). The
phylogenetic signal was also low (H 2 = 0.02%).
The mean effect size was greater than zero (z-score = 4.77;
P < 0.001; Fig. 3), indicating that contest duration increases
with mean RHP of the pair. This result provides support for
the self-assessment strategies-(E)WOA and CAM.
(3) Relationship between contest escalation and
RHP asymmetry
We obtained 20 effect sizes for 20 species (Table S3 and Fig.
S3, Supporting information). There was no evidence for pub-
lication bias (Egger’s test: estimate = −0.04; CI = −0.17 to
0.09; P = 0.55) and a low level of heterogeneity (Table 1d).
The variance component and heterogeneity due to type of
experiment was low; but in this case, most of the variance
was concentrated on phylogeny (H 2 = 99%, Table 1d).
We found that the mean effect size did not differ from
zero (Zr = −0.16; CI = −0.41 to 0.09; z-score = 1.23;
P = 0.22), indicating that there is no relationship between
contest escalation and RHP asymmetry. However, this
result was strongly affected by one study on the crab
Austruca annulipes (Bolton, Backwell & Jennions, 2013).
After removing this study, the variance component and
heterogeneity due to type of experiment increased, while
the variance explained by phylogeny decreased (Table 1e).
We found that the mean effect size was smaller than zero
(z-score = 3.10; P = 0.002; Fig. 4), indicating that for the
remaining 19 species, individuals were less prone to escalate
1438
Fig. 3. Mean Fisher’s z (Zr) value and its 95% confidence
interval for the relationship between contest duration and mean
trait value of the fighting pair for contests in which rivals with
similar resource holding potential (RHP) values were paired
(RHP-matched rivals). A Zr value of zero (dashed line) indicates
that contest duration was unrelated to mean trait values.
Fig. 4. Mean Fisher’s z (Zr) value and its 95% confidence
interval for the relationship between contest escalation (i.e.
change from less costly to costlier behaviours during the contest)
and the difference in the traits related to resource holding
potential (RHP) of the fighting pair (excluding one outlier
study). A Zr value of zero (dashed line) indicates that contest
escalation was unrelated to trait difference between rivals.
as RHP asymmetry increases. The phylogenetic signal was
low, but it was the highest among all analyses (H 2 = 4.4%).
IV. DISCUSSION
In this study, we showed that contest duration increased
with loser RHP, but not with winner RHP when individuals
are randomly matched for RHP (regardless of the number
of traits linked to RHP used in the analyses). This result is
consistent with predictions of the (E)WOA models (Fig. 1) in
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
Nelson S. Pinto and others
which the losers do not gather information about the winners’
RHP – they give up in response to the costs they accrue
based on their own actions, regardless of the occurrence of
physical contact. When considering RHP-matched rivals, we
found that contest duration increased with mean trait values
of the fighting pair, which is consistent with self-assessment
strategies such as (E)WOA and CAM (Fig. 1). Therefore,
analyses based on the framework proposed by Taylor &
Elwood (2003) support self-assessment strategies as the
models that best describe the decision to give up on a contest.
On the other hand, when we consider the predictions made
by the models themselves regarding escalation patterns, the
results were more variable. When we considered all species,
we found no relation between probability of escalation
and RHP asymmetry. However, when we removed an
outlier study, we found a negative relation between the
probability of escalation and RHP asymmetry (Fig. 4) – a
pattern that provides support for SAM. It is important to
note that non-significant results do not necessarily indicate
that the species used in our analyses do not follow any
rule – non-significant results may suggest that species vary
widely in the rules they adopt to settle contests. Hence,
significant results indicate that different species may follow
a similar assessment model which decreases variation. For
this reason, although some species may settle their contests
based on mutual assessment (e.g. Junior & Peixoto, 2013;
Schnell et al., 2015; Green & Patek, 2018), our results for the
relationships between contest duration and traits linked to
RHP suggest that the proportion of species following mutual
assessment is low and, therefore, self-assessment seems to be
the most frequent contest resolution rule adopted by species.
It is surprising that (E)WOA models are the best candidates
to explain contest resolution for most animal species for
two reasons. First, these models were not developed to
explain physical contact – they were developed to explore
bouts of displays between rivals (Bishop & Cannings, 1978).
Second, in (E)WOA, contests do not escalate to different
phases. Individuals only increase the intensity of displays.
Therefore, most studies do not agree with some or most of
the assumptions of (E)WOA. For instance, in the dragonfly
Diastatops obscura, males defend territories in which females
oviposit by performing bouts of aerial displays (Junior &
Peixoto, 2013). However, males can also escalate to physical
contact when they have similar RHP, which violates the
assumptions of (E)WOA. In fact, from the 36 species for
which we gathered information, only two could follow
(E)WOA without violating any assumption (Hamadryas and
Hermeuptychia butterflies).
Perhaps the evidence in support of (E)WOA occurs due to
other factors that are not considered in the original models.
For example, recent studies suggest that damage capacity
(Palaoro & Briffa, 2017) and self-inflicted damage (Lane &
Briffa, 2017) may affect contest dynamics and, consequently,
influence the relationship between contest duration and
winners’ and losers’ RHP. If the importance of damage
varies among species, we may find species in which the
relationship between contest duration and winner RHP is
Meta-analysis of contest resolution rules
negative (e.g. species that bear weapons) and species in
which this relationship is absent (e.g. species that do not bear
weapons). This would explain why isolated studies do not
provide evidence for (E)WOA, while meta-analytic evidence
indicates that (E)WOA explains contest decisions for most
species. Thus, we argue that a self-assessment strategy is
indeed the general rule used to resolve contests, but it is
a more flexible form of (E)WOA. Such flexibility in the
assumptions would come from adding that contests are not
a single behavioural phase, that most species use physical
contact during contests, and that damage might play an
important role in some contests, but not in others. Alternative
experimental approaches not based on correlations between
contest duration and individual traits are essential to clarify
this picture (Arnott & Elwood, 2008, 2009). For example, by
manipulating the type of visual cues exhibited by tethered
rivals in the damselfly Mnesarette pudica, it was possible to show
that, unlike the results involving correlations between fight
duration and individual traits, stronger males perform mutual
assessment while weaker males perform self-assessment
(Guillermo-Ferreira et al., 2015).
Self-assessment was advocated as the most probable
contest resolution rule among animals because it is less
cognitively complex than mutual assessment (Elwood &
Arnott, 2012; but see Fawcett & Mowles, 2013). Since
most species follow self-assessment rules, it could be that
self-assessment represents the basal assessment rule. Based
on this argument, mutual assessment would only be found in
a few closely related groups – a hallmark of a phylogenetic
signal. However, we found no evidence of phylogenetic
signal in assessment models, indicating that both self- and
mutual assessment occur scattered across the phylogeny
regardless of the phylogenetic relationships among species.
One possible explanation for this pattern is the lack of
studies in closely related species. We found only two studies
for congeneric fiddler crabs (Austruca spp.), one study for
three species of stalk-eyed flies (Cyrtodiopsis spp.), and another
study for two species of crayfish (Parastacus spp.). All other
studies were from distantly related species, which could
decrease the importance of phylogenetic history in our data.
To highlight this issue, the strongest phylogenetic signal in
our analyses indicated that only 4.4% of the variation in
escalation patterns was explained by phylogeny. Clearly, we
need more studies on closely related species to understand
the basis of contest evolution.
It is intriguing that SAM is not a general contest resolution
rule for species that signal their RHP (Jennings et al., 2004;
Briffa, 2008; Painting & Holwell, 2014; Schnell et al., 2015),
species that have high energetic expenditure during the fight
(e.g. Copeland et al., 2011) or species that fight with high
chances of injury (e.g. Rudin & Briffa, 2011). However,
signals may not necessarily be associated with mutual
evaluations of RHP. In some species individuals might signal
to show their willingness to engage or to remain in a contest
(Payne & Pagel, 1997; Jennings et al., 2012). For instance,
in contests of the hermit crab Pagurus bernhardus, losers seem
to signal that they are giving up by using within-phase
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
1439
de-escalated behaviours, while winners use escalation to
signal their willingness to persist in the contest (Briffa, Elwood
& Dick, 1998). Signals may also be used at the beginning
of the interaction to evaluate the rival. Once the decision
to engage or to continue is made, the contest may proceed
based on self-assessment. For example, in the fiddler crab,
Austruca mjoebergi, individuals seem to be able to decide when
to initiate a contest based on the difference between their size
and that of their rival (Morrell, Backwell & Metcalfe, 2005).
A third possibility is that signals might also be used as threat
displays to decrease the motivation of the rival to engage in
a contest (Számadó, 2003, 2008). Hence, signals might be
used for reasons not necessarily related to the evaluation of
the rival. Ultimately, we need first to identify the function of
each phase in a contest and only then proceed to investigate
the role of signalling.
Based on the variation in, and inconsistencies between,
model assumptions and empirical data, we suggest that
the classical models such as (E)WOA, SAM, and CAM
do not robustly explain how animals decide to give up
on a contest. We have a considerable body of evidence
suggesting that several factors might affect how the decision
to give up is made. Factors such as asymmetries in per-
ceived resource value (Mesterton-Gibbons & Heap, 2014;
Mesterton-Gibbons & Sherratt, 2016), previous experience
(i.e. winner and loser effects; Hsu & Wolf, 2001), personality
(Briffa, Sneddon & Wilson, 2015), damage capacity (Palaoro
& Briffa, 2017), self-inflicted injuries (Lane & Briffa, 2017),
and even individual fighting skill (Briffa & Fortescue, 2017)
are some of the variables that can potentially influence
how animals make decisions during contests. Among
these variables, only asymmetries in resource value have
been included in contest theory (Mesterton-Gibbons &
Heap, 2014; Mesterton-Gibbons & Sherratt, 2016). We
also have increasing empirical evidence for variation
in assessment strategies during contests (Jennings et al.,
2005; Hsu et al., 2008; Lobregat et al., 2019), and among
individuals (Guillermo-Ferreira et al., 2015; Edmonds &
Briffa, 2016). Such evidence has the potential to change
how we think about contests because assessment strategies
do not need to be fixed – individuals might vary according
to their genotype, or according to the resource value
(Mesterton-Gibbons & Heap, 2014; Mesterton-Gibbons &
Sherratt, 2016). But we still need to incorporate those ideas
into the core of contest theory. In conclusion, we highlight
that the classical models have been extremely important to
provide advances in the way we understand how individuals
settle contests. However, it may be time to move forward
and focus on new possibilities derived from such models.
V. CONCLUSIONS
(1) We tested if there was consistency among species in the
rules used by rivals to decide the winner of animal contests.
Although isolated papers show that some species settle their
contests based on the mutual assessment predicted by SAM,
1440
we found that the majority of species fight according to
self-assessment models. There was a positive relationship
between contest duration and loser RHP, but no relationship
for contest duration and winner RHP when rivals where
randomly paired. For RHP-matched rivals, contest duration
seems to increase with mean RHP of the pair. Therefore,
our results indicate that most species fight according to
(E)WOA models.
(2) We found a negligible phylogenetic signal in the
relationship between contest duration and loser and winner
RHP. This evidence indicates that closely related species do
not necessarily make similar decisions on how to give up on
a contest. Therefore, self- or mutual assessment are scattered
across the phylogeny without a clear pattern. However, the
lack of studies on closely related species might bias this result.
(3) Although relationships between contest duration and
traits linked to RHP provided support for self-assessment
models, other behavioural traits exhibited by species (such
as the occurrence of physical contact and predictable
behavioural phases during the fight) do not match
all self-assessment assumptions (particularly for (E)WOA
models). In addition, we found that for a subset of species,
escalation patterns were related to RHP asymmetry.
(4) We highlight that there is a considerable body of
evidence that could be incorporated into the core of contest
theory. The incorporation of new ideas and alternative
experimental approaches that are not based on correlations
between contest duration and traits linked to RHP might
solve the inconsistencies we identified herein. We know that
the classical models were important to understanding how
animals decide to give up on contests, but we now need
to move forward with empirical tests of new models that
incorporate new experiments and new evidence.
VI. ACKNOWLEDGEMENTS
We thank all authors that provided access to their
original data. We also thank Christina Painting, Mark
Briffa, Robert Elwood, Alastair Wilson and an anonymous
referee for helpful comments and suggestions. N.S.P.
thanks CAPES for providing a PhD scholarship.
A.V.P. has a post-doctoral grant from Fundação de
Amparo à Pesquisa do Estado de São Paulo (FAPESP
process number 2016/22679-3). P.E.C.P. has research
grants from Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq produtividade em pesquisa:
305561/2014-6 and 311212/2018-2; CNPq Edital Universal
2014: 443977/2014-3) and Fundação de Amparo à Pesquisa
do Estado da Bahia (Fapesb: JCB0034/2013).
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VIII. SUPPORTING INFORMATION
Additional supporting information may be found online in
the Supporting Information section at the end of the article.
Fig. S1. Phylogeny used to calculate the variance–
covariance matrix for meta-regression models involving the
relationship between contest duration and traits linked to
resource holding potential (RHP) for winners and losers in
randomly paired trials.
Fig. S2. Phylogeny used to calculate the variance–
covariance matrix for analyses involving the relationship
(Received 1 August 2018; revised 4 March 2019; accepted 6 March 2019; published online 27 March 2019)
Biological Reviews 94 (2019) 1430–1442 © 2019 Cambridge Philosophical Society
Nelson S. Pinto and others
between contest duration and mean trait value of the fighting
pair for contests in which rivals were paired in order to
minimize their difference in resource holding potential (RHP)
(RHP-matched rivals).
Fig. S3. Phylogeny used to calculate the variance–
covariance matrix for analyses involving the relationship
between contest escalation (i.e. change from less costly to
costlier behaviours during the contest) and the difference in
traits related to the resource holding potential (RHP) of the
fighting pair.
Table S1. Effect sizes [Fisher’s z (Zr)] and their variances (V )
calculated for all studies used in the test of the relationship
between contest duration and traits linked to resource holding
potential (RHP) for winners and losers in randomly paired
rivals.
Table S2. Effect sizes [Fisher’s z (Zr)] and their variances
(V ) calculated for studies used in the test of the relationship
between contest duration and mean trait values of the fighting
pair in resource holding potential (RHP)-matched rivals.
Table S3. Effect sizes [Fisher’s z (Zr)] and their variances
(V ) calculated for studies used in the test of the relationship
between contest escalation and trait differences between
winners and losers.