Ethology Ecology & Evolution 14: 129-139, 2002

Using behavioural characters

in phylogeny reconstruction

A.E. STUART 1,4, F.F. HUNTER 2 and D.C. CURRIE 3

1 Department of Zoology, University of Toronto, Canada

2 Department of Biological Sciences, Brock University, St. Catharines, Canada
3 CBCB, Royal Ontario Museum, Toronto, Canada

Behaviour remains underrepresented in phylogeny reconstruction, possibly

because the term ‘behaviour’ incorporates a wide range of phenomena, not all of
which are equally applicable to understanding evolutionary history. We assessed
the character homology (i.e., potential problems with coding) and homoplasy
(i.e., lability or convergence) for each of four types of behaviour (behavioural
categories, reaction stimuli traits, the specific movements of animals and quanti-
tative information relating to each of these behaviour types) and determined the
broad applicability of each behavioural type for phylogeny reconstruction. When
using behaviour to reconstruct a phylogeny we recommend the following order
of behavioural types: (1) animal movements; (2) quantitative components (pro-
viding that the animal movements are homologous); (3) reaction stimuli traits;
(4) behavioural categories.

KEY WORDS: behaviour, phylogenetics, homology, homoplasy, evolution.

Introduction . . . . . . . . . . . . . . . . . 129
Behavioural types . . . . . . . . . . . . . . . 130
Behavioural categories . . . . . . . . . . . . . 130
Reaction stimuli . . . . . . . . . . . . . . . 131
Animal movements . . . . . . . . . . . . . . 132
Quantitative information . . . . . . . . . . . . . 134
Implications . . . . . . . . . . . . . . . . . 135
Acknowledgments . . . . . . . . . . . . . . . 137
References . . . . . . . . . . . . . . . . . 137

INTRODUCTION

The application of phylogenetics has increased markedly in the past decade;

however, the use of behavioural data in phylogeny reconstruction remains under-
represented. During the 1970’s and 1980’s, questions were raised about whether

4 Send correspondence to: A.E. Stuart, Department of Ecology and Evolutionary Biology,

Haworth Hall, 1200 Sunnyside Ave., Lawrence, KS 66045. Ph: (785) 864-5788, Fax: (785) 864-5321
(E-mail: astuart@ku.edu).
130 A.E. Stuart, F.F. Hunter and D.C. Currie

behavioural data could or should be used to address macroevolutionary questions.

Some argued that behavioural homology is more difficult to establish because
behaviours were said to either be more labile (ATZ 1970, ARONSON 1981, BARONI
URBANI 1989) or more subject to convergence than other types of data (ATZ 1970,
ARONSON 1981, CARPENTER 1987, BARONI URBANI 1989). However, phylogenies recon-
structed with behavioural data have been highly congruent with phylogenies based
on morphological or molecular data (e.g., MCLENNAN et al. 1988, PRUM 1990,
MCLENNAN 1993, PATERSON et al. 1995, KENNEDY et al. 1996, STUART & HUNTER 1998,
MCLENNAN & MATTERN 2001). Further, recent studies assessing the validity of using
behaviour in phylogenetic study have shown no reason to assume a priori that
behaviour is any more subject to convergence than other types of data (WENZEL
1992, DEQUEIROZ & WIMBERGER 1993, WIMBERGER & DEQUEIROZ 1996, MCLENNAN &
MATTERN 2001). Despite these studies, there remains an ‘ingrained cultural avoid-
ance of behavior by many systematists’ (BUCHHOLZ & CLEMMONS 1997).
The term ‘behaviour’ represents a wide range of phenomena, predominantly
because animal behaviours are assessed in different ways depending on the ques-
tion under investigation. It is this broadness of the term ‘behaviour’ that may con-
tribute to the under-representation of behaviour in phylogeny reconstruction. In
this paper, we divide the term ‘behaviour’ into four separate types: behavioural cat-
egories (PROCTOR 1996), reaction stimuli traits (MAYR 1976), the specific movements
of an animal (MAYR 1976) and quantitative information of these behavioural types.
Characters from each of these four behavioural types are heritable and variable and
therefore can theoretically be used to reconstruct a phylogeny. However, it seems
probable that different types of behavioural data are not equally useful for under-
standing evolutionary history. The goal of this paper is to assess the potential of
each of these four behavioural types in phylogeny reconstruction.
In this paper, we define each behavioural type, discuss character homology
(i.e., potential problems with coding each behavioural type into phylogenetic char-
acters) and homoplasy (i.e., lability or convergence) and examine the broad applic-
ability of each behavioural type for phylogeny reconstruction. Homologous behav-
iours are those that arise from a common ancestor through descent (WENZEL 1992),
which can ultimately only be determined a posteriori from a phylogenetic analysis
(BROOKS & MCLENNAN 1991, WENZEL 1992, LAUDER 1994). Practically, phylogenetic
characters and character states are chosen based on some a priori assumptions of
homology (see WENZEL 1992 for discussion), and these assumptions are tested by
the resultant phylogeny. There are at least two possible reasons that a character
state is homoplasious: the a priori assumption of homology is incorrect or the
character state is convergent. This paper is predominantly aimed at behaviourists
or macroevolutionary biologists interested in constructing a phylogeny that uses
any behavioural characters. However, we hope other fields of behavioural biology
will benefit from the discussion of different behavioural types.

BEHAVIOURAL TYPES

Behavioural categories

Behavioural categories are general terms that describe suites of behaviours

that share some broad phenomena, usually, but not always, relating to function
Behaviour in phylogeny construction 131

(PROCTOR 1996). The specific details of these broad terms are not necessarily the
same among taxa. For example, the phrase “lekking behaviour” is a broad term that
groups all taxa in which males hold a territory without any resources for the
females. The term does not distinguish among the behaviours performed by indi-
viduals of a given species that serve to maintain a lek. The phylogenetic character
“lekking behaviour” with states, (0) present; (1) absent, would group all animals
that lek, regardless of the way the lek is established, maintained or the behaviours
that occur while on the lek. This character will be convergent among many taxa.
Cannibalism is also a behavioural category describing taxa whose individuals
eat members of their own species. The character, “cannibalism” with states, (0) pre-
sent; (1) absent, does not distinguish between adult male lions eating young after
taking over a harem, female spiders that eat males after mating, or species whose
individuals cannibalize weak or sick individuals or the manner in which individuals
prey upon, capture or eat their prey. Other similar examples include: aggressive
behaviour, monogamous vs. polygamous behaviour, cooperative breeding and gre-
garious behaviour.
Behavioural categories are rarely thought to be homologous in a phylogenetic
sense (WENZEL 1992, PROCTOR 1996). The terms represent phenomena that are likely
to be distributed among a wide range of taxa, and thus their selection is, in effect,
intended to maximize the probability of homoplasy (PROCTOR 1996), not to denote
shared derived character states. Thus, we believe that broad behavioural terms
should not be used as characters for phylogeny reconstruction.

Reaction stimuli

One obvious component of behaviour is the ability of animals to react selec-

tively to specific stimuli (‘perceptual traits’ of MAYR 1976). The objects that promote
a reaction are often easy to characterize. MAYR (1976) uses the example of predato-
ry wasps whose prey objects are spiders, caterpillars or sawfly larvae. These objects
will induce a predatory response in some species of wasps, but not in others. By
determining which species of prey is used by different wasp species, this informa-
tion can be coded as a phylogenetic character. Another example is the type of
building material used by animals to build a structure. If an individual selects and
manipulates a particular type of material appropriate for building a structure, dis-
criminating against other types of materials, then the type of building material can
be coded as a character. This component of behaviour is thus a description of the
object(s) that elicit a given behavioural reaction (e.g. species of prey, type of build-
ing material).
When coding characters for phylogeny reconstruction, there is no reason to
assume a lack of homology in traits that elicit a response. Thus, the character ‘Type
of building material: (0) organic, (1) mineral’ would potentially be homologous.
However, these traits may be prone to homoplasy (MAYR 1976). MAYR suggests that
an evolutionary switch from one type of prey to another may not be exceedingly
difficult. TABER’s (1994) data supports this hypothesis; he studied the life history
and host utilization traits in 11 species of plant lice (Rhopalosiphum) and found
that of 12 behavioural characters studied, none of the possibly conserved traits
were those involving choice of the host plant taxa. Similarly, CARPENTER (1987)
indicated that the character ‘paper type’ is not particularly informative when con-
structing a phylogeny of the wasps, because the character varies within genera of
132 A.E. Stuart, F.F. Hunter and D.C. Currie

the Vespinae. The paper in a wasp nest is either pliable or brittle depending on
whether the wood fibres (the objects that elicit nest building) collected are sound
or rotten.
Although homology is ultimately determined a posteriori from a phylogeny
reconstructed with many characters (LAUDER 1994), it is worthwhile considering the
potential homoplasy of reaction stimuli traits before reconstructing a phylogeny.

Animal movements

Another obvious type of behaviour is the specific movement of an individual

and its body parts during the course of any given behaviour (‘locomotory compo-
nents’ of MAYR 1976). Behavioural repertoires, such as mating, feeding, or building
behaviour are an aggregation of numerous smaller behavioural units. Evolution can
potentially act on each behavioural unit over time, eventually culminating in the
present-day behavioural repertoire of an animal. Thus, analyzing behavioural reper-
toires tells an ‘evolutionary story’ that can be deciphered by searching for shared
derived character states. WENZEL (1992) states “if the complex movements of sever-
al species take the same distinct form in the same context and appear to be largely
innate, they may be thought of as homologous”.
DEQUEIROZ & WIMBERGER (1993) performed a thorough comparison of the lev-
els of homoplasy of behavioural and morphological data. They compared the mean
consistency index (CI) of behavioural characters and morphological characters and
found that behaviour is no more homoplastic than morphology. Because they were
looking at the mean CI of individual characters, there was no constraint on the
number of characters included in their analyses. As such, some analyses used had
very few behavioural characters. Additionally, DEQUEIROZ & WIMBERGER (1993)
intentionally chose a broad definition of behaviour, which included all four cate-
gories described here. To assess animal movement data specifically, we searched for
papers where animal movement data was used to construct entire phylogenies and
the outcome compared to phylogenies constructed with other data (i.e., morpholog-
ical and/or molecular). Nine such studies were found representing a variety of
species and behaviours (Table 1). In all these studies, characters are delimited
based on details of the specific movements of animals during their respective
behaviours and all of these studies find behavioural data to be highly congruent
with other types of data (see references in Table 1). For example, PRUM (1990) used
29 characters relating to the detailed description of lek display behaviour in 21
species of Neotropical Manakins. When the behavioural characters were analyzed
alone, and when combined with syringeal characters (morphological traits), the
results were highly congruent with the independent syringeal hypothesis of phy-
logeny. SLIKAS (1998) was the only study to find behaviour less reliable than other
data. However, in her study, only behaviours late in the sequence of stork courtship
behaviours had higher levels of homoplasy than DNA-DNA hybridization data;
behaviours occurring early were congruent with phylogenetic relationships and
showed little homoplasy. The congruency of animal movement phylogenies with
phylogenies based on other data indicates that animal movement data are reliable
indicators of evolutionary history. Further, because there is no significant difference
(paired t-test: t = 0.54, P > 0.05) between the consistency indices (CI) of the respec-
tive datasets (Table 1), animal movements are also no more subject to homoplasy
than other types of data.
Behaviour in phylogeny construction 133

Table 1.
A list of reconstructed behavioural phylogenies including authors, taxa, behaviours studied, number
of characters in each analysis, and other data used for comparison and consistency indices (CIs).
Consistency indices for the comparison data were either found in the original paper or in cited ref-
erences within as noted. EBERHARD (1982) was not used for the statistical test. When data was not
available for other references, a CI of 1.00 was used (†). All authors found the topology of the
behavioural phylogenies to be congruent with that of the other data used.

Reference Study Behaviour # of Data of CI CI

organism characters comparison behaviour other data

EBERHARD (1982) Spiders Web-building 10 Adult morphology N/A N/A

MCLENNAN et Gasterosteid Reproductive 27 Biochemical, 0.90* 0.67*

al. (1988) and fishes morphological
MCLENNAN (1993)

ARNTZEN & Newts Courtship 16 Biochemical 0.81 N/A†

SPARREBOOM (1989)

PRUM (1990) Neotropical Display 29 Morphological 0.78! 0.90‡

manakins

PATERSON et al. Albatrosses, Behaviour, 72 Molecular 0.52 0.53

(1995) petrels, life history
penguins

KENNEDY et al. Pelecaniforms Social 37 Morphological, 0.74 0.67§

(1996) behaviour molecular

SLIKAS (1998) Storks Courtship 42 DNA-DNA 0.85 N/A†

displays hybridization

STUART & HUNTER Black Cocoon 14 Cytological, 0.92 1.00

(1998) flies spinning morphological

* MCLENNAN (2000) personal communication. ! 15 uninformative characters removed from PRUM

(1990), which reduced the behavioural CI from 0.85 to 0.78. ‡ PRUM unpubl. data in DEQUIEROZ and
WIMBERGER (1993). § CRACRAFT (1985), SIEGEL-CAUSEY (1988).

No references were found that refute the utility of animal movement charac-
ters in phylogenetic analysis even though the examples include a wide variety of
taxa and cover an array of behavioural phenomena (Table 1). Successful use of ani-
mal movements in phylogenetic analysis requires two steps. First, one must make a
detailed account about how an organism behaves during a behavioural repertoire.
This will provide an understanding of all of the movements and actions an individ-
ual makes when performing a particular behavioural repertoire (see MILLER 1988
for detailed methodology). A lack of movement is also important and should be
included in any analysis. Thus, a complete description of the feeding repertoire of a
trap door spider, for example, should not only contain information about detailed
movements of an individual during prey capture, but also information such as the
134 A.E. Stuart, F.F. Hunter and D.C. Currie

orientation and location of the spider within the trap, and the orientation of its
limbs while the individual is waiting. Reproductive and building behaviours are
typically preferred (e.g., Table 1), but there is no reason to assume that other
behaviours (e.g., feeding) cannot be studied in a similar fashion.
The second step involves the study of other closely related species in a similar
manner. Phylogeny reconstruction necessarily requires the comparative study of
similar behaviours in order to discover shared derived character states. The com-
parative approach also aids in the description of behavioural repertoires because
some phylogenetically informative movements may go undetected until some varia-
tion of the movement is observed in another taxon. As such, observation of multi-
ple taxa typically reveals aspects of behaviour that had previously gone unnoticed
(see MCLENNAN et al. 1988, PRUM 1990, PATERSON et al. 1995, KENNEDY et al. 1996,
STUART & HUNTER 1998 for examples of this methodology).
In order to use animal movement traits as phylogenetic characters there must
be a consistent definition of a ‘behavioural unit’ so that all ethologists follow the
same guidelines (WENZEL 1992). PROCTOR (1996) suggests that one reason for the
limited use of behaviour in phylogenetic studies is the difficulty in delimiting char-
acters. Her example is that of one observer recognizing three distinct behaviours in
fish, ‘mouth gape’, ‘bow’ and ‘tail fan’, whereas another considers them to be one
action: ‘fanning tail while bowing with open beak’. In order to recognize character
states effectively it is necessary to study a group of closely related species. If after
studying 20 species of fish, three species performed all three behaviours in the
same order and none performed any of the solitary behaviours, then in the example
provided above, one should consider ‘fanning tail while bowing with open beak’ as
one character state. However, if certain species performed only ‘mouth gape’, then
this character state should be recognized. A behavioural unit, therefore, is any
behavioural trait shared by two or more taxa (WENZEL 1992).

Quantitative information

Many behavioural studies concentrate on gathering quantitative information

and each of the three types of behaviour discussed above have observable quantita-
tive components. A behavioural category, such as aggressive behaviour, can be
quantified by determining the frequency of aggressive bouts for individuals of a
given species. In a trait that elicits a response, such as the species of prey captured,
one can count the number of prey captured or the rate of prey capture. For specific
movement information, each behavioural unit can be counted or timed; for exam-
ple, the length of time for each building stage during black fly cocoon spinning
(STUART & HUNTER 1998).
There is potential difficulty in determining the homology of quantitative traits
because their homology is dependent on the homology of the type of behaviour (as
discussed above) being quantified. Thus, the number of times, or length of time
that an animal behaves is only potentially homologous if the behaviour being per-
formed is first shown to be homologous. Consider, for example, black fly larval
cocoon spinning behaviour (see STUART & HUNTER 1995, 1998). Each building stage
is characterized by a specific pattern of silking and a method of moving the head
from the lateral surface of one side of its body to the other (i.e., method of cross-
ing); both of which can be quantified (e.g., number of silk strands laid down per
stage and number of times an individual crosses per stage). To demonstrate the dif-
Behaviour in phylogeny construction 135

ference in assessing homology of these two quantitative traits we use the first stage
of spinning, the ‘initial structure’ stage, as an example in three species of black fly,
Prosimulium fuscum, Simulium venustum and Ectemnia invenusta. During this
stage, larvae of all three species cross their head from one side of their body to the
other by straightening their body and moving in an arc across to the other side.
When optimized onto a phylogeny, this method of crossing is homologous; there-
fore, the quantitative information (e.g., number of crosses per stage) is potentially
homologous. There are, however, two different methods of silking during the initial
structure stage; transverse silking (i.e., larvae place strands perpendicular to the
longitudinal axis of the resting larvae) performed by P. fuscum, and longitudinal
silking (i.e., larvae place strands parallel to the longitudinal axis of the resting lar-
vae) performed by S. venustum and E. invenusta (STUART & HUNTER 1998). Since
the method of silking is not homologous among the three species, the quantitative
information (e.g., number of strands per stage) cannot be considered homologous.
Quantitative traits are potentially homoplasious, particularly across a broad
range of taxa, because there is typically sufficient genetic variation within popula-
tions to permit evolution of the range of behavioural variation observed (FOSTER
1999). Therefore, shifts in the frequency of expression of behavioural traits could be
common. However, it is currently difficult to assess amount of homoplasy in quanti-
tative traits because they are rarely associated with known behavioural homologs.
If animal movement or traits that elicit a response are shown to be homolo-
gous from a phylogenetic analysis, then quantitative information becomes valuable
phylogenetic information (see discussion in previous section). KOREF-SANTIBAÑEZ
(1972) performed an analysis of both qualitative and quantitative aspects of
courtship behaviour in the semispecies of Drosophila paulistorum. Qualitative
aspects of the animal movements were described in detail; the courtship elements
and their sequence were similar for all semispecies except for one trait termed
“shaking vibration” in males (very rapid vibrations of both wings which hardly
leave the horizontal position), which did not occur in two semispecies. The other
courtship traits (e.g., orientation, tapping vibration, scissoring), which were quali-
tatively similar, were counted, timed and compared among the semispecies.
Although no phylogeny was reconstructed with these data, each semispecies had a
characteristic courtship pattern based on relative frequencies of the courtship ele-
ments, and thus these data could potentially be used in phylogeny reconstruction.

IMPLICATIONS

Many misconceptions about the use of behavioural data in phylogenetic

analyses occur because the term ‘behaviour’ applies to a variety of phenomena that
are not equally applicable for understanding genealogical relationships. Although
any observable trait, providing that it is heritable and variable, can theoretically be
used as a character in a phylogenetic analysis, it seems prudent to be aware of the
expectations and limitations of different behavioural types before they are included
in a phylogenetic analysis. It is also important to differentiate the behavioural types
and their respective expectations when assessing the phylogenetic usefulness of
‘behaviour’ in general.
Some authors believe that behavioural characters are more labile and subject
to homoplasy than are morphological characters (e.g., ATZ 1970, WCISLO 1989,
136 A.E. Stuart, F.F. Hunter and D.C. Currie

WEST-EBERHARD 1989, HUNTINGFORD et al. 1994); whereas others assert that there is
no reason to assume a priori that behaviour is any more subject to convergence
than are other kinds of characters (e.g., LAUDER 1986, MCLENNAN et al. 1988, WEN-
ZEL 1992, DEQUEIROZ & WIMBERGER 1993, WIMBERGER & DEQUEIROZ 1996). We
believe that both sides may be correct because the behavioural lability may largely
depend on the type of behaviour studied. Although single studies do not tend to
present data from all types (i.e., behavioural categories, reaction stimuli, animal
movements and quantitative), certain trends are noticeable. Studies aiming to
address adaptation level phenomena tend to use the presence or quantification of
behavioural categories (e.g., presence or absence of cannibalism; intensity of
aggressive behaviour) or reaction stimuli traits; these studies tend to find plasticity
in behaviour (e.g., ARNOLD 1981, 1992; RIECHERT 1986; HUNTINGFORD et al. 1994;
FOSTER 1995; FOSTER et al. 1996). For example, FOSTER et al. (1996) noted that iso-
lated stickleback populations divide into limnetic and benthic sub-populations that
have certain morphological and behavioural characteristics. Comparisons among
populations suggested that adaptive change and homoplasy of behavioural pheno-
types might be very common. The behavioural characteristics used by FOSTER et al.
(1996) are either presence of behavioural categories (e.g., presence or absence of
sneaking and cannibalism), or quantified behavioural categories (e.g., amount of
boldness, conspicuousness of courtship). Similarly, HUNTINGFORD et al. (1994) stud-
ied adaptive variation in antipredatory behaviour in three-spined sticklebacks and
showed that the intensity and nature of the antipredatory responses changed when
predation regimes were varied. HUNTINGFORD et al. (1994) therefore used quantita-
tive and reaction stimuli behaviours to determine trait adaptability. The inclusion
of these traits into the construction of a phylogeny should be done with caution.
Studies aiming to address questions of genealogical history tend to use the
details of animal movements, and these studies find that behaviour is no more
labile than other types of data (e.g., references in Table 1). For example, MCLENNAN
(1993) documented 51 characters relating to the movements of sticklebacks during
courtship behaviour and found that behavioural information actually provided a
less ambiguous picture of relationships than did morphological data (MCLENNAN
1993, MCLENNAN & MATTERN 2001).
Behavioural information can be a valuable source of information for under-
standing genealogical relationships among taxa and there is no reason for it to be
avoided by systematists. If one is interested in using behavioural characters to recon-
struct a phylogeny, we recommend using the following order of behavioural types:
(1) animal movements; (2) quantitative components (providing that the animal
movements are homologous); (3) reaction stimuli traits; (4) behavioural categories.
Describing qualitative aspects of animal movements in detail for a number of
taxa have repeatedly produced phylogenies that are corroborated by other data,
regardless of taxa and/or behaviour studied (Table 1). Therefore, gathering behav-
ioural data for phylogeny reconstruction should begin with an ethogram detailing
the specific movements of individuals while performing a given behaviour. Many
species should be studied similarly, characters gleaned from this analysis and a
phylogeny reconstructed. Then, if the initial analysis of animal movements pro-
duces polytomies (i.e., unresolved regions) in the phylogeny, or if one is interested
in constructing phylogenies of populations within a species, quantitative informa-
tion can be investigated. It is only appropriate to use quantitative information as
phylogenetic characters when the behaviours being quantified are homologous.
Thus, a qualitative analysis of behaviour must precede or coincide with a quantita-
Behaviour in phylogeny construction 137

tive analysis. We suggest that reaction stimuli traits be used with caution and
behavioural categories be avoided.
Characters arising from analyses of animal movements are the most likely to
produce phylogenies that concur with other types of data, but unfortunately, this
information is rarely gathered. Although it is possible to gather quantitative infor-
mation and document stimuli that elicit a response while studying animal move-
ments, it is not possible to gather information about an animal’s movements with-
out observing them directly. Thus, we propose that the construction of behavioural
phylogenies involve, first, an analysis of animal movements and characters result-
ing from this data. Since other behavioural information can be gathered simultane-
ously, this may also be included, but one should consider the appropriateness of
behavioural types independently.
Many biologists have detailed knowledge of the behaviours performed by their
study organisms and this information can be of significant value for understanding
the evolutionary history of these taxa. We hope that this discussion encourages
behaviourists to use their data to reconstruct phylogenies and phylogeneticists to
use behavioural characters in phylogeny reconstruction.

ACKNOWLEDGMENTS

We thank NSERC for PGSA and PGSB awards to A.E. Stuart and operating grants to
F.F. Hunter and D.C. Currie. We thank C. Currie, L. Jesson, D. McLennan, J. Napolitano, J.
Rock, L. Rowe and J. Wenzel for comments on earlier versions of this manuscript.

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