Behav Ecol Sociobiol
DOI 10.1007/s00265-009-0865-8
METHODS
Sampling animal association networks with the gambit
of the group
Daniel W. Franks & Graeme D. Ruxton & Richard James
Received: 15 May 2009 / Revised: 14 September 2009 / Accepted: 16 September 2009
# Springer-Verlag 2009
Abstract Ecologists increasingly use network theory to
examine animal association patterns. The gambit of the
group (GoG) is a simple and useful assumption for
accumulating the data necessary for a network analysis.
The gambit of the group implies that each animal in a group
is associating with every other individual in that group.
Sampling is an important issue for networks in wild
populations collected assuming GoG. Due to time, effort,
and resource constraints and the difficulty of tracking
animals, sampled data are usually a subset of the actual
network. Ecologists often use association indexes to
calculate the frequency of associations between individuals.
These indexes are often transformed by applying a filter to
produce a binary network. We explore GoG sampling using
model networks. We examine assortment at the level of the
group by a single dichotomous trait, along with many other
network measures, to examine the effect of different
Communicated by J. Krause
Electronic supplementary material The online version of this article
(doi:10.1007/s00265-009-0865-8) contains supplementary material,
which is available to authorized users.
D. W. Franks (*)
YCCSA,
Department of Biology and Department of Computer Science,
University of York,
York 105, UK
e-mail: df525@york.ac.uk
G. D. Ruxton
Division of Ecology and Evolutionary Biology,
Faculty of Biomedical and Life Sciences, University of Glasgow,
Glasgow 128, Scotland
R. James
Department of Physics, University of Bath,
Bath 27, UK
sampling regimes, and choice of filter on the accuracy
and precision with which measures are estimated. We find
strong support for the use of weighted, rather than filtered,
network measures and show that different filters have
different effects depending on the nature of the sampling.
We make several practical recommendations for ecologists
planning GoG sampling.
Keywords Association networks . Gambit of the group .
Network sampling
Introduction
Ecologists are often interested in understanding social
associations and interactions between animals. To examine
these associations, ecologists are increasingly turning to
network theory, within which they represent animals as
nodes and associations as links between nodes. Network
theory offers a set of statistical measures to quantify,
describe, and compare the structure of social relations.
The availability of network analysis software has provided
statistical tools that allow biologists to readily apply
network theory to their study populations (Newman 2003;
Croft et al. 2008; Whitehead 2008). A network approach is
often applied as a descriptive tool to help understand the
social structure of a population (e.g., dolphins, Lusseau
2003; fish, Croft et al. 2004; African buffalos, Cross et al.
2004; and sea lions, Wolf et al. 2007). A key aim of animal
social network analysis is often to look for a biological
signal or biological meaning in the potentially tangled mess
of observed associations between individual animals.
The gambit of the group (GoG) is a relatively simple
method for accumulating relational data (Whitehead and
Dufault 1999). When using this technique, ecologists must

make the assumption that all individuals seen grouping
together, during an observation census, are associating with
every other individual in that group. On observing groups,
the ecologist notes the number of times each individual is
observed in the same group as each other individual to
construct indexes of associations between individuals. After
numerous censuses and numerous group observations in
each census, the ecologist will be left with relational data
representing the association strength between animals based
on group association. In a fission–fusion animal society, a
main parent group can fracture into smaller groups (such as
fish shoals forming each day), and the smaller groups, later,
coalesce to rejoin as the parent group (such as fish for
several daytime shoals collecting at a bank at night, Croft et
al. 2003). The fission–fusion process is likely to affect the
social structure of many animal societies. The GoG is a
potentially convenient way of sampling such systems.
Ecologists capture group associations over a number of
days, weeks, or years, which allow them to collect
relational data regarding associations between individuals.
Sampling is an important issue for GoG networks in wild
populations (Croft et al. 2008). Due to time, effort, and
resource constraints and the practical difficulty involved in
tracking animals, the sampled relational data produced are
usually only a subset of the actual network. That is, not all
individuals from the population will be sampled at each
census point. Incorrect sampling protocols risk creation of
artifactual results. For example, assortment by degree
(where the degree of an individual is the number of
connections from that individual to distinct other individ-
uals and assortment by degree examines whether individ-
uals are generally connected to other individuals with
similar degree scores) is typically identified by a positive
Pearson correlation coefficient between a focal individual's
degree of connectedness and the mean degree of connect-
edness of the individuals to which it is directly connected
(Lusseau and Newman 2004; Croft et al. 2005). Ecologists
often use association indexes to calculate the frequency of
associations between individuals, rather than a simple count
of the number of times individuals have been sampled in a
same group. A sensible choice of association index can go
some way to alleviating certain types of sampling bias (see
Whitehead 2008 and Croft et al. 2008, for more informa-
tion). In situations where there is a sampling bias such that
it is more likely that two individuals will be recorded in a
given census if they are in separate groups, for example,
then the half-weight index may help to compensate by
weighting separate group observations at half the weight of
other observations (Cairns and Schwager 1987). Various
indexes have been developed to account for different
sampling biases. More detail can be found in papers by
Cairns and Schwager (1987); Ginsberg and Young (1992);
Bejder et al. (1998); Whitehead (1999); and Whitehead and
Behav Ecol Sociobiol
Dufault (1999). Additionally, appropriate randomization of
the relational data can help to compensate for some types of
biases; see Croft et al. (2008) and Whitehead (2008).
In many of the early applications of network analysis to
group-derived animal associations, researchers transformed
their association indexes by applying an edge filter to
produce a binary network representing (usually) just the
stronger associations (e.g., Croft et al. 2004; Sundaresen et
al. 2007; and Wolf et al. 2007). If a filter threshold of 0.5
was used, for example, then connections with an associa-
tion index greater than or equal to 0.5 would be retained as
a network connection and connections with a weight less
than 0.5 would be removed. By transforming the data in
this way, the researcher is able to calculate a multitude of
network metrics often developed in the social sciences
(Wasserman and Faust 1994), but there are potential issues.
The choice of appropriate value for the filter threshold is a
difficult problem. Filtering a network discards data that
could improve the accuracy of any analysis (and maybe, the
weak connections are important—see Granovetter 1974).
This reduction in data resolution could prove to be
important, especially when working with limited sample
sizes. If the threshold is too high, then we lose more data. If
the threshold is too low, then we strengthen the weak
connections. Because of this problem, ecologists are
beginning to move away from the filtering approach by
keeping the initial association indexes and conducting a
weighted analysis of the network (Lusseau et al. 2008).
Even then, we cannot be sure how much of the biological
signal of interest is being measured. For example, some of
what looks like a signal might be accounted for by
sampling bias (see e.g., Croft et al. 2008). Here, we
illustrate and explore this issue using model networks in
which we insert a known biological signal. To ground the
model, we make use of calculations of Croft et al. (2008)
who looked at a sample of long-term red deer data and
select parameters that are more or less consistent with this
population (in addition to further explorations). The
biological signal is a controlled assortment at the level of
the group by a single dichotomous trait (e.g., sex). We
examine this, along with other network measures, to
examine how well network measures pick up on this signal,
given different sampling regimes, choice of weighted or
unweighted measures, and choice of filter. The aim is to
help guide field biologists in planning their data collection,
so as to collect data that makes the most effective use of
network theory.
The model
We assume a population of n animals, where each animal is
either male or female (where f is the proportion of females).
Behav Ecol Sociobiol
That is, each animal is allocated its trait/sex independently
and randomly, with the probability of an individual being
female, the value of the parameter f. Although we use male
and female as traits for the animals, our model equally
applies to a variety of traits (e.g., infant/adult). In addition to
preference by trait, each animal, i, is allocated a preference
toward grouping with each other individual animal, j. These
biases are stored as a matrix Bij, where Bij =Bji. The greater
the value of Bij, the higher the relative probability that i and
j will be allocated to the same group. We draw each Bij, i<j
from a uniform distribution ½0 ! d; 0 þ d $ for some param-
eter d ∈ [0,1], with increasing d, signifying increasing levels
of preference. Each value of Bij is also increased by b for
individuals of the same trait (in this case by sex,
representing a sex bias in preferential associations) and
decreased by b if they are the opposite trait, where b ∈ [0,1]
is a constant. Thus, the greater the value of b, the stronger the
preference for assorting with same sex individuals. Note that
for our purposes, we do not need to accurately model a
mechanism by which real animals decide to join groups; we
simply need to capture preferential associations and assorta-
tive mixing in the groups along with grouping constraints
governed by the group size distribution. Our algorithm
allows us to capture these desirable properties.
We simulate sampling by recording associations over a
number of censuses, m. In each census we define new
groups, allocate the animals to groups, and sample the
groups. The preference matrix (B) remains fixed through-
out; that is, we make the assumption that the social
dynamics of the group are at some sort of equilibrium.
This process is repeated until all censuses are complete.
Forming groups
In each census, for m censuses, new groups are formed as
follows: A group l is defined, and its size, g (representing
its capacity), is selected from a Pareto distribution:
g ¼ ð1 ! U Þ!1=K ð1Þ
where K is the shape of the distribution, and U is a random
number between zero and one selected from a uniform
distribution. Group sizes are rounded to the nearest whole
number, and when a group size of zero is drawn, we draw
again from the distribution until a nonzero rounded group
size is drawn. Many group living animals have a group size
distribution, which is right skewed and approximately,
Pareto or exponential (Okubo 1986; Krause and Ruxton
2002). Our group size distribution was selected to match
the general shape distribution in the red deer data (see
below). Additional groups (along with their group size),
continue to be defined in this manner until we have enough
total capacity (in the sum of the capacity of all groups) to
l
allocate all animals to a group Σ gl ( n. If once we define
the size of the final group, the total capacity is greater than
the number of animals, l then, the size of the final
Σ gl > n,
group is reduced, such that the capacity equals the number
l
of animals, Σ gl ¼ n. Once the groups and their respective
capacities are defined, the animals are allocated to a group
as follows: Each group l with remaining capacity (that is, to
which we have previously allocated fewer individuals than
the group’s capacity, so all groups start with remaining
capacity) is allocated a scores sli pertaining to each
unallocated (i.e., not in a group) animal i defined as:
j
X
sli ¼ Bij þ 0:001 ð2Þ
al
where ai is the number of individuals that have previously
been allocated to group l. A small value (0.001) is added to
each score to give each group a chance of being selected,
even if there are no animals in it. If a group has filled its
capacity then it cannot be selected (ski =0, for each group k
with no remaining capacity). An animal to allocate i is then
selected from all unallocated animals with the roulette
wheel selection algorithm (Mitchell 1998), where their
proportion of the roulette wheel is the maximum score
for that animal to be allocated to any group. We select
individuals in this manner, rather than from a uniform
distribution, because it results in a stronger affect of the
bias to be grouped with other individuals. This means
that animals with stronger association preferences will,
generally, be allocated to groups earlier than those with
weak preferences. A group is then selected with the
roulette wheel selection algorithm, where the group’s
proportion of the roulette wheel is their score for the
selected animal sli, and the selected animal is then allocated
to the selected group (and its remaining capacity is
updated). This process is repeated until all animals are
allocated to a group (and thus, all groups are full). When
b=0, then allocation to groups is not assorted by trait and as
b increases towards b=1, there is an increased bias for the
group to be assorted by trait (because b is reflected in each
group’s score for an individual through Bij). Likewise,
increasing d would affect Bij and thus, the probability of
individuals being allocated to a group with other particular
individuals.
Sampling the groups
After the groups have been allocated, we simulate gambit
of the group sampling. We simulate the sampling process
where we have complete information in that we sample all
groups and when we only have partial information where
we sample a proportion p of the animals (when p=1.0 we
have complete information). Our assumption that groups

reform for each census corresponds to the sampling of a
fission–fusion system where successive samples of a
population are taken at time intervals long enough to allow,
in principle, complete mixing. Thus, we do not model
timescale issues related to short-term switching of individ-
uals between groups during a census or the fracturing of
groups. We simply capture biases in the allocation of
individuals to groups driven by their association prefer-
ences (captured in the matrix B).
Sampling proceeds as follows: A group l is selected at
random (from groups we have not previously sampled in
this census, each with the same probability of being
selected) and added to our list of groups to observe g. We
repeat this process until we l have observed our sample
proportion of the population Σ gl ( p:n.
We then model gambit of the group sampling by
recording the group allocations of observed individuals in
each census. This models the way that biologists usually
record this information. From these data, we then construct
our association index. We used the simple ratio index (SRI)
(Ginsberg and Young 1992) in the standard manner (Croft
et al. 2008) to calculate the association weight wab between
animal a and animal b:
X
wab ¼ : ð3Þ
X þ Yab þ Ya þ Yb
where X is the number of times a pair of animals were
observed in the same group, Ya is the number of times a
(but not b) was observed in a census, Yb is the number of
times b (but not a) was observed in a census, and Yab is the
number of times both a and b were observed in different
groups in a census.
Parameter values
Croft et al. (2008) analyzed a small slice of the long-term
investigation of the red deer (Cervus elaphus) population
on the island of Rum (Clutton-Brock et al. 1982). One of
the pieces of information gathered in this study has been the
identity of members of groups of deer during regular visual
censuses. This is exactly the sort of data for which we can
construct networks via the gambit of the group. Croft et al
(2008) used as an illustrative data set 26 censuses of group
membership between January and May 1990. There were
342 deer (212 female and 130 male) in their network with
all links included. Croft et al. (2008) used this data set to
explore the effect of filtering on simple node-based binary
network measures and whether these measures were
sensitive to trait segregation. For the present study, we use
similar population parameters to the earlier study. We,
therefore, set n=400 (the number of individuals), f=0.6 (the
proportion of individuals that are female), and set both the
mode and shape of the Pareto distribution to one to fit the
Behav Ecol Sociobiol
red deer group distribution. We examine sampling for a trait
preference (b=1) and no trait preference (b=0). Unless
otherwise stated, there is no individual pairing bias (d=0).
For all cases, we vary m (the number of censuses) from one
to 23, and p (the proportion of individuals sampled in each
census) from 0.1 to one and average the results over 30
runs.
Network analysis
We first analyze the network using the raw SRI values and
use weighted network measures that take into account the
values of the association indexes. Next, we examine the
impact of selecting different threshold values and compare
the results to our weighted analysis. Although biologists
usually collect weighted association data (as in our model),
they often apply a threshold to the data so that they can
analyze the network using binary measures (the process of
filtering discussed in the “Introduction” section). For
example, if the threshold is set to 0.5, any association
strength greater than or equal to 0.5 will become one, while
any other association strength becomes zero. Thus, this
process eliminates weaker network connections and (usu-
ally) strengthens stronger ones. What values should be used
for the threshold is an open question, and may vary
depending on the type of system being examined. Typical-
ly, ecologists will experiment with the threshold until it
produces a network that matches their intuition. Implement-
ing both a weighted and binary analysis allows us to
compare the two approaches and to look for any differences
in sampling effects between them.
Results: weighted networks
Figure 1 shows the average (over 30 replications for each data
point) of a number of weighted network measures taken from
our model. Each figure shows the consequence of the number
of censuses and the proportion of individuals sampled in each
census. The value of the measure with the maximum number
of censuses and all individuals sampled can be taken as the
most reliable indicator of the true value of the measure. The
results, generally, show (with the exception of the mean
degree) that the number of censuses is much more important
than the proportion of individuals sampled for most measures
(although, both affect the results). Figure 1g shows a measure
of trait assortativity where there is no assortativity in the
population being sampled (b=0). Conversely, Fig. 1h shows
the same measure where there is assortativity by trait in the
population (b=1). For all sampling levels (even very low
levels), we do not find assortativity by trait when it does not
exist. Additionally, we always find assortativity by trait when
it does exist. However, if the sampling is inadequate—
Behav Ecol Sociobiol

Fig. 1 Mean network measures

for weighted networks over the
number of censuses and propor-
tion sampled. Each data-point is
averaged over 30 replications
taken from networks each with a
different random seeds. a Mean
degree. b Mean path length. c
The clustering. d The degree
correlation. e The betweenness.
f Mean C.V. of the betweenness.
g The mean weighted trait
assortativity where there is no
trait bias (b=0). h The trait
assortativity where there is a
strong trait bias (b=1). There is
no individual bias (d=0)

particularly, if not enough censuses are taken—we record a
level of assortativity that is very much artificially high (note
that the estimate with most samples settles on ∼0.3, while the
estimate with the least samples is ∼0.7). A similar
observation applies to most of the other measures. We can,
therefore, see that without enough samples, researchers
sampling real populations are unlikely to find assortativity
where it does not exist, but are likely to find highly
exaggerated values where a population is moderately
assorted. For most measures, however, increasing the
number of censuses gives reasonable results.
Results: binary networks
Figures 2, 3, and 4 show the results of setting the threshold
to 0.2, 0.5, and 0.8, respectively. We select these threshold
values to show the impact of selecting high medium and
low values. It is easy to see from comparing these figures
that the choice of threshold can drastically change the
resulting network measures. Additionally, the different
threshold levels have different effects depending on the
level of sampling (conversely, the level of sampling has a
different effect depending on the threshold).
A threshold of 0.2 gives results that a qualitative
visual inspection shows are, easily, most similar to our
weighted analysis (Fig. 2). However, the level of trait
assortativity remains high (∼0.7; Fig. 2h) even with our
best sampling effort. We know from our weighted analysis
that assortativity is closer to 0.3. Thus, there is nothing we
can do with our sampling effort to correct this: it is simply
a result of our threshold selection. Assortativity becomes
falsely exaggerated further as we increase the threshold to
0.5, where our best sampling effort shows assortativity of
one (Fig. 3h). This strongly indicates that no males or
females group together, although, this is untrue. Addition-
ally, this threshold has the highest variation in assortativ-
ity, for different levels of sampling, for the case where the
population actually has no assortativity (including an
average false positive of ∼0.2 at our best sampling effort).
Only when we increase the threshold to 0.8 (Fig. 4h) does
increased sampling effort improve our assortativity estimate
(although, the best estimate gives a slightly lower measure
than our weighted analysis). However, even here, we need a
very high sampling effort (in terms of both censuses and the
proportion sampled) before we get a realistic measure
(otherwise, we get a highly exaggerated value).
Compared to the weighted analysis, there is a clear
difference in the type of sampling effort required to get
reasonable results. Whereas, the number of censuses was
more important than the proportion sampled in the
weighted analysis, both become equally important for the
binary analysis.
Behav Ecol Sociobiol
Results: biased grouping
We tested the above condition with an individual bias
(d = 1) after all censuses with the entire population
sampled. We checked that the individual bias worked by
comparing the association value (SRI) of individuals with
the top ten highest biases, with ten randomly selected
associations, replicated over 10 runs and found that the
group allocation did assign biased individual pairings into
groups more often (a 2-tailed t test revealed p<0.001).
However, we found that individual pairing, biases make
no qualitative difference to the overall network statistics
(thus, no graphs shown).
Further analyses
In the model above, we sample a proportion of the
individuals in the population (still, always sampling an
entire group when encountered). We have also examined
the effect of basing our sampling on a proportion of
groups, rather than proportion the population. We found
similar results, except that with low proportions sampled,
the measure, generally, became less accurate, meaning that
when we base our sampling on the proportion of groups
sampled, rather than the proportion of the population
sampled, the proportion sampled becomes more important
even for our weighted measures (Electronic appendix
figure A1). We examined our findings when varying the
exponent K of the group size distribution and also with a
much higher population size (1,000 animals). In both cases,
we found no significant difference in the results (e.g.,
Electronic appendix figure A2).
We tested a situation where the population comprises
80% females and found that a small amount of
sampling is robust (Electronic appendix figure A3).
However, this is not surprising because we know (from
Fig. 1h) that there is an initial sampling bias for high
levels of assortativity. A sexually assorted population
comprising of mainly females is clearly going to give a
high level of assortativity. Thus, the bias towards high
assortativity at low levels of sampling is unlikely to provide
us with a discrepancy when the actual assortativity is high.
However, we are still unable to have confidence in the
assortment measure at low levels of sampling because, with
real populations, we would not know a priori whether our
population is highly assorted by trait (and so, would not
know if the high assortativity of our small sample is a true
representation).
Although animal group size distributions are typically
right skewed (such as our red deer group size distribution),
we also configured our model with group-sizes selected
from a normal distribution (μ=10, σ=4) to examine the
Behav Ecol Sociobiol

Fig. 2 Mean network measures

for binary networks with a
threshold of 0.2. Each data-point
is averaged over 30 replications
taken from networks each with a
different random seeds. g Shows
the mean weighted trait assorta-
tivity where there is no trait bias
(b=0) and h shows the trait
assortativity where there is a
strong trait bias (b=1). There is
no individual bias (d=0)
 Behav Ecol Sociobiol

Fig. 3 Mean network measures

for binary networks with a
threshold of 0.5. Each data-point
is averaged over 30 replications
taken from networks each with a
different random seeds.
g Shows the mean weighted trait
assortativity where there is no
trait bias (b=0) and h shows the
trait assortativity where there is
a strong trait bias (b=1). There is
no individual bias (d=0)
Behav Ecol Sociobiol

Fig. 4 Mean network measures

for binary networks with a
threshold of 0.8. Each data-point
is averaged over 30 replications
taken from networks each with a
different random seeds. g Shows
the mean weighted trait assorta-
tivity where there is no trait bias
(b=0) and h shows the trait
assortativity where there is a
strong trait bias (b=1). There is
no individual bias (d=0)

significance of the distribution. We generally found
qualitatively similar results when we used the Pareto
distribution (Electronic appendix figure A4). When group
formation was biased by sex, however, we found a much
higher level of assortment (even with the highest level of
sampling). Given that we used the same algorithm for
group allocation, this suggests that assortment by a
dichotomous trait such as sex may be more likely to occur
strongly when group sizes are normally distributed, rather
than when they are right skewed.
Discussion
We offer strong, independent support to the argument made
by Lusseau et al. (2008) advocating the use of weighted
network measures, rather than filtering data prior to
analysis. We show that the value of the threshold used to
filter association data can drastically change the resulting
network measures. Crucially, we show that different filters
have different effects depending on the level of sampling.
Researchers commonly use mid-valued thresholds to filter
network association data (see Croft et al. 2008). Often, this
value is arrived at using an intuitive approach: exploring
network plots with different threshold levels and selecting
the filter that appears to give the best core structure.
However, it is often difficult to determine which individuals
or connections are important (or which connections are
collectively important). Our comparison of binary thresh-
olded networks with weighted networks shows that for our
simulated gambit of the group associations, lower values of
the threshold are a better match to our weighted analysis.
This does not mean that lower thresholds are always going
to be the best option, but it does mean that threshold
selection may not be as simple as picking out core
structures manually. Given that weighted analyses over-
come such issues, they seem to be a sensible choice.
Filtered networks are, however, a sensible choice for the
graphical display of network structures as they can provide
researchers with a clear view of the core structure of the
network, rather than a typically visually incomprehensible
mass of weighted connections. Weighted analyses also have
the advantage of increasing data resolution without increasing
sampling effort. When we filter a network, we are discarding
data and therefore, reducing the resolution of our data,
meaning that we might need to sample for slightly longer
when performing a binary analysis. Another point to keep in
mind is that rare (how rare depends on the threshold level) or
random associations might be important (Granovetter 1974)
depending on the purpose of our network investigation. This
might particularly be the case if there are more of these
discarded connections than there are common connections.
If, for example, we are interested in disease transmission
Behav Ecol Sociobiol
through a network, then chance encounters with many
individuals might be as (or more) important than capturing
preferential association. Filtered networks are also useful for
looking for trait assortment by association type. For example,
Croft et al. (2009) studied associations in a wild population
of Trinidadian guppies, finding that strong ties were
positively assorted and weak ties were negatively assorted
by behavioral trait. This finding is only possible by thresh-
olding the network to preserve only strong associations or
only weak associations. However, when doing this, sampling
issues become more pertinent.
Our simulations show that little can be deduced from
gambit of the group networks with limited sampling.
Comparison to a randomized null model—that controls
for group size distributions, the type of sampling, and the
number of samples—would greatly help matters. However,
even when using an appropriate null model, an inadequate
level of sampling means it will only reveal whether
observed measures are higher or lower than we might
expect. We must be more cautious regarding our interpre-
tation of the absolute structure of the network. We might,
for example, find that our data show a higher positive trait
assortativity (in dichotomous traits) than is shown to be
likely by a null model where we keep the connections
constant and randomly shuffle the traits. However, in our
model, trait assortativity is artificially high with inadequate
sampling, showing positive trait assortativity where there is
no assortment. Although our null model comparison would
reveal that the assortativity is higher than expected, it does
not tell us the extent of the assortativity. Our weighted
analysis for the assorted animals, for example, shows a
much higher level of assortativity (for our dichotomous
trait) for a low level of sampling than for a (more accurate)
high level of sampling (Fig. 1g). Yet, the null model
(essentially, Fig. 1h) would, on average, show zero
assortativity for each. So, although, we would have
confidence that we have assorted data, we do not know
the extent of the assortment. For this, there is little
substitute for a good level of sampling.
We can cautiously derive several further points of advice
for ecologists sampling animal association networks—at
least with the SRI associations—from our simulation study.
We recommend that when sampling effort is limited and a
weighted analysis will be used, then effort is better invested
more heavily in the number of censuses (the number of
sampling events after fission–fusion is expected to occur)
than the proportion sampled in each census (although, this
should not be neglected). This approach can typically
resolve most sampling issues quite easily. If, however,
sampling effort is limited and a binary analysis will be
performed, then caution needs to be taken when selecting a
threshold and much more effort needs to be placed into
sampling. With a binary analysis, both the number of
Behav Ecol Sociobiol
censuses and the proportion of individuals observed in each
census become equally crucial (i.e., you need a high level
of each). Further research is needed to fully assess the
generality of our results. For example, the inclusion of a
few well-connected individuals in the network might
change the relative importance of the number of censuses
versus the proportion of the network sampled. It will also
be useful to explore the biases that different association
indexes might cause (and biases they might solve).
As animal association data collected with the gambit of
the group is weighted, it is possible for researchers who
have previously conducted a binary analysis to conduct a
weighted analysis on their current data. Our simulation
results suggest that this would be a particularly useful
exercise if the proportion of individuals sampled in each
census is estimated to have not been very high when the
data were originally collected.
Acknowledgments We would like to thank two anonymous referees,
Darren Croft, Jason Noble, and Jon Pitchford for their valuable input.
DWF is funded by NERC grant NE/E016111/1.
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