BEHAVIORAL SAMPLING IN THE FIELD: CONTINUOUS FOCAL

VERSUS FOCAL INTERVAL SAMPLING

by
LISA M. ROSE 1,2)
(Department Anthropology, Campus Box 1114, Washington University, St. Louis, MO
63130-4899, USA)
(Acc. 16-VIII-1999)

Summary
I compared data collection rates for continuous and interval focal samples during a twoyear, single-observer eld study of white-faced capuchins (Cebus capucinus) in Costa Rica.
I also compared the basic activity budgets generated by the two sampling methods, estimates
of numbers in proximity, and rates at which additional ad libitum observations could be
recorded. I collected 1238 hours of focal data (620 hr continuous, 618 hr interval). I found
focal interval sampling to be 25% more time ef cient, despite higher rate of sample loss,
partly because interval samples are easier to obtain in dif cult conditions. I found no
evidence that interval sampling provided better opportunities for ad libitum observation
than continuous sampling. Overall, the two methods yielded similar estimates of activity
budgets. However, continuous sampling resulted in somewhat higher estimates of time spent
eating, while interval data gave somewhat lower estimates of time spent foraging (looking
for or handling food items) and moving, resulting in lower estimates of foraging success.
Interval sampling also yielded slightly lower estimates of time spent vigilant. I attribute
these patterns to two major effects: (1) errors of omission (missing rare behaviors of short

1) Current address: Department of Ecology and Evolution, State University of New York,

Stony Brook, NY 11794-5245, USA; e-mail: address: lrose@life.bio.sunysb.edu

2) I thank the Area de Conservacin Guanacaste for permission to conduct eld research
at Santa Rosa, research director Roger Blanco for his assistance and support, and Linda
Fedigan for permitting me to continue work with groups LV and CP. Funding for this project
was provided by the National Science Foundation, National Geographic Society, L.S.B.
Leakey Foundation and Sigma XI. Additional support and logistical assistance was provided
by Ammonite Ltd. I thank Sue Boinski for her encouragement and advice on sampling
design, and Bob Sussman, Tab Rasmussen, Jane Phillips-Conroy, Stan Braude, Zuleyma
Tang-Martinez, Fiona Marshall and an anonymous reviewer for helpful comments on earlier
drafts of the manuscript.
c Koninklijke Brill NV, Leiden, 2000

Behaviour 137, 153-180

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LISA M. ROSE

duration) during interval samples and (2) a greater tendency toward conditional sampling
bias (under-representing behaviors due to dif cult sampling conditions such as rapid travel)
under a continuous sampling regime.

Introduction
A common concern among eld researchers is to obtain as much accurate
data as possible in a limited period of time. In a seminal discussion of
sampling methods, Altmann (1974) concluded that for most behavioral
studies, focal animal sampling will be the method of choice because of the
relative lack of bias and the wide range of questions that can be addressed.
Indeed, where study questions require identi cation of speci c individuals
e.g. questions regarding social relationships, age or sex differences, or
effects of dominance rank focal animal sampling is the only practical
choice. However, focal data (i.e. data collected on one known individual at
a time) may be recorded either continuously or at pre-determined intervals.
Two related sets of questions which I address in this study are (1) what are
the relative advantages or disadvantages of each method in terms of data
collection rates, representation of rare behaviors, and opportunities to collect
additional ad libitum (non-focal) data; and (2) do the two sampling methods
yield similar estimates of behavioral time budgets. Activity budgets derived
from the two methods have rarely been directly compared, although see
Doran (1992) and Kawanaka (1996), and relative sampling ef ciency has
not been previously examined.
The possible effects of sampling method are of critical importance when
we compare behavioral data obtained from different studies, sites, or species.
Models of primate behavioral ecology are best tested through cross-site or
cross-species comparison, but where different sampling methods are used,
we cannot be con dent that apparent differences are not simply artifacts of
differences in methodology. I address one aspect of this problem by direct
comparison of activity budgets derived from continuous focal and focal
interval samples obtained during a single study of white-faced capuchins
(Cebus capucinus ) in Costa Rica. Because all data were collected by a single
observer on the same social groups, many of the potentially confounding
factors present in cross-study comparisons are eliminated, allowing the
effects of sampling method to be systematically examined.

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155

Methods
Study species, study site and study groups
White-faced capuchins are medium-sized arboreal monkeys, adult males weighing an average
of 3.2 kg and adult females 2.3 kg (Glander et al., 1991). They are highly exible in terms of
behavior, habitat use and diet (Fragaszy et al., 1990), and have a geographic range extending
from Belize to northern Colombia. Their diet typically consists of 65-80% fruit and 2030% animal foods (including vertebrate prey), with considerable local and seasonal variation
(Freese, 1977; Chapman, 1987; Chapman & Fedigan, 1990; Janson & Boinski, 1992). Groups
are typically multi-male, multi-female with an average size of about 15 (range 3-30) and a
female-bonded form of social organization (Fedigan et al., 1996).
The study was conducted at Santa Rosa National Park in Northwestern Costa Rica
(approx. 10 50 N, 85 37 W). Established in 1971, the 108 km2 park is now part of the much
larger (1100 km2 ) Area de Conservacin Guanacaste (Janzen, 1986). It is a mosaic habitat
consisting primarily of tropical dry forest in varying stages of regeneration, patches of semievergreen forest, riparian strips, and fragments of oak forest (Janzen, 1983; Chapman, 1987).
Biological station Nancite, a restricted area within the park created to protect an important
turtle nesting beach, lies about 18 km from the central administrative area of Santa Rosa,
and also includes mangrove forest. Average annual rainfall at the park is 1528 mm (range
8880-2560 mm), almost all of which falls between mid-May and mid-December (Janzen &
Hallwachs, 1995).
Two of my three study groups, Los Valles (LV) and Cerco de Piedra (CP) had been
subjects of earlier behavioral studies (Fedigan, 1993; Rose, 1994a, b; Rose & Fedigan, 1995;
MacKinnon, 1995; Hall & Fedigan, 1997), and were well-habituated at the start of this study.
I collected focal data on groups LV and CP from January to September 1995 and January
through July 1996. Group LV ranged in size from 15 to 22 individuals, excluding 3 infants
born during the study (2-5 adult or subadult males, 4-6 adult females and 9-11 immatures)
and group CP ranged in size from 8 to 13 individuals, excluding 2 infants born during the
study (3-4 adult or subadult males, 2-4 adult females and 3-5 immatures). I habituated Nancite
group (NC) during 1995, and collected focal data collection from January through May 1996.
During the focal data collection period, the group consisted of 14 individuals, excluding one
infant born in January (2 adult and 1 subadult male, 4 adult females and 7 immatures).
Focal subjects and data collection
I collected focal samples on all adults and subadults in groups LV and CP in 1995 (N = 19
individuals), and in all three social groups in 1996 (N = 20 individuals). In total, 26
individuals were focal subjects, 13 being present in both years (Table 1). I was able to
reliably identify all individuals by physical characteristics such as scars, hair patterns and
facial features. All subjects were well habituated to my presence, and observation distances
of < 5 m were common where terrain and height in the canopy permitted. I scored behaviors
according to a prepared ethogram (Rose, 1998), with the direction and target of behaviors
indicated where applicable. All behaviors were assigned to one of ten higher-level activity
classes (Table 2). Both during and between focal sessions, I recorded ad libitum observations
(Altmann, 1974) of events such as intergroup encounters, mating, ghts, predator encounters,
mobbing, and predation by the monkeys, interrupting focal sampling if necessary.

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LISA M. ROSE

TABLE 1. Data hours for each individual and group/year by sampling

method
Focal subject

Group

Sex

BO
BQ
CA
GR
KL
SG
BU
LO
MT
MY
SK

LV
LV
LV
LV
LV
LV
LV
LV
LV
LV
LV

F
F
F
F
F
F
M
M
M
M
M

CP
CP
CP
CP
CP
CP
CP
CP

F
F
F
F
M
M
M
M

Subtotal
CF
DE
SP
TT
MB
ME
MO
Subtotal
Total

59.0
58.5
59.0
19.2
58.5
24.2
59.0
58.5
24.2
19.8
44.2

NC
NC
NC
NC
NC
NC
NC

F
F
F
F
M
M
M

1995
continuous

1995
interval

12.2
12.2
12.2
9.7
12.2
12.2
12.2
12.2
12.2
9.8
9.7

12.0
12.0
12.0
9.5
12.0
12.0
12.0
12.0
12.0
10.0
10.0

126.5

125.5

31.3
76.5
76.5
31.3
73.2
76.5
73.2
62.3

15.8
15.8
15.8
15.8
15.8
15.8
15.8
15.8

15.5
15.5
15.5
15.5
15.5
15.5
15.5
15.5

500.8

126.7

484

Subtotal
BC
LI
SE
TU
DI
NO
PI
PN

Total
data hr

36.2
36.2
36.2
36.2
36.2
36.2
36.2

253

17.3
17.3
17.3

17.3

17.3
17.3

11.0
115

23.2
23.2

19.8
23.2
19.8
16.0

124

125.2

18.2
18.2
18.2
18.2
18.2
18.2
18.2

253.2
1238

1996
continuous

250

1996
interval
17.0
17.5
17.5

17.0

17.0
17.5

13.5
117

22.0
22.0

22.0
22.0
22.0
15.0
125
18.0
18.0
18.0
18.0
18.0
18.0
18.0

127.2

126

367.3

368

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TABLE 2. Ethogram used for study of Santa Rosa capuchins

Activity class 1

Af liation

Groom

One monkey picks through or manipulates the fur of another with

hands and/or mouth.

In contact

One monkey sits or lies in body contact with another.

Huddle

Three or more monkeys sit or lie in close body contact.

Play

Two or more monkeys, usually immatures but occasionally adults,

mutually engage in sequences of spontaneous, apparently non goaloriented behaviors that include chasing, lunging, tagging, wrestling,
rough and tumble, gentle mouthing and rolling, tickling and play biting. Many components resemble those described under Aggression,
but are typically accompanied by a characteristic play face and performed in a relaxed manner without bodily tension or piloerection,
usually without accompanying vocalizations.

Double-play

Two adults simultaneously play with one or more immatures while

maintaining body contact with one another, typically in the Overlord position (Oppenheimer, 1973).

Approach

One monkey approaches to within one body length of another.

Follow

One monkey closely follows another, reaching toward or gently

touching its body.

Present for groom

One monkey sits or lies in front of, beside, or on top of another,

inviting grooming. Sometimes request is reinforced by pushing or
leaning against the body of the monkey being solicited.

Sniff

One monkey approaches and intently sniffs the hand or mouth of

another. In some cases, one monkey places a hand over the nose and
mouth of another, often with one or two ngers inserted in the nostrils
(= Hand sniff, Fedigan 1993).

Hug

One monkey embraces another, usually front-to-front.

Gargle

Rasping, throaty vocalization typically directed by infants toward

adult males, particularly the alpha male, but occasionally directed by
adult females towards males (Perry, 1997).

Activity class 2

Agonism

Open-mouthed
threat

One monkey opens the mouth so that the teeth, particularly canines,
are exposed, and stares at another. Typically performed with tense,
stiff-legged posture, laid-back ears, and often accompanied by slight
piloerection.

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LISA M. ROSE

TABLE 2. (Continued)
Stare

One monkey stares intently at another, body posture slightly tensed

and eyebrows raised. Often mouth is partly open and teeth slightly
exposed.

Chase

One monkey chases another.

Push or lunge

One monkey shoves another away with the hands, or lunges as if to

do so.

Branch shake,
break or bounce

A monkey pulls rapidly and repeatedly on a branch or small tree,

intermittently staring at or threatening another. Often accompanied
by vigorous bouncing. Branches may be broken off and dropped or
thrown toward the opponent.

Bite

One monkey bites another.

Supplant

One monkey approaches another and takes its place as the second
moves away.

Huh

Brief vocalization associated with maintaining space (Boinski &

Campbell, 1996).

Avoid

One monkey moves away as another approaches. Typically accompanied by tense body posture and nervous glances toward the approaching individual.

Fear grimace

Mouth is slightly open, teeth bared and lips stretched back in a

wide grimace. Often accompanied by quiet chattering and cowering.
(= grin, Oppenheimer, 1973).
One monkey grabs its own tail and kneads, licks or chews it rapidly.
Typically occurs after receiving aggression or during prolonged
ghts.

Chew tail

Scream

Loud screaming vocalization.

Alliance

Two or more monkeys join in an agonistic interaction against one or

more others (speci c behavior Double threat usually occurs within
this context).

Double threat

Two monkeys in body contact, either side by side or on top of

one another (= overlord, Oppenheimer, 1973) direct open-mouthed
threats to one or more others.

Activity class 3

Sexual behavior

Mild solicit

One monkey stares at another with lips protruded (= duckface,

Perry, 1997). May be accompanied by quiet sex vocalizations (low
trills, squeaks, and a warbling sound that resembles gargling) and
slight piloerection. Typically directed to a monkey of the opposite
sex.

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159

TABLE 2. (Continued)
Strong solicit

One monkey approaches another with duck face and sex vocalizations. Often accompanied by a slightly weaving gait; males are typically piloerected. Typically directed to a monkey of the opposite sex.

Courtship dance

Two individuals (typically male and female, but occasionally two females) engage in courtship display. The dance incorporates mutual
pacing, parallel movement or shadowing, oblique approaches, circling and pirouettes. Both participants have pronounced duck faces
and give intermittent and often intense sex vocalizations (see Manson et al., 1997; Perry, 1997)

Mount

One monkey mounts another and thrusts repeatedly against the genital area. Includes male-male, female-female, female-male and juvenile male-female mounts. Male-male mounts may involve apparent
intromission. Male-female mounts involving intromission are notes
as Mate.

Mate

A male mounts a females and thrusts repeatedly. Intromission is

apparent or can be reasonably inferred. Performed with duckface and
squeaking vocalizations.

Activity class 4

Other species aggression

Threat

One monkey threatens, lunges at or chases one or more individuals of

another species, including humans. May include branch breaking or
shaking.

Alliance

Two or more monkeys simultaneously threaten one or more individuals of another species while in body contact (typically double threats
in overlord or side by side position).

Mob

A group of three or more monkeys closely approach, bark at and

repeatedly threaten one or more individuals of another species,
typically a potential predator.

Activity class 5

Eat

Pick direct

Gather and consume food items (typically fruit, nuts or seeds, caterpillars) that require minimal or no manipulation prior to consumption.

Pick and process

Gather and consume food items that require considerable manipulation and processing prior to consumption (e.g. banging, rubbing
on branch, tearing and picking through matrix with hands and/or
teeth). Excludes multiple small invertebrates in stems (see Embedded Invertebrates).

Catch and eat

Catch and consume a mobile or potentially mobile invertebrate prey

that requires minimal or no consumption prior to consumption.

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LISA M. ROSE

TABLE 2. (Continued)
Continue eating

Continue eating food obtained before an intervening behavior.

Embedded
invertebrates

Extract and consume multiple small invertebrates encased

in branches, stems, or other plant matrices.

Glean invertebrates

Gather and consume multiple small invertebrates from vegetation.

Eat vertebrate prey

Consume meat or eggs.

Drink

Ingest water directly from source using hands or mouth.

Lick water

Lick water from hands, fur, or vegetation.

Eat dirt

Consume earth or crumbled rock.

Activity class 6

Forage

Visual forage

Visually inspect vegetation or food while stationary or moving

< 1 m.

Manual forage
(light)

Manipulate substrate in search of food. Includes poking, prying,

breaking small twigs, gentle pulling, removing light bark.
Excludes explore nest.

Manual forage
(heavy)

Vigorously manipulate substrate in search of food using body

strength as well as hands or teeth. Includes ripping off bark, breaking
open large branches.

Explore nest

Investigate contents of bird, squirrel, or coati nest.

Sample

Smell, poke or bite (but not consume) food (typically unripe fruit).

Army anting

Closely watch or follow army ant raiding party ushing invertebrate

prey (excludes prey captures, each noted as catch and eat).

Grab and miss

Failed attempts to catch and eat invertebrate prey.

Chase or stalk
(vertebrate) prey

Attempt to capture vertebrate prey. Includes squirrel chases, chasing

prey ushed from nests, birds, large lizards, bats.

Food interest

One monkey approaches another that has a food item and solicits
sharing by placing hand or mouth on the food, or begs with hand
outstretched (see Rose, 1997).

Other foraging
activity

Behaviors involving attempts to obtain food not included above.

Activity class 7

Movement

Move

Slow, sporadic locomotion over short distances, usually associated

with foraging.

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161

TABLE 2. (Continued)
Travel

Fast, sustained locomotion over longer distances, usually associated

with movement between distant food patches or water.

Activity class 8

Inactive

Inactive

A monkey rests, sitting or lying, and is not in body contact with

another or performing any other behavior.

Activity class 9

Scan

Vigilance

A monkey gazes intently beyond vegetation in immediate vicinity

with the head up, body stationary and slightly or conspicuously
tensed.

Scan and call

A monkey gazes beyond the immediate vegetation, intermittently

giving loud contact calls (= arrawh, Oppenheimer, 1973) and
listening intently. Typically involves lone individuals temporarily
separated from the group or responses to the calls of others.

Watch group

A monkey gazes intently toward its own social group, watching the
group as a whole rather than any speci c individual, and not engaging
in any other activity.

Activity class 10

Other

Alarm call

Brief barking vocalization (= gyrrah, Oppenheimer, 1973) given in

response to an actual or potential threat. Includes at least three distinct
types of call, to large snakes, raptors, and humans or other monkey
groups (see Perry, 1995).

Chuck

Staccato, guttural cough/bark vocalization given by adult males and

typically associated with intergroup encounters, or contact with males
that have temporarily been away from the group.

Fur rub

A monkey chews a substance and rubs it into its own fur or the fur of
others.

Leave

One monkey moves away after having been in contact with another.

Nurse

A female suckles an infant.

Caregiver call

A lactating female calls an infant to nurse (Boinski & Mitchell, 1995).

Self-groom

A monkey manipulates and picks through its own fur with hands or
mouth.

Sniff substrate

A monkey intently sniffs a substrate, often rubbing its face against it.

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LISA M. ROSE

TABLE 2. (Continued)
Share food

Share food with another individual. Includes offering food and allowing food to be taken without reprisal (tolerated scrounge; see Rose,
1997).

Startle

A monkey is startled by a sudden movement or noise, jumps abruptly

and turns toward the source, but does not give a threat or alarm call.

Trill

Vocalization associated with travel and initiation of group movement

(Boinski, 1993).

Urine wash

While urinating, a monkey places a hand under the ow and splashes

urine onto the body, usually onto a raised foot. The urine soaked foot
is typically rubbed against the opposite thigh, lower belly or genital
area.

Watch

One monkey watches another, or watches another animal or object,

without aggression or attempted predation.

Wheeze dance

A mutual display performed by adult males that includes many elements of courtship dancing, but lacks the duckface and is accompanied by a wheezing vocalization. First described by Perry (1995) in
the context of reconciliation; see also Perry, 1998.

I conducted continuous focal and focal interval sampling on alternate days, typically
spending six consecutive sampling days with each group in turn. I collected data from dawn
until dusk whenever possible. Continuous samples were ten minutes in duration, timed by
stopwatch, and followed previous protocol for this site (Fedigan, 1993; Rose, 1994a, b).
Interval samples were 30 minutes in duration. I recorded the subjects predominant activity
at the start of each session, and subsequently at the sound of an alarm set for one minute
intervals (or within a maximum of 10 seconds if the subject was temporarily obscured from
view), yielding 31 data points per session. This method is patterned after Boinski (1988) and
Fragaszy et al. (1992). During focal interval sessions, I also recorded all observed occurrences
of speci c behaviors of interest, including agonism and vigilance, but these data are not
included here. During both continuous and interval focal sessions, I recorded the number
and identity of individuals in proximity (within a 10 m radius of the focal subject) at 5-min
intervals.
I kept a cumulative tally of samples for each group and each individual to facilitate
equal sampling (except for individuals who disappeared during the course of the study),
and alternated samples between males and females as far as group composition permitted.
Visibility and group dispersion mitigated against a rigid sampling order, but I attempted to
sample each focal subject in the group before beginning the next round of samples, with
subsequent priority given to any whose tallies were short.
If I lost contact with a subject for longer than 20 seconds for continuous samples or 60
seconds for interval samples, I paused the stopwatch. If I was able to re-establish contact
within 5 minutes, I restarted the stopwatch and continued the sample, noting the duration
of the pause. If I was unable to re-establish contact within ve minutes, the sample was

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

163

scored as lost and I located a new focal subject, with priority given to those not yet included
in the current round of samples. I occasionally paused focal interval samples for longer
periods (maximum 2.5 hours) if at least the rst 10 minutes of the sample had been completed.
Permitted conditions for these prolonged pauses included torrential rain, subject temporarily
inaccessible (e.g. below dangerous cliff face), observer attacked by wasps, and priority given
to ad libitum observations of predation or other events of particular interest.
Data preparation and analyses
For continuous focal data, I rst calculated the total time (sum of all completed 10-min focal
sessions) for each focal subject in each group and year. I then calculated total time spent
engaged in each activity for each subject and generated individual time budgets (time in
min/observation min 100). Initial calculations were made on the basis of activity class,
with subsequent calculations for speci c behaviors of interest. I treated focal interval data
similarly, but used the number of interval scans on which an activity was scored/total intervals
as the basis for activity budgets. Average values across all groups and years represent data
from the 26 individual focal subjects, with results for 1995 and 1996 combined for the 13
individuals that were included in both years.
I used unpaired t -tests to compare hourly data collection rates for continuous and interval
sampling. To compare activity budgets, I rst performed chi square tests on overall activity
class distributions, based on mean values across subjects. I then used paired t -tests to compare
mean values for individual activity classes, with continuous and interval representing the
paired conditions for each focal subject. As not all variables were normally distributed, t -tests
were performed on arcsine transformed values. I also performed equivalent non-parametric
statistical tests (Wilcoxon signed ranks tests for related samples), reporting any discrepancies
in results. To determine the effects of sampling method on variance in activity classes, I
performed linear regressions of method against activity class (N = 26 individuals 2 values
= 52 cases for all groups and years). After examining the data at the level of activity class, I
extended the same procedures of paired t -tests followed by regression analysis to speci c
behaviors within activity classes found to be signi cantly affected by sampling method.
Signi cance level was set at p < 0.05, two-tailed.
Observation hours and data collection rates
During 1,886 observation hours (time spent in visual contact with at least one member of a
study group, excluding time spent habituating group NC in 1995) I collected 1,238 hours of
focal data, an average of 43.6 data hours for each subject (range 19.2-76.5 hr; Table 1). Data
hours were equally divided between continuous and interval sampling (620 and 618 hr). My
average daily data collection rate was 0.62 data hr per observation hr. Daily data collection
rates were signi cantly correlated with daily observation time, which ranged from 1 to 13 hrs
(r = 0.46, p < 0.001). Data collection rates were slightly higher for group CP (501 data
hr/727 observation hr = 0.69) than for group LV (484/753 = 0.64) or group NC (253/406 =
0.62).

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LISA M. ROSE

Time distribution of samples

The time range of focal samples was from 5.10 to 18.12 hr. Sampling was slightly biased
toward the middle of the day: morning (< 9.30) 32%, midday (9.30 to 13.30) 36%, and
afternoon (> 13.30) 32%. The distribution is similar in both data sets (continuous: morning
31%, midday 36%, afternoon 33%; interval: morning 32.5%, midday 36%, afternoon 31.5%).
Only in group NC was sampling biased more toward the morning hours, especially in
the continuous data set (continuous: morning 40%, midday 31%, afternoon 29%; interval:
morning 35.5%, midday 33.5%, afternoon 31%). This re ects the groups tendency to spend
the midday and early afternoon hours in dense mangroves, where sampling is dif cult.

Results
Continuous versus interval data collection rates
The overall rate of data collection was 25% higher for interval than for
continuous sampling (617.5 data hr/834.5 observation hr = 0.74 versus
620.5/1046 = 0.59). Overall data collection rates for interval sampling
exceeded those for continuous sampling in all groups and years, with the
difference ranging from 17% for group CP in 1995 to 36% for group NC
(Table 3). Average daily data collection rates were signi cantly higher for
TABLE 3. Productivity of continuous versus interval sampling
Group/year Continuous sampling
obs hr 1 data hr rate2
LV 1995
CP 1995
LV 1996
CP 1996
NC 1996
Overall

Interval sampling

Rate comparison

obs hr 3 data hr rate2

increase4

p5

126.5
126.7
115.0
125.2
127.2

0.55
0.58
0.61
0.71
0.54

180.0
181.5
151.5
150.5
171.0

125.5
124.0
117.0
125.0
126.0

0.70
0.68
0.77
0.83
0.74

28%
17%
28%
18%
36%

4.42 < 0.001*

2.05
0.046*
3.74
0.001*
3.34
0.002*
3.46
0.001*

1051.5 620.5

0.59

834.5

617.5

0.74

25%

7.10 < 0.001*

231.5
217.5
190.0
177.5
235.0

1 Observation hours devoted to collecting continuous samples.

2 Rate of data collection = data hours/observation hours.
3 Observation hours devoted to collecting focal interval samples.
4 Increase in focal interval data collection rates over continuous rates = (interval rate cont

rate)/cont rate 100.

5 Unpaired t -test, mean daily rates for continuous versus interval sampling, * indicates
p < 0.05, 2-tailed.

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

165

interval data than for continuous data (0.72 versus 0.57, t = 7.10, p < 0.001,
N = 213 days), and signi cantly higher in each group and year (Table 3).
Because data collection rates were correlated with observation hours,
I examined rates for full days only (minimum observation time 11 hr,
N = 114 days). Overall data collection rates for full days were 24% higher
for interval sampling (483 data hr/650.5 observation hr = 0.74) than for
continuous sampling (366/6121.5 = 0.60), with differences ranging from
13% for CP in 1995 to 35% for NC in 1996. Average full day data collection
rates were signi cantly higher for interval than for continuous data (0.73
versus 0.61, t = 4.91, p < 0.001).
Rates of sample loss
In total, 99 continuous samples and 45 interval samples were lost (subject
could not be relocated within the required parameters), and 25 interval
samples were paused for more than 5 minutes. Overall, 2.6% of attempted
continuous samples were lost, compared with 3.5% of attempted interval
samples. The rate of sample loss was higher for interval than for continuous
sampling in all groups and years, except for group CP in 1996. If paused
interval samples are included with lost samples, the rate of interval sample
loss increases to 5.5%, double the rate for continuous sampling. Again, the
pattern of higher losses for interval samples was consistent in all groups and
years except for group CP in 1996. Two-thirds of sample losses occurred
during rapid travel, and this was the most common condition for sample loss
in every group/year.
Number of observations
The continuous focal data set included 56,262 observations, with a mean duration of 0.66 min (40 s). The focal interval data set included 38,285 observations (one-minute interval points), excluding all-occurrences observations.
More behaviors have fewer focal observations than focal subjects (N = 26)
in the interval data set than in the continuous data set (interval 36/74 = 49%;
continuous 20/74 = 27%). The 18 behaviors that are uncommon by this standard in the interval data set but not in the continuous data set tend to be brief
(average 13.5 s), with the greatest under-representation appearing in behaviors that typically occupy only a few seconds: approach, leave, present
for groom, avoid, supplant, and trill (Table 4).

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LISA M. ROSE

TABLE 4. Behaviors under-represented during interval samples 1

Continuous
observations

Interval
observations

Duration2
(s)

1194
184
51
32
30

5
10
21
14
12

3
4
12
16
51

154
86
54
42

3
1
3
6

5
3
13
11

Sexual behavior
Mount

43

24

30

Aggression to other species

Alliance

57

20

30

Forage
Food interest
Grab and miss
Other foraging activity

88
59
28

1
1
4

9
7
15

928
491
51
29

0
1
2
2

2
3
17
13

Activity class/behavior
Af liation
Approach
Present for groom
Sniff
Hug
Double play
Agonism
Avoid
Supplant
Scream
Fear face

Other behaviors
Leave
Trill
Alarm call
Sniff substrate

1 Scored < 26 times during interval samples and > 26 during continuous samples.
2 Average duration based on continuous samples.

Activity budgets
Based on averages across focal subjects in all groups and years, the overall
distribution of time (for continuous samples) and scans (for interval samples)
among activity classes does not differ signi cantly between the two sampling
methods (X 2 = 1.45, p = 0.229, one tailed). However, among individual
activity classes, mean values for Eat and Scan are signi cantly higher in

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

167

the continuous data set, and the mean for Forage is signi cantly higher in
the interval data set (Table 5). Wilcoxon tests give the same results, except
that the difference in Movement (interval > continuous) is also signi cant
(z = 1.97, p = 0.049). The differences in Scan and Movement would not
reach signi cance if results were adjusted to re ect multiple tests (N = 10),
but as the null hypothesis in each case is that sampling method does not affect
estimates of activity budgets, the results provided here are appropriately
conservative.
Within the ve individual group/year data sets, interval values for activity
class Forage are signi cantly higher than continuous values in all cases
except for NC 1996. Continuous values are signi cantly higher for Eat and
lower for Movement in LV 1995 and CP 1996, with a similar trend in the
remaining three group/years. There are no signi cant differences in Scan
in any group/year, but there is a consistent trend toward slightly higher mean
values in the continuous data set.
On average, sampling method explains 5.7% of individual variance in
activity classes, with only estimates of time devoted to activity classes
Eat and Forage being signi cantly affected (Table 6). Sampling method
explains 13.6% of the variance in Eat (F = 7.89, p = 0.007) and 32.4%
of variance in Forage (F = 23.93, p < 0.001).
Activity classes signi cantly affected by sampling method
Movement
Movement has two components: move (slow, sporadic movement over
short distances) and travel (fast, sustained movement over longer distances). The source of higher values for Movement in the interval data set
is travel (8.5 versus 6.7% in the continuous data set, t = 4.26, p < 0.001).
This represents a difference of about 10 mins per day for the average individual (interval 60 mins per day spent traveling versus continuous 50 mins).
The difference in move is smaller and in the opposite direction (interval
11.4 versus continuous 12.4%, t = 4.70, p < 0.001). Regression tests indicate that sampling method accounts for 26.8% of the variance in travel
(F = 18.30, p < 0.001), and 6.1% of the variance in move (F = 3.26,
p = 0.077).
The pattern of higher interval values for travel, lower interval values for
move, and larger differences in travel than in move is consistent across

7.6%
0.49%
0.16%
0.26%
40.0%
12.2%
19.2%
16.1%
1.5%
2.4%

8.3%
52.2%

Af liation
Agonism
Sexual
Other species
Eat
Forage
Movement
Inactive
Scan
Other

All social4
All forage5

9.0%
51.1%

8.2%
0.56%
0.26%
0.25%
35.9%
15.3%
19.9%
16.3%
1.2%
2.2%

Interval
mean

5.3%
5.1%

5.1%
0.41%
0.17%
0.23%
5.5%
2.1%
3.0%
5.9%
1.9%
0.8%

Continuous
SD

5.9%
5.0%

5.7%
0.52%
0.42%
0.26%
5.2%
2.5%
2.0%
5.6%
1.4%
0.8%

Interval
SD

1.31
1.15

1.07
0.83
1.37
1.16
4.53
7.98
1.65
0.20
2.17
1.02

t -test
t

0.202
0.260

0.279
0.417
0.184
0.259
< 0.001*
< 0.001*
0.111
0.841
0.039*
0.319

t -test
signi cance2

X 2 test of overall distribution, continuous versus interval sampling: X 2 = 1.45, p = 0.229 (one tailed).
1 See Table 2 for description of activity classes; values are mean % time/intervals for N = 26 individuals.
2 Paired t -test, continuous versus interval means, arcsine transformed data, * indicates p < 0.05, 2-tailed.
3 Wilcoxon signed-ranks matched pairs test, * indicates p < 0.05, 2-tailed.
4 Af liation + Agonism + Sexual behavior.
5 Eat plus Forage.

Continuous
mean

1.19
0.37

1.16
0.17
0.43
1.08
3.47
4.36
1.97
0.17
2.22
0.42

Wilcoxon
z

TABLE 5. Activity budget comparisons, continuous versus interval sampling

Activity class1

0.125
0.370

0.248
0.869
0.664
0.280
0.001*
< 0.001*
0.049*
0.869
0.026*
0.675

Wilcoxon
signi cance3

168
LISA M. ROSE

169

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

TABLE 6. Proportion of activity class variance accounted for by sampling

method
Activity class

Variance1
in category

% explained
by method2

ANOVA
F -value

Signi cance3

Af liation
Agonism
Sexual
Other species
Eat
Forage
Movement
Inactive
Scan
Other

0.290%
0.002%
0.001%
0.001%
0.325%
0.076%
0.064%
0.309%
0.028%
0.006%

0.3%
0.6%
2.7%
2.6%
13.6%
32.4%
2.4%
0.0%
0.9%
1.4%

0.13
0.31
1.40
1.32
7.89
23.93
1.24
0.01
0.48
0.72

0.724
0.579
0.242
0.256
0.007*
< 0.001 *
0.271
0.913
0.493
0.402

All social
All forage

0.310%
0.256%

0.4%
1.1%

0.21
0.58

0.648
0.449

1 Total variance in continuous and interval data (N = 26 individuals 2 values = 52 values).

2 r 2 value from linear regression of sampling method against activity class. Constant =

sampling method, dependent variable = activity class.

3 * Indicates p < 0.05, 2-tailed.

groups and years, although not all individual differences are signi cant. I
considered the possibility that higher values for travel in the interval data
set may be due poor discrimination from move. The two behaviors are
typically distinct, but they do form a continuum within which rapid foraging
movement can grade into slow travel. However, there is no consistent trend
for groups or individuals with higher values for travel to have lower values
for move, and no correlation between individual differences in travel and
individual differences in move (r = 0.07, p = 0.721, N = 26).
Scan
Scan has three components: scan and lost call and watch group (behaviors primarily directed towards the social group) and vigilance (primarily
directed outside the social group). The major component, vigilance, accounts for 80% of Scan in both data sets. Across all groups and years,
mean values for vigilance are higher for continuous than for interval data
(1.23 versus 0.96%, t = 2.85, p = 0.009). There are no signi cant differences in scan and call (0.10 versus 0.11%, t = 0.12, p = 0.904) or watch

170

LISA M. ROSE

group (0.22 versus 0.16%, t = 1.76, p = 0.090). Within group/year data

sets, there is a trend for vigilance to have higher values in the continuous
than in the interval data set, although the differences are signi cant only in
LV 1995 and CP 1995. Across groups and years, sampling method explains
1.1% of the variance in vigilance (F = 0.57, p = 0.453), 0.5% of the variance in watch group (F = 0.27, p = 0.606) and < 0.1% of the variance in
scan and call (F = 0.01, p = 0.917).
Eat and forage
Paired t-tests, Wilcoxon tests, and regression tests all indicate that Eat
and Forage are signi cantly affected by sampling method. Mean values
for Eat are 4.2% higher in the continuous data set than in the interval
data set (40 versus 35.9%), and values for Forage are 3.1% lower (12.2
versus 15.3%; see Table 4). It is noteworthy that total foraging time (Eat
plus Forage) does not differ signi cantly between sampling methods
(continuous 51.6%, interval 50.8%, t = 0.91, p = 0.370), a pattern that
is consistent in all groups and years. However, foraging success (eat/eat
+ forage + movement) is consistently higher when based on continuous
data than when based on interval data. Across all groups and years, average
foraging success based on continuous data is 56.0%, compared with 50.3%
based on interval data (paired t-test: t = 5.82, p < 0.001). Within
group/year data sets, differences in foraging success (continuous > interval)
range from 2.4% for NC 1996 to 8.8% for CP 1996.
Within activity class Eat, values for pick direct and pick and process
are signi cantly higher in the continuous data set than in the interval data
set (Table 7). Within activity class Forage, values for visual forage and
manual forage are signi cantly higher in the interval data set, with the
major difference being in visual forage (interval 12.6 versus continuous
9.8%, t = 8.12, p < 0.001). Paired t-tests and Wilcoxon tests give the
same results. For the average individual, this represents a difference of about
20 mins per day spent in visual foraging (90 mins interval versus 70 mins
continuous).
Within individual group/year data sets, there is a consistent trend for
visual forage to have higher values in the interval data set than in the
continuous data set. Results for pick direct, pick and process and manual
forage are less consistent, but there is a general trend toward higher
estimates of manual forage in the interval data and lower estimates of

9.8%
1.7%
0.7%

Activity class Forage

Visual forage
Manual forage
Other forage5

12.6%
2.0%
0.7%

23.0%
3.2%
1.9%
4.4%
2.8%
0.7%

Interval
mean

1.7%
0.5%
0.7%

4.9%
2.4%
0.8%
2.9%
0.7%
0.8%

Continuous
SD

2.1%
0.6%
0.7%

5.8%
2.4%
1.1%
2.4%
0.7%
0.7%

Interval
SD

8.12
2.38
0.21

3.12
2.28
1.50
0.94
0.41
0.64

t -test
t

< 0.001*
< 0.001*
0.834

0.005*
0.031*
0.146
0.358
0.687
0.527

t -test
signi cance2

1 See Table 2 for description of foraging activities; values are mean % time/intervals for N = 26 individuals.
2 Paired t -test, continuous versus interval means, arcsine transformed data. * indicates p < 0.05, 2-tailed.
3 Wilcoxon signed-ranks matched pairs test, * indicates p < 0.05, 2-tailed.
4 Other eat includes eat meat or eggs, eat dirt, drink or lick water.
5 Other forage includes sample, sniff, grab and miss, chase or stalk prey.

25.8%
4.3%
1.6%
4.7%
2.8%
0.8%

Continuous
mean

Activity class Eat

Pick direct
Pick and process
Glean inverts
Embedded inverts
Catch inverts
Other eat4

Foraging activity1

TABLE 7. Foraging time budgets: continuous versus interval sampling

4.31
2.30
0.27

2.93
2.17
1.46
0.60
0.57
0.55

Wilcoxon
z

< 0.001*
0.022*
0.790

0.003*
0.030*
0.144
0.551
0.568
0.585

Wilcoxon
signi cance3

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

171

172

LISA M. ROSE

TABLE 8. Proportion of variance in foraging activities explained by sampling method

Foraging activity

Variance2
in category

% explained
by method2

ANOVA
F -value

Signi cance

Activity class Eat

Pick direct
Pick and process
Glean inverts
Embedded inverts
Catch inverts
Other eat

0.31%
0.59%
0.09%
0.07%
0.04%
0.01%

6.8%
5.5%
2.2%
0.3%
0.2%
0.8%

3.63
2.91
1.14
0.16
0.09
0.42

0.062
0.094
0.291
0.688
0.767
0.519

Activity class Forage

Visual forage
Manual forage
Other forage

0.05%
0.03%
0.04%

37.2%
7.9%
0.1%

29.56
4.28
0.03

< 0.001 *
0.044 *
0.857

1 Total variance in continuous and interval data (N = 26 individuals 2 values = 52 values).

2 r 2 value from linear regression of sampling method against foraging activity. Constant =
sampling method, dependent variable = foraging behavior. * indicates p < 0.05, 2-tailed.

pick direct in the continuous data. Regression of sampling method against

speci c foraging activities indicates that on average, sampling method
explains 6.8% of observed variance, and only manual forage and visual
forage are signi cantly affected (Table 8). Sampling method explains 7.9%
of the variance in manual forage (F = 4.28, p = 0.044) and 37.2% of
variance in visual forage (F = 29.56, p < 0.001).
It is possible that high values for visual forage in the interval data set
are due to poor discrimination from vigilance, as short vigilance bouts can
sometimes be dif cult to distinguish from visual foraging. However, there
is no correlation between vigilance and visual forage in either data set
(continuous r = 0.03, p = 0.856; interval r = 0.15, p = 0.350, N = 26
individuals), and no correlation between differences in visual forage and
vigilance (r = 0.05, p = 0.803). Also, the difference in vigilance
(continuous interval = 0.27% of observed time) is far too low to account
for the overall mean difference in visual forage (interval continuous =
2.8% of observed time).

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

173

Numbers in proximity
The continuous focal data set included 11,169 proximity scans (3 per 10min session), and the focal interval data included 8,645 proximity scans
(7 per 30-min session). I was unable to obtain proximity data on 3.2% of
continuous scans and 3.7% of interval scans, typically because of restricted
visibility. The average proportion of scans with no individual in proximity to
the focal subject was signi cantly higher during interval sessions than during
continuous sessions (41.5 versus 36.6%, paired t-test t = 3.74, p = 0.001;
Wilcoxon signed-ranks matched pairs test z = 3.65, p < 0.001). The
average number of individuals in proximity was slightly but not signi cantly
lower during interval sessions than during continuous sessions (1.4 versus
1.3, t = 1.64, p = 0.114; z = 1.79, p = 0.073).
Ad libitum observations
To compare my ability to record events of interest within the group under
continuous and interval sampling regimes, I grouped ad libitum data collected during and between focal sessions into ve major categories (Table 9).
In some cases, these observations were subsequently represented in the focal
data (e.g. predator mobbing or large ght in which a focal subject participated), but most involved individuals other than the focal subject. Hourly
rates of ad libitum observation are similar in four categories ( ghts, predator or other species encounters, vertebrate predation and sexual behavior) but
food sharing was observed more often during or between continuous samples
than during or between interval samples. However, most of the difference in
food sharing can be traced to two major predation events (a currassow nest
raid and a coati nest raid) that occurred on one continuous sampling day
in group NC. Overall, rates of ad libitum event scoring were similar during continuous and interval samples (1.14 versus 1.06 events/hr), and almost
identical when I included events observed between samples (continuous 0.95
versus interval 0.93 events/hr).

Discussion
This study clearly indicates that interval sampling is a more ef cient method
of data collection than continuous sampling in terms of the amount of focal

41

Food sharing/theft
1.14

0.07

0.08

0.13

0.20

0.66

649

12

47

79

112

399

1.06

0.02

0.08

0.13

0.18

0.65

992

71

60

104

185

572

0.95

0.07

0.06

0.10

0.18

0.54

Continuous
rate/hr

1 Hourly rates based on number of observations during 620 hrs continuous data, 618 hrs interval data.
2 Hourly rates based on number observations during 1051 continuous observation time, 835 hrs interval observation time.
3 Excludes threats to human observers and interspeci c aggression during predation attempts.

705

49

Sexual behavior
(courtship, mounts, mating)

All events

78

126

Predators/other species3
(mob or other aggression)

Vertebrate predation
(attempts and events)

411

Aonistic interactions
(evoking audible screams)

Interval
rate/hr

Continuous
no.

Interval
no.

Continuous
no.

Continuous
rate/hr

During and between sessions 2

During sessions 1

TABLE 9. Ad libitum observations: continuous versus interval sampling

776

18

61

107

135

455

Interval
no.

0.93

0.02

0.07

0.13

0.16

0.55

Interval
rate/hr

174
LISA M. ROSE

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

175

data that can be obtained in a given time. Interval sampling requires about
25% less observation time than continuous sampling to obtain similar data
yields. This is an important consideration when eld time is limited, as
it generally is. Part of the difference is probably due to extended session
length, which reduces the number of starts (48 for 10-min continuous
sessions versus 16 for 30-min interval sessions for a typical 8-hr data day),
and the associated time spent locating new focal individuals and setting up
samples. However, this advantage is partly offset by the greater risk of losing
samples during the longer interval sessions. Relative ease of data collection
is probably a more important factor, especially where visibility is poor or
terrain is dif cult. It is often more feasible to collect data at intervals than to
record data continuously when observation conditions are unfavorable, and
observer fatigue is also reduced by some relaxation of the need to continually
watch the focal subject, monitor time, and record data.
The greater ef ciency of interval sampling under dif cult conditions
is con rmed by comparing data collection rates across the study groups.
The most striking difference between data collection rates (36%) is in
group NC, which has a high proportion of old forest and dense mangroves
where visibility is poor. The least difference (17-18%) is in group CP,
which has a more open and virtually at home range, and many areas of
young forest where visibility is excellent. The difference is intermediate
(28%) in LV group, where terrain and visibility are highly variable. Group
contrasts in sampling conditions are con rmed by patterns of sample loss and
comparative data collection rates. The proportions of sample loss associated
with poor visibility or rough terrain mirror the order of differences between
sampling methods (NC 33% > LV 17% > CP 10%), and data collection
rates follow the reverse order (CP 69% > LV 64% > NC 62%).
Although continuous sampling is less time-effective than interval sampling, it offers an advantage in yielding a larger number of observations for
most behaviors. Continuous data also provide a full record of an individuals activity during the sample, including exact durations and sequences of
events, and there is a much greater likelihood that infrequent or brief behaviors will be represented (see also Doran, 1992, for similar remarks on
locomotor sampling). Missing events of very short duration may have little
effect on overall time budgets, but can seriously limit analyses of speci c
activities. For example, in this study, interval samples provided very few instances of approach, leave or present for groom behaviors that were

176

LISA M. ROSE

well-represented in the continuous data set. Of course such behaviors can

be recorded on an all-occurrences basis if necessary, but this adds time and
complexity to data management, and one must anticipate the need before
beginning data collection.
A potential advantage of interval sampling over continuous sampling is
that it allows an observer to take greater note of events in the group as
a whole, and to collect other types of data during focal sessions (Doran,
1992; Fragaszy et al., 1992). However, in this study, interval sampling did
not yield more ad libitum observations than continuous sampling the
rates were remarkably similar. This is probably because my attention was
typically focused on the focal subject throughout interval sessions in order
to avoid losing contact or missing all occurrence behaviors. In addition,
capuchin groups are often widely dispersed, and many events scored ad
libitum were based on audible cues (screams, alarm calls, barks) that could be
noted equally well during continuous or interval sessions. Where events were
visible without my moving more than a few meters from a focal subject, I was
often able to record more detailed data during interval sessions than during
continuous sessions. However, my protocol of pausing or ending sessions in
order to focus closely on major events makes it dif cult to assess the degree
of difference. Based on the limited analyses suitable for this study, I found no
quantitative evidence that interval sampling permits more ef cient collection
of ad libitum data than continuous sampling.
Overall, continuous and interval sampling provide very similar estimates
of activity budgets. The major differences are in visual forage and travel,
which both have higher values in the interval data set. To place these
differences in real time perspective, continuous sampling indicates that the
average individual spends about 70 min each day visual foraging (9.8% of
12 hr) and about 50 min traveling (6.7% of 12 hr). Interval sampling indicates
about 90 min visual foraging (12.6% of 12 hr) and about 60 min traveling
(8.5% of 12 hr). The discrepancies may be partly due to poor discrimination
from similar behaviors ( travel and move; visual forage and vigilance)
on interval scans. Fragaszy et al. (1992) found that group scans gave less
stringent discrimination of activities than focal interval scans, and a similar
problem may affect focal interval versus continuous focal observations.
However, the differences in move and especially vigilance are insuf cient
to account for the differences in travel and visual forage, and there
is little evidence of compensating differences in either pair of behaviors

CONTINUOUS FOCAL VERSUS FOCAL INTERVAL SAMPLING

177

at the individual or group level. While this does not rule out differential
discrimination between sampling methods, it does suggest that differences
in discrimination are not systematic and that the effects are relatively minor.
I propose that the small differences in activity budgets derived from continuous and interval sampling methods arise from two basic causes: errors of
omission (underestimating infrequent or brief activities in the interval data)
and conditional sampling bias (under-sampling activities in dif cult conditions or peripheral locations in the continuous data). Errors of omission include missing short bouts of vigilance or a subject quickly obtaining and consuming a small food item that requires little or no processing. An associated
effect is that extremely common behaviors such as visual forage become
over-represented in the interval data, due to cumulative under-representation
of various less common activities. Conditional sampling bias offers the best
explanation for the greater representation of travel in the interval data. Because it is easier to collect interval data than continuous data on a rapidly
moving focal subject, one is more likely to undertake and less likely to lose
an interval sample than a continuous sample when the subject is traveling. In
addition (all other things being equal) there is a greater chance that a 30-min
sample will include some rapid travel than a 10-min sample. This can affect
other aspects of the focal data. For example, travel often involves one or a
few individuals moving away from the group, or the entire group dispersing widely to look for new food patches. Samples under these conditions
will tend to include more foraging than eating and have fewer individuals in
proximity, a pattern found in my interval versus continuous data sets.
A related aspect of conditional sampling bias is our natural tendency to
sample focal subjects that are relatively easy to nd. For example, if sample
tallies are roughly even and most group members are feeding in or around
a large g tree, one is more likely to sample one of these individuals than
to search outlying areas for a potential subject. Similarly, one is more likely
to sample individuals that are visible and accessible rather than those that
are hidden from view or in dif cult to reach locations. These tendencies
will be particularly marked if one risks losing the entire group by moving
away to search for outliers, as is often the case with rapidly-moving arboreal
subjects. The bias toward easier subjects is less likely to affect data from
interval than from continuous sampling. This is partly due to the relative
ease of data collection, but also because longer sessions are more likely to
include peripheral or dif cult to follow activities, and a greater sensitivity to

178

LISA M. ROSE

unequal data hours promotes earlier (and probably more rigorous) searches
for missing subjects. Conditional sampling bias could probably be reduced
by using a more stringent sampling protocol than I used in this study. As
in the collection of all-occurrences data, there is a trade-off involved. At
Santa Rosa, a fully random or predetermined focal sampling order would
almost certainly increase search time and reduce sampling ef ciency to
an unacceptable level, especially where visibility is restricted and group
dispersion is high. However, protocols that approximate random selection
(Boinski, 1988) or prohibit sampling individuals that have recently interacted
(Perry, 1996) offer a reasonable compromise.
An important implication of conditional sampling bias is that continuous focal sampling is not necessarily the ideal standard against which other
sampling methods should be measured. It is true that continuous sampling
provides a fuller and richer data set, and this is clearly the method of choice
when a complete record of behavioral sequences or social interactions is required. However, the belief that continuous focal sampling will always provide the best, most real data in return for the more arduous work involved
may be ill founded. For example, in this study, although continuous sampling
probably provided the better estimate of time spent vigilant, interval sampling probably provided the better estimate of time spent traveling, and the
two methods provided very similar estimates of social and rest time. As others have noted previously (Fragaszy et al., 1992; Doran, 1992; Kawanaka,
1996), the best sampling method depends on the nature of the study, eld
conditions, time available, and above all the questions of interest. Often a
mixed sampling regime is the most useful, because different questions can
be addressed and allowances made for the different biases involved.

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