Anim. Behav.

, 1982, 30, 327-338

A FIELD STUDY OF THE HEADBOB DISPLAYS OF MALE GREEN

IGUANAS (IGUANA IGUANA): VARIATION IN FORM AND CONTEXT
BY BEVERLY DUGAN*

Department of Psychology, The University of Tennessee, Knoxville, TN 37916

Abstract. Headbob displays play an important role in the social behaviour of male green iguanas
(Iguana iguana). Four types of headbobs were identified, and the variability in form and context were
determined from quantitative analyses of displays filmed in the field. The Roll is a conspicuous advertisement display. The Shudder, the most variable in form, occurred during close contact between males and
females. The Roll-shudder is intermediate between the Roll and Shudder in its form and use. The
Signature Bob, the most stereotyped display in form, punctuates much activity and is the most variable
in contextual use. This display encodes species identity, and possibly individual identity. The conspicuous nature of the headbob displays was enhanced by the intensified colour of territorial males, and
by the selection of highly visible display posts. This system of visual displays is viewed as the product of
related sets of constraints, including the iguanas' physical capacities, the forest environment, and the
social environment.
(Distel & Veazey, 1982), although data on
variability in form and context have not been
reported. The observations of iguana display
behaviour reported here are part of a more
general field study of the social behaviour
of adult iguanas (Dugan 1980; 1982). The
objectives in this report are (1) to describe the
(headbob) display repertoire of L iguana; (2) to
identify the situations in which each type of
display is performed in the field; (3) to quantify
the variability of each display type; and (4) to
relate the variation in form and context of each
display type to the iguana's natural history.

Central to the study of animal communication

is the idea that the structure of a display represents the optimum form for transmitting particular types of information in a particular
environment. One aspect of display form that
has received much attention is variability. Many
investigators have sought correlations among
the variability of a display, its contextual use or
information content, and the species' social and
ecological environment in an attempt to understand the functions of relative differences in
variability (see reviews in Barlow 1977; Marler
1977). The objective of the present investigation
was to apply this approach in a field study of
the headbob displays of the green iguana,
Iguana iguana (Sauria, Iguanidae).
The most thoroughly studied displays among
lizards in the family Iguanidae are the headbob
displays, vertical head and/or body movements
that are usually accompanied by extension of
the dewlap, a flap of skin under the throat. Most
iguanids studied perform a display termed the
'signature bob' (Stamps & Barlow 1973): it is
stereotyped in form at the species, population
and/or individual level but occurs in a variety
of contexts such as territory advertisement, threat
and courtship. At least one other display in
addition to the signature bob has been described
for the majority of species investigated (see
reviews in Jenssen 1977; Ferguson 1977).
Iguanas have been observed to perform headbob displays in both the field (Mtiller 1972;
Lazelle 1973; Hazlett 1980) and the laboratory
*Address correspondence to: BeverlyDugan, Route 15,
Box 367, Gray, TN 37615, USA.

Methods

The social behaviour of L iguana was studied

from October 1977 to January 1979 on Flamenco
Island, a 13.5 ha island (maximum elevation
75 m) at the Pacific entrance to the Panama
Canal. The annual climate is marked by distinct
wet (May through November) and dry (December through April) seasons. Observations were
confined to a 5.5 ha area on the northwest side
of the island. The study area contained 200 to
300 iguanas (Dugan 1980).
Direct observation and filming were employed.
The iguanas were habituated to an observer
sitting quietly within a minimum distance of
about 20 m. I did not detect any differences in
behaviour observed at this range and that
observed with a telescope from 200 m and more.
During 15-min observation sessions, verbal
accounts of the behaviour of a focal animal
(Altmann 1974) were recorded with a cassette
tape recorder. The iguanas were filmed with a
327

328

ANIMAL

BEHAVIOUR,

Nizo S-80 Super 8 mm movie camera equipped

with a zoom lens and set at an advance speed
of 18 frames/s. Occasionally it was possible to
film and conduct focal animal samples simultaneously; however, during periods of intense
activity this was not possible.
Displays were filmed between October and
January, a period during which display rates
were high. Although displays were given by all,
large males displayed much more frequently
than other members of the population (see
below). Thus, the display analysis is based on
observations of large males only.
Display-action-pattern (DAP) graphs (Carpenter & Grubitz 1961) were generated from
frame-by-frame analysis of filmed headbobs.
The location of the tip of the snout in each
frame was plotted on graph paper. Time in
seconds was plotted on the X-axis; the Y-axis
represented the amplitude of each vertical head
movement. Since camera-to-subject distance
varied from 25 to 60 m, it was not possible to
measure absolute amplitude. Hence a measure
of relative amplitude was used to compare amplitudes within a display performance; the high
and low head positions were expressed as a
proportion of the highest in each display. Only
those displays in which the animal was roughly
lateral to the camera, and in which snout
position could be clearly seen in each frame,
were graphed.
Two measures of context, 'distance' and
'social', were recorded for each display. Distance
context refers to the distance between a displaying male and the nearest female(s). Estimates
of distances were classified into one of four
categories: (a) 0.0 to 0.5 m; (b) 0.5 to 1.5 m;
(c) 1.5 to 3.0 m; (d) greater than 3 m. Social
context categories were as follows: (1) Solo,
the male was the only occupant of the tree;
(2) Not Directed, other iguanas were present in
the same tree but at a distance of greater than
1 m, and the displaying iguana was not interacting with another individual; (3) General
Social, a male and female(s) were within 1 m of
one another, but no obvious interaction was
occurring; (4) Male-Female Interaction, an actresponse-act sequence occurred between an
initiator of the interaction and the respondent.
Distance and social context categories are not
mutually exclusive. Distance category 'd' applied
when the tree was occupied by females that
were more than 3 m from the male, as well as
when the male was the only occupant of the
tree. Thus, this category at times overlapped the

30,

social category Solo. However, social context

categories contain important information about
the nature of the momentary relationship
between the male and female that is lacking in
the distance categories. For example, males and
females basking at close distances typically did
not interact, even though physical contact was
maintained for longer than 15 min. However,
most interactions did occur within distances of
1.5 m. Although it was desirable to distinguish
among such situations, small sample size did
not permit the use of one system of independent
categories. Thus, all data were initially analysed
according to distance. If this analysis suggested
that consideration of social context would provide more information, then these categories
were explored.
Iguanas headbobbed during male-male interactions. Agonistic encounters usually involved
males of different sizes. These encounters were
brief, consisting of lunges or chases by the
larger male, and a rapid retreat by the smaller.
The larger male then moved rapidly throughout
the tree, pausing frequently to display. In interactions between males of equal size, an exchange
of headbob displays accompanied by lateral
body compression, as described in other iguanid
lizards (Carpenter 1967), might continue for as
long as 1.5 h. The terminal event in these encounters, the retreat of one lizard, was occasionally preceded by a snout-to-snout pushing
battle. Such encounters were extremely rare;
only three were observed in more than 1000 h
of observation. Due to its rarity, this context
was not induded in the analysis. Thus the
function of headbob displays in the challenge
phase of an agonistic encounter is not addressed
in this report.
Five types of head movements were identified:
the Head Jerk, the Roll, the Shudder, the
Roll-shudder, and the Signature Bob. Filmed
Head Jerks were too few in number to permit
quantitative analysis of form and context. Because the movements of the Roll, Shudder and
Roll-shudder were of such low amplitude,
subject-to-camera distance precluded detailed
treatment of the form of these displays. Only
overall durations and contextual use of the
three will be discussed. Ten components of the
Signature Bob were measured, and variations in
duration and amplitude measures, as well as
contextual use, were analysed.
Since all displays were filmed in the field, it
was not possible to have equal numbers of observations in each classification category. There-

DUGAN: FIELD STUDY OF IGUANA HEADBOB DISPLAYS

fore, the General Linear Models Procedure for
unbalanced designs (Statistical Analysis System,
Barr et al. 1976) was used for analyses of
variance. Other statistical methods and references are detailed in the following sections as
appropriate.
Results
Behavioural Observations

The following summary of the ecology and

social behaviour of the iguanas on Flamenco
provides a background within which to interpret
the use of the headbob displays. More detailed
accounts can be found in Dugan 1980, and
1982.
Iguanas are sexually dimorphic in body size,
shape, and coloration. Males have larger heads,
longer spines on the dorsal crest, larger femoral
pores, and a larger dewlap. Most females and
small male iguanas are green; larger males are
dull grey, gold or tan. The male's colour intensifies during the breeding season, when the head,
forelegs, and/or body of most large males are
bright gold or red-orange. Secondary sex
characteristics are most well-developed in males
larger than 36 cm body length.
During the non-breeding months of February
to October, the home ranges of all sex and size
classes overlapped in both space and time. The
largest males confined their movements to an
average area of 0.08 ha; the home ranges of
smaller males and females were roughly three
times that size.
In both 1977 and 1978 mating activity began
in the last half of October with increasing rates
of display by large males. Courtship began in
early November. Females visited and courted
several males, and less frequently, males left
their home tree to court nearby females. By late
November, most large males had established a
territory in a tall, conspicuous tree or group of
trees. Territorial males alternated periods of rest
with display bouts (non-directed displays given
while basking or resting), courtship bouts, and
territory patrols. A mating territory contained
from one to four resident females. Each female
was courted for at least two weeks before
becoming receptive. Copulation occurred during
the first six weeks of the dry season, from early
December through mid-January.
Head Movements

1. Head Jerk. The behaviour consists of a

variable number of abrupt up and down movements of the head (Fig. 1A). This movement was

329

performed by individuals in exposed locations

(nesting females, animals basking on the
ground) and in response to the approach of a
human or (less frequently) another iguana. Rand
(1968) noted that this simple bob is performed
by nesting iguanas that seem to be nervous, and
argued that it functions as an aid to vision
rather than as a social signal. My observations
are consistent with Rand's, and as the Head Jerk
was not associated with any particular social
situation(s), it was not considered to be a
display.
2. Shudder. The Shudder corresponds to Distel
& Veazey's (1982) 'vibratory head nodding'.
This bob type consists of very low amplitude
vertical head movements that are much more
rapid than those of the Head Jerk (Fig. 1B).
Head position is variable, but the head is most
often parallel to the substrate. Neck movement
is more reduced in this bob than in the others.
Occasionally, the animal raises the head or
swings it from side to side while performing the
Shudder. More commonly, the neck is motionless and only the head moves vertically, producing a jiggling appearance. During the
Shudder, the dewlap is maximally extended, the
forelegs are one-half to fully extended. Two
hundred and twenty-nine filmed Shudders performed by 16 individuals were suitable for
analysis. Of these 64, or 28 %, were performed
while the iguana was walking.
3. Roll. This movement corresponds to
Miiller's (1972) 'high frequency nodding' and
to Distel & Veazey's (1982) 'rotatory head
nodding'. In performing the Roll, the head is
lifted at an approximately 45 ~ angle to the substrate and rotated along the longitudinal axis of
the body. Vertical movements varying in duration and amplitude may occur simultaneously
with the head rotations (Fig. 1C, 1E). The dewlap is maximally extended, the forelegs are onehalf to fully extended. The Roll is much more
exaggerated than the Shudder. During the high
amplitude head rotations of the Roll, the long,
pendulous dewlap undulates as the head moves
from side to side. The iguanas typically performed Rolls from exposed perches, thereby
enhancing the conspicuous nature of this headbob. Locomotion was never observed to occur
simultaneously with the Roll. Forty-nine Roils
performed by 12 individuals were analysed.
4. Roll-shudder. This bob type consists of
rapid alternations of Rolls and Shudders in
highly variable sequences (Fig. 1D). Body

330

ANIMAL

BEHAVIOUR,

30,

[_._

|
I sec.

Fig. 1. Patterns of iguana head movements in which the horizontal axis rep
resents time in seconds, the vertical axis represents amplitude: (A) Head jerks
from three individuals; (B) Shudders from two individuals: (C) Two Rolls performed by the same individual; (D) Roll-shudders from two individuals;
(E) Rotatory head movements performed during the Roll. Top graph of D is a
Shudder-Roll-Shudder sequence; bottom graph is a Roll-Shudder sequence.
position is the same as during the Shudder and
Roll. Thirty-four Roll-shudders performed by
14 individuals were analysed. In four of these,
locomotion occurred simultaneously with the
Shudder component of this bob type.
5. Signature Bob. This behaviour corresponds
to Distel & Veazey's (1982) 'stereotyped headnodding', and to MiiUer's (1972) 'low frequency
nodding'. Hazlett (1980) referred to this movement as the assertion display. The Signature Bob
is a high amplitude vertical head movement
during which the head is elevated at a 60 to 90 ~
angle to the substrate, followed by a second
high amplitude movement of longer duration
(the plateau). A series of shorter, low amplitude
bobs follows (Fig. 2). During the plateau, the
head is held up for about 1 s, and head rotations
occur. These head rotations are much reduced
compared to those of the Roll. The Signature
Bob is performed from both sitting (forelegs
extended) and lying (ventral surface in contact
with substrate) positions. The dewlap is maximally extended. The analysis included 258
filmed displays performed by 13 individuals.
Contextual Correlates
Display rates varied with the sex and size of
the individual. Females displayed much less
than males; virtually all displays given by females
occurred during interactions. Large males
(longer than 36 cm body length) displayed more
often than smaller males throughout the year.
Large males were more likely than medium

males (32 to 36 cm) to perform at least one display during an observation period (%2 = 26.9,
df = 1, P < 0.001).
Overall display rates also varied seasonally,
being greatest when mating activity was most
IGUANA

0.5

1.0

1.5

F,~O. 19

r z , ~,

I 1

, E I I

2.0

2.5

~.0

A NO. I0

"I/~

1.0

1.5

0.5

I r I I I,L.?

1.0

2.0

. /~/~ 9

t t I | t

0.5

I ]
1.5

1
2.0

P v I

2.5

IGUANA

q I

~,.0

NO, 2

t I [ I f f I

~ II
2:5

III

]
3.0

TIME IN SECONDS

Fig. 2. Samples of two DAP graphs of Signature Bobs

from each of three different male iguanas. Displays for
each iguana were filmed on the same day.

DUGAN: FIELD STUDY OF IGUANA HEADBOB DISPLAYS

intense (Fig. 3). The proportion of displays observed that were Roll-shudders and Signature
Bobs, however, remained relatively constant
throughout the year, with the Roll-shudder
accounting for 3.2% overall, and the Signature
Bob for 65.0%, of the total. The Roll and
Shudder, on the other hand, showed more drastic fluctuations (Fig. 3). The relative frequency
of Shudders increased during all breeding season
months. The Roll showed the greatest increase
in October, when the number of Rolls exceeded
the number of Shudders. The fluctuation in
relative frequency significantly deviated from
the expected for the Roll (;(2--_ 23.6, dr= 9,
P < 0.01) and the Shudder (%2 = 34.6, dr= 9,
P < 0.001).
The Shudder was most likely to occur at close
distances (0.0 to 1.5 m). Display performance
was much more variable at intermediate distances. When the distance between a male and
female was greater than 3 m, the Roll tended to
be more frequent than either the Shudder or the
Roll-shudder (Fig. 4). The relative proportions
o f the three headbobs differed significantly from
the overall proportions between November and
January at all distances except 1.5 to 3.0 m (0.0
to 0.5 m, ;(2 = 37.2, df = 2, P < 0.001 ; 0.5 to
[ ] Roll
r~ Shudder

1.5 m, %2 = 20.3, df = 2, P < 0.001 ; greater

than 3 m, K o l m o g o r o v - S m i r n o v test, P < 0.01).
Bob performance also varied according to the
social context. Solitary males usually performed
Rolls; over 95% of displays were Shudders
during Male-Female Interaction (Fig. 5). Solitary males did perform the Shudder or Rollshudder; however, these were rare events that
never occurred during filming sessions. The frequency of the three bobs differed significantly
from the overall proportions in all cases (Solo,
K o l m o g o r o v - S m i r n o v test, P < 0.01; N o t Directed, ;(2 _= 10.2, df= 2, P < 0.01; General
Social, ;(2 = 11.9, df= 2, P < 0.01; M a l e Female Interaction, ;(9 = 46.5, df= 2 , P < 0 . 0 0 1 ) .
In the N o t Directed category, differences were
primarily due to a greater than expected number
of Roll-shudders, while the low frequency of
Rolls contributed most to the differences in the
General Social category.
In summary, the Roll is more likely to occur
in 'non-social' situations; the Shudder is more
likely in social contexts. Although low in frequency in all contexts, the Roll-shudder tends
to occur most often when a recipient is just outside the displayer's individual distance (I observed it at between 1 and 1.5 m, the distance at
[ ] Roll-Shudder

[ ] SignatureBob

IO

<

J
(3rj}

2-

o
0
( 55)(

N
[D
46 ) ( ~ 3 9

d
)( 25 )

IBREEDINGACTIVITYI
( No.

Mar
Apr

331

Jul
Aug

0
N
D"
(53)(59)(80)(37)

(40) ( 44>

I BREEDINGACTIVITYI

Month
15-rain. intervals)

Fig. 3. Mean number of headbob displays observed per 15-min observation

interval, and proportion of total displays observed that were Rolls, Shudders,
Roll-shudders, and Signature Bobs. Large males only, 1977-1979. Vertical lines
represent :E SD a/2/N,where N = number of intervals. Non-overlapping lines
indicate a significant difference (P = 0.05) between any two pairs (see Burghardt
1969).

ANIMAL

332

BEHAVIOUR,

which escape or avoidance responses occurred),

and in the intermediate social contexts (Not
Directed and General Social). It is at these distances and in these contexts that the apparent
social nature of the large male's behaviour may
change (between N o t Directed and General
Social, or General Social and Male-Female
Interaction). In order to determine whether the
Roll-shudder is restricted to transitions between
social contexts, the situations in which the Rollshudder occurred were further sub-divided into
(1) transitional categories, including Rollshudders performed during transitions between
0 ROLL

1.0

H ROLL-SHUDDSR

0.8

F0
I--

ta_ 0.6

0
Z
0

F- o.a
2O

0.2

O- 0.Srn
03,120)

0.5- I.Sm
{9,88)

[i

1.5 - 3.Ore
(6,461

> &Ore
(6,41)

D I S T A N C E FROM FEMALE
(NO, INDIVIDUALS, NO. DISPLAYS)

Fig. 4. Median proportion of total Roll, Shudder, and

Roll-shudder displays given by males at varying distances
from females. Line denotes interquartile range; numbers
in parentheses refer to the number of individuals and
number of displays, respectively.
I.O

0 ROLL
$ A SHUDDER
t

-.10.8

D ROLL-SHUDDER

30,

sequences of Rolls and sequences of Shudders,

between social contexts, and between patrols
and social contexts, and (2) non-transitional
categories, including Roll-shudders interspersed
among sequences of Shudders or among sequences of Rolls, and (3) indeterminate contexts, in which the activity preceding and/or
following the display was not observed. Twentytwo Roll-shudders occurred in transitional situations, and only nine in non-transitional situations (Z2 = 5.45, df= 1, P < 0.02). Three
were indeterminate. Roll-shudders were more
likely to occur during sequences of Shudders
than sequences of Rolls, but were never mixed
with Shudders in the course of a social interaction.
The Signature Bob punctuated activity in all
contexts. F r o m the filmed record, I noted the
event immediately preceding 723 displays. No
behaviour preceded 54 of the 723 displays. Some
of the displays m a y have followed an event that
was not recorded on film. Twenty-eight of the
displays occurred at the approach of a female,
and the remaining 641 followed some other
behaviour by the displayer (Table I). These
Signature Bobs were not proportionately associated with all preceding acts, but rather tended
to follow more perch changes and fewer Shudders and Roll-shudders than expected (Z z = 48.4,
df= 5, P < 0.001). Sequences o f these acts
often preceded Signature Bobs. The extent to
which the sequential ordering of behaviour influenced the association between the Signature Bob
and other activities is not known. Nevertheless,
it is clear that the Signature Bob occurred during
pauses in, and at the termination of, ahnost any
activity in which a large male engaged. After
performing a Signature Bob, an individual might
continue the same activity, change to another
activity, or cease activity.

I.l. 0.6
0

Table I. Frequency with which Varions Behavioural Patterns by the D[splayer Preceded Signature Bobs, and the
Overall Frequency of the Behaviour. Proportional Frequencies are in Parentheses

0.4
o

~0 0.2
fl..

Solo
(3,14)

__
I~.
~ L o
Not Directed GeneralSocial
(9,91)

M/F Interaction

(12~78)

(10,112)

SOCIAL CONTEXT
(NO. INDIVIDUALS, NO. DISPLAYS)

Fig. 5. Median proportion of total Roll, Shudder, and

Roll-shudder displays given by males in different social
contexts. Line denotes interquartile range; numbers in
parentheses refer to the number of individuals and number of displays, respectively.

Behaviour
Roll
Shudder
Roll-shudder
Perch change
Position change
Tongue-touch

No. that preceded

Signature Bobs

Overall
frequency

60 (0.094)
173 (0.270)
31 (0.048)
233 (0.363)
114 (0.178)
30 (0.047)

80 (0.089)
308 (0.342)
46 (0.128)
283 (0.314)
141 (0.157)
42 (0.047)

DUGAN: FIELD STUDY OF IGUANA HEADBOB DISPLAYS

Variation in Form
The Roll, Shudder, and Roll-shudder. The
Roll, averaging 1.26 s in duration (SD = 0.41),
tended to be shorter and less variable than the
Shudder ( ~ = 2.79, SO = 1.24) and Roll-shudder
(-~ = 2.68, so = 1.09). Individual differences in
display performance followed this overall trend,
with the relative duration of the Roll being
shorter and less variable than that of the
Shudder (Fig. 6). Inter-individual comparisons
in the performance of the Roll-shudder are
difficult to interpret due to its low frequency.
With an overall coefficient of variation (CV)
of 32.38, the Roll falls within the typical range
of variability of modal action patterns (MAPs)
that have been quantified for a large number of
species representing different taxonomic groups:
CVs of durations based on inter-individual
variability range from 15 to 35 (summarized in
Barlow 1977). The Shudder (CV = 44.46) and
Roll-shudder (CV = 40.84) exceed these values.
Although the differences among the three are
not large, the Shudder is significantly more
variable than the Roll (c statistic, Dawkins &
Dawkins 1973).
The relationship between display duration
and distance between, the displaying male and
female accounts for some of the differences
among displays in temporal variability (Fig. 7).
The Shudder became shorter and less variable
as distance from the female increased. Shudder
duration was influenced by both individual
identity (F = 2.70, df= 15, 193, P < 0.001)
and distance from the female (F = 5.11, df=
3, 15, P < 0.025). The duration of the Roll was
little affected by distance from the female, but

was significantly related to individual identity

(F = 2.91, df= 10, 32, P < 0.025). The Rollshudder increased slightly in duration at 0.51.5 m, but otherwise followed the same general
trend as the Shudder. No significant effects were
found for the Roll-shudder, although again
small sample size makes interpretation difficult.
Since the Shudder tended to decrease in duration as distance from the female increased, it
was of interest to ask whether or not significant
differences in the duration of this display might
be found among the three social context categories (recall that Shudders did not occur when
the male was solitary). Differences among the
social contexts did occur (F = 10.02, df = 2, 17,
P < 0.001). The duration of the Shudder was
significantly greater during interactions (,~ =
3.19 s) than in the General Social ()? = 2.56 s)
and Not Directed (J? = 2.23 s) categories (Duncart's test). This pattern was the same in all
individuals. The General Social and Not Directed categories include instances where males
and females are close together but not interacting. Thus it appears that Shudder duration is
more influenced by the occurrence/non-occurrence of an interaction than by distance.
The Signature Bob. The pattern o f head movements characteristic of the Signature Bob was
divided into two units (Fig. 8). Unit 1,
consisting of the pre-plateau bob, the plateau,
and the post-plateau bob, occurred in all Signature Bobs and was considered to be the
species-typical component of this display. Unit 2
was more variable; any number of Unit 2 bobs
might follow Unit 1. Only one o f 259 displays
analysed stopped after Unit 1.

8.0

[ ] ROLL
[ ] sHu~R

[ ] ROLL-$HUODER

6.0

4.0

2.0

2o

Toc

333

SPMF

H$

CR

SPST

I NDI VI DUALS

Fig. 6. Mean duration 4- SD of Rolls, S~"-dders,and Roll-shudders

performed by six individuals. N of each bob type is above the
vertical line.

334

ANIMAL

BEHAVIOUR,

The initial analysis considered 10 components

of the Signature Bob. Duration measures mctuded the duration of the plateau (PLT), the
summed durations of the pre- and post-plateau
bobs (U1-P), and Unit 2 (U2). Amplitude
measures included the pre-plateau peak (A-l)
and dip (A-2), the plateau peak (A-3), the postplateau dip (A-4) and peak (A-5), and the mean
amplitude of Unit 2 bobs (A-6). The number of
Unit 2 bobs ( # U2b) was also counted. The mean,
standard deviation, and coefficient of variation
of each component and o f total duration are
presented in Table II.
Neither distance from female nor preceding
behaviour (Roll, Shudder, Roll-shudder, locomotion, other, or unidentified) influenced the
variability of the I0 components. Only individual identity (F = 8.39, d f = 120, 1334, P <
0.0001) and t h e individual-by-distance interaction (F = 1.58, d f : 170, 1494, P < 0.0001)
were significantly related to variability of bob
components (MANOVA, Tatsuoka 1971). Univariate analyses of variance indicated which
components contributed to the MANOVA
results. All components differed significantly
among individuals. The relative amplitudes of
the pre-plateau dip (A-2), the plateau (A-3),
and the post-plateau peak (A-5), varied with
the indivldual and the distance between the
male and female. None of the duration measures
and none of the other amplitude measures were
influenced by the interaction of these two
variables.
A Signature Bob is a continuous movement;
measurements made on one segment will depend
in part on durations and amplitudes of other
segments. Thus it was not clear to what extent
the low intra-individual variability of all
measures might be due to dependence among

30,

display components. Nor was the strength of

each component as a unique discriminator
among individuals known. If the Signature Bob
does communicate individual identity, must the
iguanas witness the overall pattern of movement,
or might a certain segment be the important cue ?
A stepwise discriminant analysis was used to
identify the components that best separated
individuals.
In discriminant analysis, the values of discriminating variables are weighted and combined to form one or more linear functions that
maximize the variance between groups while
minimizing that within groups:
Di =

dizZ1 + d~2Z2 + . . . . . .

--

TOTAL

UNIT:

......

~!
--,-,--UNIT 2--- ~-[

A i

0 ROLL

I ^

6SHUDDER
OROLL'SHUDDER

dipZp

where Df is the score on the discriminant function i, the di's are weighting coefficients, and
the Z's are the standardized values of the p discriminating variables used in the analysis. The
coefficient reflects the relative contribution of its
associated variable to that function.
A stepwise procedure first selects the strongest
predictor variable, and then enters the 'next best'
discriminator at each step ( N i e e t al. 1975).
Dependencies among variables are written into
the model; the relative strength of each variable
as an independent predictor is indicated by its
conditional F-value. The conditional F ' s are
difficult to interpret, however, if extreme dependencies exist between two or more variables.
Pearson product-moment correlation coefficients
revealed that although 28 of 45 pairs were significantly correlated, the correlation between
only one pair, U2 and # U 2 b (r = 0.89) was
greater than 0.60, an arbitrarily selected cut-off

. . . .

50

PLATEAUI~
A-J~
~

r~A-6----

- l

40

Vi

3o

A4

-_z
Z
O

2,0

OC
C~

l.O

vI v v v \ I
I
I

R E L A T I V E AMPLITUDES

AI
0 " 05m

I[
0 5 - 1.Sm
DISTANCE

[l
1.5- 5,Orn
FROM

.__.J I
> .~.Om

__
OVERAt,t

FEMALE

Fig. 7. Mean durations + SD of displays performed by

male iguanas at varying distances from females.

A-I : Pre-ploteou Peak

A - 4 : Post-plateau Dip

A-E: Pre-plateau Dip

A - 5 : Post-plateau Peak

A-3=

Plateau Peak.

A-6: X

amplitude of Unit?
bobs

Fig. 8. Components of the Signature Bob used in analysis.

See text for explanation.

DUGAN: FIELD STUDY OF IGUANA HEAl)BOB DISPLAYS

value. The univariate F-values indicated that
# U2b (F = 12.84) might be a stronger predictor
than U2 (F = 4.55), and thus U2 was omitted
from the analysis.
The stepwise procedure used all of the remaining nine variables in forming the discriminant
functions. The functions significantly discriminated a m o n g individuals (F = 25.1, d f = 108,
1738, P < 0.0001). Nine discriminant functions
were formed. The duration of the plateau and
of the remainder of Unit 1 contribute most to
the separation among individuals on the first
four functions, which account for 90.6 % of the
variability among individuals (Table III). Thus,
although # U 2 b and the amplitude measures
contribute to the separation of individuals, when
all variables are considered together individual
differences are primarily due to duration
measurements made on Unit 1.
The conditional F-values (Table IV) indicate
that plateau duration is by far the best predictor
o f an individual's bob performance, followed by
the amplitude of the pre-plateau dip, postplateau dip, and # U2b, respectively. Thus, when
overlap among the variables is taken into consideration, the strongest independent discriminators are to be .found within Unit 1.
The discriminant functions correctly classified
88.4 % of the Signature Bobs. In a more exacting
test, the jack-knife classification, 83.7% of the
displays were classified correctly. The expected
percentage of correct classifications based on
chance (determined from the proportional distribution of sampled displays among individuals)
was 9.1.
Discussion

Each iguana headbob type represents a unique

combination of a particular form, degree of
Table H. Mean, Standard Deviation, and Coefficient of
Variation (CV) of Signature Bob Components. Duration
Measures (D) are Seconds; Amplitude Measures (A) are
Percentage of Highest

Component

Mean

so

CV

(D) PLT
(D) U1-P
(D) U2
Total duration
A-I
A-2
A-3
A-4
A-5
A-6
~U2b

1.061
0.722
1.235
3.012
98.043
56.310
92.752
28.911
69.857
62.120
4.725

0.107
0.076
0.322
0.419
3.346
9.203
5.245
11.909
11.289
13.006
0.993

10.068
10.588
26.095
13.900
3.413
16.344
5.655
41.192
16.160
20.937
21.025

335

stereotypy, and pattern of occurrence. In order

to understand the different associations among
these display features, it is helpful to further
consider the use of the displays, the problems
that an iguana displaying in a forest habitat
must solve, and general selective pressures
thought to influence stereotypy.
The Roll. Because of its high relative frequency
in the early stage of the breeding season, its
exaggerated movement patterns, and its performance from highly conspicuous display posts,
I consider the Roll to be an advertisement display, alerting neighbours to the presence of a
resident male. In the early part of the breeding
season, a male successful in establishing a territory and attracting females must make his
location and status known. As this period
coincides with the end of the rainy season, the
foliage is much thicker than at other times of the
year. The Roll appears to be uniquely suited to
overcome obstacles to vision. The constant
rhythm of the head and dewlap stands out
against the background of other environmental
'noise', such as leaves swaying in the breeze. As
iguanas are dispersed at this time, an advertisement display must transmit over some distance.
Variance in signal performance should be low
in a long distance signal, thereby facilitating
accurate detection of the signal from its background (Marler 1977). In keeping with this prediction, the Roll is more stereotyped than the
Shudder, a display used primarily in interactions.
The close temporal relationship between the
Roll and the Signature Bob suggests a second
Table HI. Standardized Discriminant Function Coefficients (d~'s) Showing the Relative Contribution of the
Variables to Each Discriminant Function (dr's)

d~
Components

df 1

U1 -P
--5.349
PLT
--7.936
A-I
--0.027
A-2
0.005
A-3
--0.033
A-4
--0.036
A-5
0.002
A-6
0.036
~U2b
0.416
Percentage of
variance among
individuals
55.5
Cumulative percentage

df 2

df 3

df 4

--3.431 --2.017
0.141
3.945 --1.934
0.981
--0.018 --0.070
0.133
0 . 0 7 7 0.033 --0.037
--0.023
0 . 0 8 2 0.013
0.015 --0.087
0.006
--0.028
0.027 --0.052
--0.005 --0.020 --0.052
0.314 --0.269 --0.069
16.7
72.2

11.6
83.8

6.8
90.6

336

ANIMAL

BEHAVIOUR,

function for the Roll: it may introduce the

Signature Bob that invariably follows. Stamps
& Barlow (1973) reported that in Anolis aeneus,
introductory head movements were more likely
to precede a Signature Bob with increasing distance between interactants. They suggested that
introductory movements function to call attention to the Signature Bob. The Roll may serve
a similar function in L iguana, saying 'here I am,
Signature Bob to follow'. The appropriate response would depend on the sex of the recipient
of the signal. Males might avoid the displaying
male or approach to challenge him; females
might avoid, or approach to inspect, a potential
mate. This interpretation is consistent with the
change in relative proportions of each display
type across seasons. Only in October, in the
initial stages of the breeding season when many
large males were solitary, did the Roll exceed
the Shudder in proportion of total displays given.
The Shudder. Shallow, rapid headbobbing has
been reported in a variety of iguanid lizards (see
Jenssen 1977). In L iguana, the Shudder occurs
in male-female interactions both during and outside of the mating season, and thus is not considered to be a courtship bob. Outside of the
mating season, Shudders were most commonly
observed when a male and female moved close
to one another while basking. This display may
signal that threat or attack is unlikely, an
explanation that is in keeping with suggestions
that the Shudder evolved to allow close contact
between males and females (Stamps & Barlow
1973; Barlow 1977).
As observed in other iguanids (see review in
Jenssen 1977), locomotion may occur simultaneously with Shudder movements, but never
with other headbob displays. In 61 of 68 filmed
cases in which locomotion and Shudders occurred together, the displaying iguana was
approaching a female. If the Shudder indicates a
male's lack of aggressiveness, thereby reducing
the probability that the female will retreat at his
approach, the ability to display and approach
simultaneously would have an obvious advantage.
The Shudder is the least stereotyped of iguana
headbob displays. Marler (1977) suggested that

30, 2

greater variability, as with graded displays, provides the potential to communicate more refined
information over short distances and to adapt
to feedback from a respondent. Since the interactions between male and female iguanas were
not analysed, Marler's hypothesis cannot be
evaluated fully. However, unlike social signals
that exhibit a 'typical intensity' (Morris 1957),
the duration of the Shudder is not constant
across social contexts. Its duration is inversely
related to distance and is significantly greater
during Male-Female Interactions than in noninteractional contexts. Assuming that the male's
level of arousal is greater during courtship than
while resting near a female, then the duration of
the Shudder may provide information about the
male's level of arousal. Shuddering seems to
reflect arousal levels in sceloporine iguanids as
well (Ruby 1977; Rothblum & Jenssen 1978).
Shudder duration may also reduce ambiguity
regarding the male's intention. The most distinctive difference observed in a large male's behaviour in the General Social and Male-Female
Interaction contexts was the duration of the
Shudder. Otherwise the male behaved similarly
in both contexts: the Shudder was the most
frequently occurring display, the male was often
close to more than one female, and movement
toward females occurred in a variety of contexts.
In effect, the greater duration of the Shudder may
be equivalent to the male's saying: 'I am talking
to you'. This message, along with other contextual cues (e.g. distance from male, male
approaches female) may facilitate the female's
recognition that she is the recipient of the male's
attention.
The Roll-shudder. A gross examination of the
contextual correlates of the Roll-shudder indicates that it does convey information about the
response tendencies of the displaying iguana: a
change from one set of activities to another set
of activities is highly likely. My impression was
that one could predict the direction of the
behavioural transition to be made by the displayer based on the bob sequence. For example,
a Roll to Shudder sequence is likely to precede
movement into a more social context, whereas a
Shudder to Roll sequence is more likely to pre-

Table IV. Conditional F-values of Signature Bob Components, Ranked in Descending

Order. All Values are Significant at P < 0.001, d f = 12,237

PLT
71.38

A-2
A-4 7~U2b A-3
A-5
U1-P
28.11 26.46 22.47 13.67 13.42 10.27

A-6
7.29

A-I
5.87

DUGAN: FIELD STUDY OF IGUANA HEADBOB DISPLAYS

cede leaving a social situation. However, more
complex sequences occurred, such as the
Shudder-Roll-Shudder sequence shown in Fig.
1D. The performance of such a sequence while
close to, but not interacting with, a female may
reflect ambivalence on the part of the male
regarding approach versus withdrawal.
Since the Roll-shudder is an apparently variable intergradation between the Roll and the
Shudder in its form, variability and use, one
might ask whether it should be classified as a
separate display. Due to the small sample of
filmed Roll-shudders, it could not be determined
whether the behaviour consists of a very few
fixed sequences or represents a range of variable
combinations. It seemed that the form(s) of the
Roll-shudder would most likely fit Smith's (1977)
category of variable display sequencing, in which
patterns of behaviour have not been formalized,
and that the head movements should not be
classified as a separate display.
The Signature Bob. Signature Bobs in L iguana,
as in other iguanids, occur in a variety of circumstances such as aggressive and courtship
encounters, territory advertisement or assertion,
and after locomotion. Based on the display's
pattern of occurrence, it fits the message category that Smith (1977) termed 'general set
alternatives'. These displays are always associated with some definable activity, but give no
information about the activities that are being
changed or the alternatives replacing them. They
occur in a wide variety of circumstances, and
information concerning the probability of future
behaviour is derived from the context.
The Signature Bob is more stereotyped than
most MAPs (Barlow 1977) in duration measures
made on Unit 1, total duration, and two of the
amplitude measures (Table II). Stereotypy and
species uniqneness are common characteristics
of iguanid Signature Bobs, and a species recognition function is usually ascribed to these displays (see review in Ferguson 1977).
Although experimental support is lacking,
Signature Bobs may communicate individual as
well as species identity. Individual recognition
should evolve if advantages are derived from
discrimination between comparably detected
signals. Iguana iguana is territorial only during
the breeding season, when nearest neighbours
are 25 to 100 m distant. Although movements of
large males overlap spatially at other times of the
year, temporal overlap is rare. Individual recognition might allow a male to avoid a larger,
dominant animal, and thereby avoid a potentially

337

costly encounter. Individual recognition might

also facilitate mate acquisition. Where males
are spread apart and females pass through the
territories of several males before mating, the
female must presumably be able to remember
and compare displays from several suitors
(Brander 1967; Shepard 1975). Since much early
courtship between iguanas occurs in locations
other than those selected for territories in late
November, remembering the male's identity,
rather than the locations of early courtship
interactions, would aid the female in her final
choice of mate.
The Signature Bob resembles the Roll in that
it is an exaggerated headbob that is conspicuous
in its patterned rhythm, high amplitude head
movements and dewlap motion. Given this
similarity, one might ask why the Roll has
evolved, and why it prefaces a similarly conspicuous display in territorial advertisement.
However, although each Signature Bob component reliably discriminated among individuals,
the most distinctive part of the Signature Bob,
Unit 1, occurs at the beginning of the display.
If individual recognition is important, then the
introduction of the Signature Bob with a Roll
should ensure that Unit I is noticed by conspecific observers from a distance. The question
remains, why were not all Signature Bobs preceded by a Roll? Presumably, activities performed in other contexts serve to introduce the
Signature Bob. In contrast, the Roll appears to
be specialized to introduce the Signature Bob
in one context only: advertisement.
Conclusion. Signal transmission in L iguana is
accomplished with a highly redundant system
of visual signals, including selection of conspicuous trees and conspicuous perches in those
trees, frequent performance of conspicuous displays, and colour intensification. The redundancy
in the system may be viewed as an adaptation to
a very complex habitat, where both distance and
visual obstacles must be overcome. Among forest
birds and monkeys that require signals to maintain intergroup spacing, auditory exchange is
common since it is less hindered than vision by
obstacles. The importance of visual signals and
the variability of the signals tend to increase in
close-range, intragroup communication (Marler
1977). Iguanid lizards do not have the option of
using auditory signals. They must depend for
the most part on visual communication, particularly for long distance signaling. However, it
appears that patterns of variation in visually
communicating iguanas parallel such patterns

338

ANIMAL

BEHAVIOUR,

in species living in similar environments but

employing other sensory modalities for communication.

Acknowledgments
Based in part upon a dissertation submitted in
partial fulfillment of the requirements for the
Ph.D. degree at The University of Tennessee.
This research was supported by NSF Grant
BNS-77-11344, by grants from the Smithsonian
Tropical Research Institute, Sigma Xi, The University of Tennessee Department of Psychology,
and by NSF grants BNS-75-02333 and BNS-7814196 awarded to G. M. Burghardt. I owe many
of the ideas in this paper to discussions with
A. S. Rand. 1 thank H. Molina and J. Philpot
for statistical advice, and G. M. Burghardt, A. S.
Rand, and T. A. Jenssen for helpful comments
on various drafts of the manuscript.
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(Received 25 February 1981 ; revised 15 July 1981 ;

MS. number: A2614)