Sie sind auf Seite 1von 55

Behaviour of a harbour porpoise (Phocoena phocoena)

mother-calf pair in captivity

(C) Fjord & Baelt. Photo by Ewa Borowska

Individual study activity by Lara Delgado García


Supervisor Magnus Wahlberg
Institute of Biology
University of Southern Denmark
Behaviour of a harbour porpoise (Phocoena phocoena)
mother-calf pair in captivity

Lara Delgado1* and Magnus Wahlberg1,2

1 Institute of Biology, University of Southern Denmark, Campusvej 55, 5230 Odense M, Denmark.
2 Fjord & Baelt, Margrethes Plads 1, 5300 Kerteminde, Denmark.

ABSTRACT

In this study the social and spatial interactions between a harbour porpoise cow and calf
are described. The study started three months after the birth and presents the
development of their interactions during 10 months in a captive situation. This is the
first time a porpoise calf is documented to have been born and survived in captivity.
The relative spatial positions of the mother and the calf were measured, the apparition
of new behaviours and the variation in their occurrence along time, was described. The
study was done by means of quantitative measures using focal sampling. Parent-
offspring conflict (avoidances and aggressions when the calf was close to the mother),
tended to increase when the calf was 8 months old, while potential fishing increased
around that time. The total time the calf and her mother spent in close proximity varied
but an overall tendency to decrease was seen, significantly decreasing around the 10th
month of the calf. The longest interval of time calf and mother spent separated increased
from 6 minutes when the calf was 3 months, to up to 35 minutes when it was 8 months
old. But nursing stayed constant over all the study, postponing the expected weaning
time for the species. Additionally differences in mother and calf time budgets, and
durations of behaviours showed probable age related differences. The study of the
development of the calf during its first year has capital importance for understanding its
survival in the age class with the highest mortality in the species. Additionally there is
an almost absolute lack on harbour porpoise mother calf behaviour. Even though the
study is based on only a single mother-calf pair, it sets a basis of mother and infant
interactions for the first time.

Keywords: behaviour, harbour porpoise, Phocoena phocoena, captivity, time together,


infant position, behavioural development.

*Corresponding author: Tel: 0034 646 11 58 01. E-mail address: lara_6_a_secas@hotmail.com

1
INTRODUCTION

In all mammalian social systems the mother-calf bond is the core unit (Gubbins
et al. 1999). In the case of harbour porpoises the mother-calf bond, as in other marine
mammals, is expected to be very strong. Harbour porpoises are long-lived animals — an
average of 12 years (Lockyer 2003) —, invest highly in offspring — 10 to 11 months of
gestation (Lockyer 2003)—, and sociality in this species has been shown to be, as in the
majority of marine mammals, an important feature of their biology, with an interesting
diversity of displays performed towards different individuals depending on their sex,
age or season (Delgado et al. in prep.b). However any study has been done until today
that described the mother-infant relation along time, focusing on developmental changes
in their behaviour. There have been only three studies on behaviour that included a
harbour porpoise mother and calf pair (Oleksenko & Lyamin 1996; Delgado et al. in
prep.b; Borowska 2009). Oleksenko & Lyamin’s study focused on the identification of
resting states in a captive situation, having the animals quite space-limited conditions
(Oleksenko & Lyamin 1996). In Borowska (2009) the same mother-calf pair studied
here was observed in September 2008, two weeks later to the end of this study, during
two weeks, quantifying their behaviour and breathing rates. And in Delgado et al. in
prep.b., another study made by the author in captivity, the relationship among different
harbour porpoises was compared, including the same mother-calf for a period of a
month. The infant was one year and a half old by the time of that study.
There are two punctual observations of mother-infants in the wild, one being the
first qualitative description of mother and infant behaviour (Amundin and Amundin
1974) and one study of the swimming patterns, that included an 11 months old calf and
her mother during some hours (Teilmann et al. 2007). The first study gives some input
in the close-bond maintained among mother-calf, and some distances descriptions and
reaction to other individuals. The second, providing the first direct empirical
observation of mother-calf bond lasting, swimming together at the infant age of 11
months, which gives an indication for the longevity for the social bonds mother and
calf, which may last at least until 11 months.
Lockyer (2003) presented the biological parameters of this species obtained
from studies done on directly caught, by-caught or stranded animals. This information
provided insights that can give a basis for setting the mother-infant relation as duration

2
of lactation, size or age at weaning, pregnancy rates or age at sexual maturity. However
the first two parameters (duration of lactation and weaning time) are uncertain, and only
observation on one or two individuals set the ranges that follow. The likely duration of
lactation was at least 8 months (the end of the milk intake it is not clear, but the
beginning of fish eaten). In relation with size or age at weaning, it was observed fish
eating in a captive calf at 10 months of age, but the mother was not present, and so the
continuation of nursing could have been possible together with the fish consumption as
weaning is a gradual process. There are many observations of stranded alive calves
consuming fish in captive conditions, with lack of the mother. But those results do not
define the natural weaning range. At 4 months of age calves were observed to eat
shrimps in Lockyer (2003).
In other marine mammals multiple studies demonstrate that mothers tend to
decrease they role in maintaining proximity to their calves as they age; and that infants
spend less time close to their mothers. In addition, infants spend more time travelling
and socializing as they age. And also they also separate form their mother more often
and for longer periods of time, and also increase foraging as they age (in dolphins: e.g.
McBride & Kritzler 1951; Chirighin, 1987; Mann & Smuts 1998; Mann & Smuts 1999;
Mello et al. 2005; belugas: e.g. Krasnova et al. 2009). All this have been studied in
other marine mammals, and set on time for the species. However, nothing is known
about harbour porpoises in this aspect, and this study will try to provide some basis.
This study describes the behaviour of a harbour porpoise (Phocoena phocoena)
mother-cow pair in captivity during 10 months. It focuses on developmental trends and
signs of weaning and parental investment, as well as on other aspects of mother-infant
interactions, spatial relations, and behavioural differences among them. And the
obtained basis will be compared with the developmental trends observed in other
marine mammals, and conditioning factors that could have affect them in the specific
studied environment will be discussed.

3
MATERIALS AND METHODS

Animals
The subjects of this study were a harbour porpoise (Phocoena phocoena) calf
and cow kept at the Fjord&Bælt (FoB) research and outreach centre in Kerteminde,
Denmark. The calf, Frigg, is the first harbour porpoise that has been reported to be born
and survived in captivity. It is a female born on the 9th of August of 2007 and parented
by the female Freja and the male Eigil (Blanchet et al. 2008). The calf was three months
old at the beginning of the study. The mother, Freja, was approximately 11 years old at
the time of the birth — estimated to be born in the summer of 1996 by means of her
growth curves and length at arrival (Wahlberg et al. in prep.). The father and another
younger female were kept in a contiguous pen separated from mother and calf; both
groups were able to maintain acoustical communication towards the separation nets.

Environment
The behavioural observations were conducted at FoB, where the harbour
porpoises are housed in a semi-natural outdoor enclosure in the harbour of Kerteminde.
The pen where mother and calf were enclosed is separated by the harbour by means of
nets which provides natural environmental conditions of water flow and quality, tidal
currents, temperature, and also allows the entry of small-sized flora and fauna into the
enclosure (Lockyer et al. 2003).
There were three changes in the environment when the animals were kept in the
same pen for the long term study.
· The first environment was present from the beginning of the study, 3rd month of
the calf, and until the beginning of the 5th month of the calf, i.e. the environment
was kept constant for 2 months. The first environment consisted of a pen
delimited by nets of a dimension of 8 by 13 meters and around 2 meters deep.
The pen contained a floating pool of aluminium walls and bottom and an
opening in one side. The pool had the dimensions 4x4 m. In this first
environment the animals could swim under the floating pool where they were
out of sight.
· The second environment consisted of the same pen but without the floating pool.
This situation was kept from the begging of the 5th month of the calf, until the

4
beginning of the 8th month of the calf. The animals were visible 100% of the
time, in the 8x13 meters area.
· The third environment was kept from the beginning of the 8th month of the calf
age until the end of the study at 12 months old, i.e. the environment was kept for
5 months. It consisted of the same pen where the inner nets were removed and
the animals were able to swim underneath the floating pontoons that covered the
sides of the pen. The dimensions of this pen were 11x14 m and a depth that
varied with the tide, ranging from 2 to 4 meters, with an averaged depth of 3.5
meters. The bottom of the harbour was reachable, being natural rocky and sandy
bottom, still separated from the harbour by nets. The animals were visible within
the area with an open water surface covering 8x13 meters, the same dimensions
of the first environment, but were not visible when swimming underneath the
pontoons.

Data collection
Observations of the two harbour porpoises were made a total of 52 days during a
period of 10 months, done at least once every week, sometimes twice a week,
depending on the weather. The porpoises, cow and calf, were observed for 2 hours
every observation day (a total of 104 observed hours). Being each of them focal animals
(Altmann 1974) during alternate 15 minutes, therefore, per day each animal was
observed during one hour in 15 minutes intervals; with 15 minutes pause between each,
where the other animal was observed. The data was collected from direct observation of
the two animals that were enclosed together in the pen described above. And was
directly introduced in the laptop by means of a custom made program “Eventcounter
3.0” (by courtesy of Kristian Beedholm, Aarhus University). The animals were
observed from a height of 5m overlooking the experimental pool. As there were three
changes in the environment (se above), two non-visible options were also collected
together with the behaviours. In the first (floating pool and net) and third (floating pool
and net removed) environments, the animals were able to swim under the floating pool
and under the pontoons respectively, that not-visible time was collected for the mother
alone, the calf alone and mother-calf underneath. To be able to take out the not-visible
time for some of the analysis (see below).The 2h observations were started at sunset,
when the centre was closed, to be able to see the animals better with external lights out

5
of the pool, as during the day reflections did not allow to see the animals permanently,
and also in this way no external factors disturbed the animals’ behaviour.

Ethogram
In Tables 1 and 2 the ethogram or classification of behaviours that were
observed and quantified in the mother and calf harbour porpoises are shown. The spatial
relations of the mother-calf were defined as mother-calf together or individual alone
being mutually exclusive. The mother calf were considered together when they were in
physically contact, or being close to one another (within one body length) for more than
5 seconds. The 5 seconds arbitrarily defined as the minimum time for the observer to be
able to record all the spatial distances. The animals were considered alone when being
at a distance greater than one body length. They were considered alone even if during
their different activities/directions they passed by one another if not staying longer than
5 seconds within one body length.
The most well described and common positions that dolphin calves maintain
towards their mothers are the so called echelon position, and infant position (e.g.
McBride & Kritzler 1951; Tavolga & Essapian 1957; Amundin 1986) (see Table 2 for
definition). In this study we have distinguished different activities of the mother while
the calf is in infant position. The behaviours have been classified according to their
function under the names and categories seen in Tables 1 and 2.
Table 1. Ethogram for individuals alone. Showing the spatial state, the functional category at which the behaviours have been
assigned, the name and description of the behaviours observed.
Spatial
Category Behaviour Description of behaviour
state
Swimming at a distance greater than one body length in different direction of other animals during
Swimming
more than 5 seconds.
RESULTS Resting “logging on the surface with the blowhole out of the water and the tail bent ventrally”
Resting (Andersen & Dziedzic 1964) at a distance greater than one body length from another animal.
Swimming at very high speed, slightly leaping and almost jumping out of the water, and
Fear-stress sometimes being submerged for a longer time until the animal jumps, all at high speed (Andersen
& Dziedzic 1964; Amundin & Amundin 1971).
Coughing Breathing explosive and violently, producing a deep hoarse sound.
Scanning the sand and rocks of the shore probably searching for food (crabs, fishes, ….), the
INIDIVIDUAL ALONE

Bottom grubbing animal stays upside down, with the head pointing the bottom, of the pen in this case, and the tail
towards the surface (Amundin & Amundin 1974)
The animal scanned a separation net, it is probably scanning in the search of food as in bottom
Wall scanning grubbing; but could also be sometimes acoustical or even visual communication with the other
two porpoises in the contiguous pen.
An animal is specifically observed chasing a fish, with quick and directed movements, changing
Playing/feeding

Fishing direction at higher speed than normal swimming. It was not possible to observe if the fish was
cached or swallowed.
A porpoise turns and swims towards another porpoise (the calf towards the mother in this case),
Approaching-
and spins approaches and escapes in front of her, the mother not reacting by leaving, linking it
turning-spinning with a playing function (not aggressive).
Carrying objects around with the mouth, splitting them out and re-catching them. Usually diverse
fauna and flora (leafs, crabs, mussel shelves), crabs and mussels after bottom grubbing; it might
be that the content was eaten, as the empty shelves have been found by the trainers (Wahlberg
Object handling personal comment). Sometimes the object was not observed but the pattern was identified because
of the calf directing the
6 head towards the surface, splitting water (probably the object) out of the
water, swimming around with open mouth, floating at the surface with the mouth open, like
chewing, splitting and re-catching something.
Table 2. Ethogram of the mother-calf together. Showing the spatial state, the functional category at which the behaviours have
been assigned, the name and description of the behaviours observed.

Spatial state Behaviour Description of behaviour Category


The calf swims under the tailstock of the mother keeping pace with her momentum, placing the
Swimming top of its head just touching the posterior extremity of her abdomen. Called infant position and
underneath described in bottlenose dolphin related with nursing and sleeping (e.g. McBride & Kritzler 1951;
Tavolga & Essapian 1957; Reid et al. 1995).
Fear-stress The calf swims in infant position, but at high speed (keeping the same momentum as the mother),
underneath as a response to a fearful or stressful situation.
Calf underneath the mother

Fishing The mother fishes while the calf stays close by, usually trying to keep the infant position under
underneath her.
Bottom
The mother bottom grubs and the calf stays close, trying to maintain infant position under the
grubbing
cow.
underneath
Net scanning The mother scans the net and the calf stays underneath her, maintain infant position under the
underneath cow.
The calf is attached to one of the mother mammaries and presumably nursing. Nursing is
Nursing equivalent in this study as each time the infant is locked to one teat of her mother (lockons). The
calf was always seen to roll and get attached to one mammary with its belly up.
“The calf underneath the mother repeatedly raises and lowers its head contacting the mammary
Bumping region” (in belugas e.g. Recchia 1994), also more intense rubs of the mother’s belly were

AFFILIATIVE
performed by the calf with the head, only observed during resting underneath.
Resting The mother rests logging at the surface (Andersen & Dziedzic 1964) while the calf stays under
underneath her in infant position. Presumably both rest.
The calf swims by the side of the mother in close proximity, sometimes touching, on one side of
Echelon the cow slightly over her trunk, just below her dorsal fin Providing the calf hydrodynamic
swimming advantages. Observed to breathe in synchrony. Observed in dolphins (e.g. McBride and Kritzler
MOTHER-CALF TOGETHER

1951, Tavolga & Essapian 1957).


Mother and calf beside each other

Beside Resting “logging on the surface with the blowhole out of the water and the tail bent ventrally”
resting (Andersen & Dziedzic 1964) near another animal, at a distance no greater than one body length.

Two animals swim beside each other, not in contact – at a distance no greater than one body
length – in the same direction during > 5seconds, usually surfacing in synchrony when breathing.
Beside
Described by Andersen & Dziedzic (1964) as companionship between male-female harbour
swimming porpoises giving it a sexual function. It has been defined as well in many delphinids with a social
affiliative function (e.g. Perelberg & Schuster 2008).
Stroking of two porpoises “by swimming one by another, where the passer-by touches the other’s
dorsal fin or side” the other porpoise contributes by slowing down its speed (Amundin &
Grooming Amundin 1971). Grooming tends to occur in bouts, where both animals groom each other.
Grooming was considered in this study as grooming bouts.

Beside fear- The animals swim beside each other (same definition as above) but at high speed, jumping at the
stress surface, as a response to a stressful-fear reaction.

An animal twists horizontally its body 90 degrees staying with the belly towards the surface, or
Turning 180 degrees coming back to the initial position. Only the mother was observed to do this when
EVASIVE

the calf was below her.


An animal swims away from another at high speed, trying to avoid the contact with the other.
Escaping- Only the mother escaped from the calf, when the calf was underneath her. As a response of the
following escape, the other animal swims after the one swimming away, trying to get close also swimming
at high speed. Only the calf followed the mother trying to achieve infant position again.
Sideward turn: Snouting towards another animal turning the head and frontal part of the body
with closed and less often open mouth towards the head of the threatened porpoise; having an
aggressive function (Amundin & Amundin 1971). This aggressive behaviour has a duration of
AGGRESSIVE

about 1second (Amundin & Amundin 1971).


Pushing: The sideward turn ended in the aggressor touching the other animal with its snout.
Supposed to be a more intense nature of aggression (Amundin & Amnundin 1971) also described
Aggression by Andersen & Dziedzic (1964).
Chasing: Aggressive interaction, where one animal chases another one in a high speed, proceeded
by a sideward turn or a pushing, or ended by them. Not to be confused with escaping-following,
where the animal following does not intend to be aggressive as in chasing. And not to be
confounded with the chasing described by Andersen and Dziedic (1964) during sexual encounters
as the animal following was also not aggressive.

7
Some behaviours (nursing, grooming, evasions and aggression, as well as the
spatial position, i.e. the animals together or alone) were observed for 10 months.
However the rest of the behaviours were observed the first 9 months, and not observed
the last month of study.
The duration of sideward turn, one type of aggression was counted to be of 1
second, as that is the average duration of the behaviour according to Amundin &
Amundin (1971). Turning was also seen to have a duration shorter than 5 seconds most
of the times. Nursing usually had a duration shorter than 5 seconds. The duration of
coughing was not measured, only counts of number of events and presence or absence
of a motor-boat were collected. A “control” of motor-boat presence, to contrast with
coughing-motor boat, was not done, as not all the motor boats affect the animals, the
properties of the sound produced by the boat explains that. The presence of motor-boats
when the animals were scared or coughing was collected along the 10 months of study.

Data Analysis
The data was sorted by means of custom made programming with Matlab (ver.
6.2, MathSoft, Inc.) and Excel (ver. 2000, Microsoft Corporation). And the statistical
analyses were done by means of SAS (ver. 9.1.3. SAS Institute Inc.). The different
types of data were tested for Normality distributions (Goodness-of-Fit Tests for Normal
Distribution, SAS) and for homogeneity of variances (Levene’s Test for Homogeneity
of Variances, SAS) to apply parametric or non-parametric tests. The analysis were
chosen consulting Dytham (2003), Fowler et al. (1998) and Zar (1999).

Accumulated duration of behaviours or spatial distances along time


The behaviours presented in this way are the ones that were observed the 2 hours
observation per day (i.e. the behaviours that were only observed to be performed by the
mother and calf together).
Accumulated time together-alone: The data was divided depending on the
spatial state (i.e. when the mother or the calf were alone, and when they were together).
These observations were done during 10 months. The analysis was done summing in the
time together and alone within the 2 hours observation day. The non-visible time was
not taken out, assuming continuity of the state previous to swimming under the cage or
the pontoons until being visible again. This was done not to break artificially the
important developmental measure of time together or alone.

8
Accumulated duration of behaviours: Some behaviours were observed for 10
months (grooming, nursing, avoidances) and some for the first 9 months (beside
swimming, beside resting) (see methods).
The accumulated duration of behaviours and time-together alone were analysed
as follows. First, the accumulated duration of behaviours or time together-alone was
analysed for differences among months. By means of One-way ANOVA test with
posterior pairwise comparisons with Ryan-Einot-Gabriel-Welsch Multiple Range Test
(SAS), if the data was normally distributed and with homogeneous variances; or by
means of the non-parametric Kruskall-Wallis test (SAS) if not normally distributed.
Second, if significant differences were seen along time (among months) in the previous
analysis, the 10 or 9 months of study were divided in two periods. The period division
was moved along the 10 or 9 months (9 or 8 analyses were made moving the division of
the two periods, for instance, the first month versus the rest of the months, the first two
months versus the other months, etc) to find a significant difference between the two
periods, a breaking point along time. This second kind of analysis was done by means
of a One-Way ANOVA (SAS) tests if the data was normally distributed. For non-
normal data we a non-parametric Mann-Whitney test was used (Wilcoxon Two-Sample
Test in SAS). Bonferroni corrections were applied to the obtained p-values (with either
ANOVAS or Mann-Whitney tests) to correct for testing several times on the same data
(P’= P * number of comparisons). However, if not significant differences were found
among months (first analysis), the data was tested for a correlation with time, to check
for an increasing or decreasing tendency of co variation with time. This was done by
means of the Spearman Rank Correlation Test (SAS) as applied to not normally
distributed data. Finally, if some behaviours seemed to vary similarly along time, were
tested for a correlation between behaviours, done as well by means of the Spearman
Rank Correlation Test (SAS).

Percentage of time in activities of mother alone, calf alone, and together


The behaviours presented in this way are the ones that were observed performed
by the calf alone, and by the mother alone, and some behaviours as well by mother-calf
underneath. This behaviours have been transformed to percentages to be able to
compare between the behaviour when done by the mother alone, calf alone or mother-
calf underneath, as each condition was observed for a different duration of time.

9
The behaviours performed by the animals alone were observed for 1 hour per
day each animal. However the behaviours performed by mother-calf underneath were
observed for a total of 2 hours. However, in this 1hour each animal was observed they
did switch from being together to being alone, and therefore the total time in each
condition varied depending on the animals’ choice. Therefore, to be able to compare
among the accumulated duration of each behaviour performed by the mother alone, the
calf alone or the mother-calf underneath needed to be standardized as a percentage of
the observed time in each condition per day. Additionally, in two of the environments
(some months) the animals were non-visible a percentage of the time. This non-visible
time was taken out from the total observed time in each condition (as it was written
down if each animal was not visible alone or both as a couple).
A percentage of the accumulated time inverted in different behaviours by the
mother, the calf and both as a couple was obtained from the total observed (visible) time
each day in the three conditions and it is the one analysed and presented in the plots.
Only the first 9 months of data were considered for comparisons. Some of the
behaviours were only observed during 9 months.
All this behaviours: swimming, resting, fishing, object, bottom grubbing, wall
where performed in the three conditions (mother and calf alone, and mother-calf
underneath), were tested for differences. First, for differences within behaviour
depending on the actor/position considering all the study period as a whole. And
second, for differences within behaviour depending on the actor/position along time.
For the first type of analysis (all the studied period as a whole), each behaviour
was tested for differences depending on the actor by means of One-Way ANOVA tests
with posterior multiple pairwise (Ryan-Einot-Gabriel-Welsch Multiple Range Test) for
the normally distributed with homogeneous variances’ data. If significant deviations
from normality and homogeneous variances were found, Kruskall Wallis tests were
used instead (using SAS, ). If significant differences were found then posterior pairwise
comparisons were done by means of Mann-Whitney tests (SAS) to check for specific
pairwise differences among actors/positions.
For the second type of analysis (behaviours along months), each behaviour and
condition (calf alone, mother alone, or mother-cal together) was tested for differences
among months. With the first and second type of analysis used for the “accumulated
duration of behaviours or spatial distances along time” type of data. Except for wall
scanning that an extra analysis (Mann-Whitney test) comparing each month against

10
mother and calf wall scanning percentage, to reject a similar change along time in both
animals.

Mean and median duration of behaviour events


The duration of the first and last behaviour events of each sampling interval (15
minutes) were not taken into account to not include durations of behaviours artificially
divided by the sampling method. All the studied period was considered as a whole.
The duration of the behaviour events whose duration could have biological
significance is presented. Only the duration of certain behaviours was measured (see
methods above). The duration of the behaviour events whose duration could have
biological significance is shown. The duration of the diverse behaviours was classified
differentiating the type of behaviour, the actor or their relative position (calf alone,
mother alone; mother-calf underneath, or mother-calf beside each other). The data used
comprises all the durations of behaviours events in the 10 month study. Each kind of
behaviour was statistically tested for differences among actors and their relative
position. The data was not normally distributed, and therefore a non-parametric Mann-
Whitney test was used to compare pairwise, positions or actors in the different
behaviours (Wilcoxon Two-Sample Test in SAS). To correct for multiple comparisons
on the same data, the Bonferroni corrections was applied to the P-values (P’= P *
number of comparisons).

11
RESULTS

DESCRIPTION OF BEHAVIOURS NOT PLOTTED IN GRAPHS


Information on the fear-stress reaction (see methods), was collected along the
ten-month study period. It was observed only simultaneously with the close pass of
motor boats by the harbour, beside the pen where mother and calf were enclosed. This
demonstrates that sounds made by boats can scare the animals, probably due to pain or
startle caused by the sound. A differentiation of a reaction of fear by the mother or the
calf alone, and a reaction of the mother-calf pair beside each other was done initially to
check whether the calf tried to keep close to the mother in those moments or if it stayed
by its own. It was observed that when the fear/stress started the animals started
swimming at a high speed by their own, the calf was not able to keep a constant
momentum with the mother. They would pass nearby each other, underneath, beside
and at a distance greater than one body length in a quickly and random way, until the
level of stress decreased, then the calf swam underneath the mother. This showed that
the calf achieved a position underneath the mother at the end of the episode, but whether
they would have stayed in close proximity in the wild during the most stressful moment
could not be predicted. However, they did try to find each other after the fear reaction
was over.
Coughing, which could initially be just a physiological response to a respiratory
obstruction, was only observed in the mother or calf simultaneously with the fear
reactions and the pass of motor boats. This shows that coughing, not only explosive
breathing but the production of a deep hoarse sound, is a fear/stress response. Coughing
was not performed every time a fear-stress response to the pass of a motor boat was
seen, which could reflect that it is a more intense nature, or a different one, to show
fear-stress.
Echelon swimming was included in the ethogram, due to the hypothesis obtained
from other marine mammals. It was never observed in the study period from the 3rd
month of calf age to the 10th. In some videos of the mother-calf of the week posterior to
the birth, echelon swimming was observed (personal observation). This shows that this
behaviour did occur for the species but would have disappeared by the time of the
beginning of the study (3rd month).

12
ACCUMULATED DURATION OF BEHAVIORS AS A FUNCTION OF AGE

Time together-alone accumulated duration

7000 Alone
Together
6000 A AB AB
ABC
Total time 2h (seconds)

ABC
5000 ABCD ABCD
4000 BCD CD
3000 D

2000

1000

0
3 4 5 6 7 8 9 10 11 12
Months (calf age)

Figure 1. Accumulated time together-alone in each observation day, averaged per month. Standard error
of the mean, as error bars. The months with all the letters different significantly differ from each other.

The total time per observation (2h) mother and calf spent together-alone
averaged per month is shown in Figure 1. A decrease is seen along the period, and
significant differences are found between months (One-Way ANOVA, P<0.001).
Posterior pairwise comparisons (with Ryan-Einot-Gabriel-Welsch Multiple Range
method) showed not clear differences between all the months. Significant differences
among all the months are indicated by different letters in Figure 1; For instance, in the
3rd month the total time together-alone
significantly differs form the mean time
together-alone in the 10th, 11th and 12th
month, etc. Being the 3rd month of calf age
the one where the duration of time spent
together was longest, and having the 10th
the shortest amount of time together, being
this two months significantly different
form each other.
Another analysis was done on the Figure 2. Average accumulated time mother-calf spent
together in the two significantly different time periods. The
total time spent together by mother and period A comprising: 3, 4, 5, 6, 7, 8 and 9 months of infant
age; and the period B comprising: the month 10, 11, and 12.
The error bars correspond to the standard error of the mean.

13
calf, to try to find a clearer breaking point. The developmental period studied (10
months), was divided into two (several times), setting the time division on all months
and testing for differences among the first (A) and second (B) time interval (Several
One-Way ANOVA with Bonferroni correction). There was only one significant
difference between A and B, when the time line was set between 9 and 10 months of
calf age (Figure 2). Therefore, this shows there is a significant decrease in the time
mother and calf spend together at 10 months of age. In Figure 2 the mean duration of
the time mother and calf spent together in the first period (A, form the 3rd to 9thmonth)
and the significantly different second period (B, from the 10th to 12th month) is
presented.
A significant negative correlation was found with month (Spearman Rank
Correlation Test, r= -0.30, P= 0.03), which shows that time together decreased along the
study period.

Grooming accumulated duration


1200
Accumulated duration
per 2h observation

1000
(seconds)

800

600

400

200

0
3 4 5 6 7 8 9 10 11 12
Months

Figure 3. Accumulated duration of grooming bouts per hour, averaged per month. The duration of
grooming is shown along the studied period. The error bars correspond to the standard error of the mean.

Accumulated grooming time per observation (2 hours) averaged per month is


shown in Figure 3. No significant differences in the accumulated duration of grooming
were found among the months due to the great variation within each month.
However, comparing the total time the mother and the calf spent together
(Figure 1) and the total grooming time (Figure 3) it can be seen that the shapes seem
similar, and greater values are seen in the 3rd , 7th and 9th month of calf age. A positive
significant correlation was found between grooming and time together (Spearman Rank
Correlation, r= 0.73, P<.0001). The 53% (r2= 0.53) of the variation of grooming time is
accounted by variation in time together, and inversely.

14
Beside swimming and beside resting accumulated time per observation (2 hours)
is presented in Figure 4 averaged per month. No significant differences were found by
month, in beside swimming accumulated, nor in beside resting accumulated time.

Beside swimming accumulated duration Beside resting accumulated duration


300

Accumulated duration
500
Accumulated duration

per 2h (seconds)
per 2h (seconds)

400
200
300

200 100
100

0 0
3 4 5 6 7 8 9 10 11 3 4 5 6 7 8 9 10 11
Months Months

Figure 4. Accumulated duration of beside swimming and beside resting per hour, averaged per month. The duration of both
behaviours is shown along the studied period. The error bars correspond to the standard error of the mean.

However, again correlations are found. There is a significant positive correlation


between beside resting and beside swimming (Spearman Rank Correlation Test, r=0.31 ,
P= 0.03). And both, are significantly and positively correlated with grooming
(Spearman Rank Correlation Test, r=0.45 , P=0.0007 for beside swimming and r=0.51,
P=0.004 for beside resting) and time together (Spearmen Rank Correlation Test,
r=0.43, P= 0.001 for beside swimming and r=0.47, P= 0.0004 for beside resting). This
shows that to some grooming and time together vary together, and that all the above-
mentioned variables are positively related. Additionally, time-together, beside resting
and beside swimming are negatively correlated with months (Spearmen Rank
Correlation Test, r=-0.40, P=0.003 for beside swimming and r=-0.40, P= 0.003 for
beside resting). This shows that the accumulated duration of these behaviours decrease
along the study period.

In Figure 5 the accumulated duration of evasive and aggressive behaviours from


the mother towards the calf, showing the proportion of each kind of avoidance are
shown per observed day during the 10-month study. These evasive and aggressive
behaviours: turning and aggression, were only observed from the mother towards the
calf, never in the other direction. The mother was the only one observed escaping from
the calf when the calf was underneath her, and as response immediately after the calf
followed her. Tuning and aggression were also seen when the calf was underneath the
mother.

15
Accumulated duration of avoidances from the mother towards the calf
500
A B

A B
400 A B
Total time in 2h (Seconds)

300 aggression
escaping-following
turning
200

100

0
3 4 5 6 7 8 9 10 11 12
Months

Figure 5. Accumulated duration of avoidance in each observed day (2h per day). The proportion of each
specific type of evasion to the total avoidance duration is shown per day. Three possible significant
divisions of the studied period in two are shown with different letters on the arrows on top, the 3 possible
divisions marked by the three doted lines. Further explanations see text

When testing for differences among months of the accumulated duration of


avoidances, significant differences were found among at least two months (Chi29 = 19.6,
P= 0.02). Therefore, the study period was divided into two, setting the division in each
month (as done with time together-alone, see methods), and the significant results were
found when dividing the study period between the 6th and 7th month (Mann-Whitney
Wilcoxon test: z = -3.0, P’= 0.02); when dividing the study period between the 7th and
8th month (Mann-Whitney Wilcoxon test: z = 3.5, P’= 0.005) and when dividing the
study period between the 8th and 9th month (Mann-Whitney Wilcoxon test: z = 3.6, P’=
0,003). The most significant result being when setting the division between the 8th and
9th month.
The same procedure, but considering as evasions only aggression and escape-
following behaviours. The same results were found about the period breakpoint.
Significant results were found when dividing the study period between the 6th and 7th
month (Mann-Whitney Wilcoxon test: z = 5.0, P’<0.0009); when dividing the study
period between the 7th and 8th month (Mann-Whitney Wilcoxon test: z = 4.4
P’<0.0009) and when dividing the study period between the 8th and 9th month (Mann-
Whitney Wilcoxon test: z =2.9, P’<0.0009). And the p-values showing a highly
significance level, bigger than when using turning as well. This shows that aggression

16
and escape-following do reflect a more intense nature of active avoidance. These results
show altogether that avoidance from the mother towards the calf starts/increases rapidly
from around the 7-8th and definitively in the 9th month of calf age.
Additionally the total duration of avoidances (turning, escape-following and
aggression grouped together) were significantly and positively correlated with day
(Spearmen Rank Correlation Test, r=0.53, P<0.0001). This shows that avoidances
increased with time. If only aggression and escape-following are summed in, and
turning is kept out as done before the correlation of avoidances (aggression and escape-
following) with time, in this case month, is a better correlation than if including turning
(Spearman Rank Correlation Test, r=0.55, P<0.0001).
No correlation was found between avoidances and any of the other behaviours
described above: time together-alone, grooming, beside resting nor beside swimming.

Bumping accumulated duration


1200
Accumulated duration

1000
per 2h observation

800
(seconds)

600
400

200
0
3 4 5 6 7 8 9 1 1
Months

Figure 6. Accumulated duration of bumping in each observed day (2h per day). The duration of bumping
is shown along the studied period.

Bumping accumulated duration per observation day is presented in Figure 6. It


can be seen that this behaviour was not commonly observed, seen only 8 days from the
50 observed. It is not correlated with evasions and aggressions (Spearman Rank
Correlation Test), even though it is presented mostly at the end of the period. And if the
total studied period is divided in two, there are significant differences between periods
setting the division in the 8th (Mann-Whitney Wilcoxon test: z =2.8, P’=0.05) and 9th
month (Mann-Whitney Wilcoxon test: z =3.2, P’=0.01).

17
This behaviour was probably difficult to observe and the results presented here,
probably an underestimation of the real amounts, this observations accounted for the
most obvious performances.

Nursing accumulated duration


150
Accumulated duration

125
per 2h observation

100
(seconds)

75
50
25
0
3 4 5 6 7 8 9 10 11 12
Months
Figure 7. Accumulated duration of nursing per hour, averaged per month. The duration of nursing is
shown along the studied period. The error bars correspond to the standard error of the mean.

Nursing accumulated duration along time is presented in Figure 7, averaging


durations per month. There were no significant differences among months in all the
observed period (One-Way ANOVA test). A more thorough analysis on the nursing
data will be presented in another paper.
In Figure 8, the accumulated duration of the accumulated time the animals were
not visible during the observations is represented in the three environments.

Total time not visible together


1400
Accumulated time in 2h

1200
1000
(Seconds)

800 under floating pool


600 under pontoons
400
200
0
3 4 Total5time not6visible7 mother and
8 calf alone9 10 11 12
Months
1400
Accumulated time in 1h

1200
1000
(Seconds)

calf under floating pool


800 calf under pontoons
600 mother under floating pool
400
mother under pontoons

200
0
3 4 5 6 7 8 9 10 11 12
Months

Figure 8. Accumulated not visible time in the three environmental conditions (with floating pool and
net, without floating pool but net, without net) and in the three spatial positions of the animals
(mother alone, calf alone, and mother-calf together).
18
A floating pool present from the 3rd to the 5th month (environment 1), from the
5th the floating pool was removed (environment 2), and from the 8th the inner net which
made the pen smaller and the access under pontoons not possible, was removed
(environment 3). In the first environment the animals could swim under the floating
pool, where they were not visible to the observer. In the second environment they were
permanently in sight. In the third environment they could swim under the pontoons,
being the not-visible space larger than the floating pool. The non-visible amount of time
was taken out to analyse the duration of some behaviours (see Methods).

PERCENTAGE OF TIME IN ACTIVITIES OF MOTHER ALONE, CALF ALONE,


AND TOGETHER

Differences among animals or positions


The time budget of the calf when alone, of the mother at her time alone, and of
the mother with the calf together, are shown in Figure 9.
The percentage duration of behaviours that were performed by the two animals
alone and/or the mother-calf together were tested for statistical differences due to the
actor, and statistical results are shown in the plots. All the studied period was
considered as a whole (showing differences among actors/positions).
All the behaviours performed by the calf alone, and the mother alone, were
shown in the time budget made with the percentage of time observed (see methods) for
each of them. Some behaviours presented in the plots were not tested, where the
behaviours were only performed by one animal or by the mother-calf underneath (only
one actor or spatial position), or that differences among animals were not due to the
animal (for instance fear). The fear reactions were observed in the mother alone, in the
calf alone, and in mother-calf underneath; however, the differences in percentage did
not depend on the animal but on the amount of motor-boats passing by, being that
amount different in each condition observed. Both animals were scared the same
amount of times by the same motor-boats.
In the first environment, where the animals had a floating pool inside the pen,
the animals could freely swim under it or inside it. The mother, as seen in Figure 8,
swam underneath the pool, and inside it (Figure 9). The calf never swam inside it.

19
The percentage of swimming was significantly different (One-Way ANOVA,
F=40.7, P <0.0001). Swimming was significantly longer when the mother swam alone
than when the mother-calf swam together and when the calf swam alone.

Mother alone time budget Calf alone time budget

A A
A
A

A
A

C
A B

A
B

sw imming resting fear


sw imming resting fear
bottom grubbing object w all bottom grubbing object w all
fishing cage escaping fishing follow ing

Mother-calf underneath budget

B
B

swimming resting fear bottom grubbing


wall fishing beside fear beside sw
beside rest grooming nursing turning
bumping

Figure 9. Time budgets of the mother alone, the calf alone and mother-calf together. Percentage of time inverted in the
different behaviours by the calf alone, the mother alone, or mother with the calf underneath. Behaviours with different
letters in the time budgets significantly differ from the same behaviour performed by another animal.

20
The percentage of swimming was significantly shorter for the calf swimming
alone than for the mother-calf underneath (Ryan-Einot-Gabriel-Welsch Multiple Range
Test, in the ANOVA test), Figure 9 shows this in the sectors with light blue colour
The percentage duration of resting did also differ depending on if resting was
done by the calf alone, the mother alone or the mother-calf underneath (Kruskall Wallis
test, Chi29 = 65, P <0.0001). Resting was significantly longer when the mother-calf
underneath were resting than when the calf rested alone (Wilcoxon Two-Sample Test,
z= -5.6, P’<0.0003) and when the mother rested alone (Wilcoxon Two-Sample Test, z=
-3.7, P’=0.006). Resting was significantly shorter when the mother was resting alone
than when the calf was resting alone (Wilcoxon Two-Sample Test, z= -5.6, P’<0.0003).
See Figure 9 shows this in the sectors with purple colour.
Bottom grubbing, fishing and wall did also differ if performed by the mother, the
calf and both together (Kruskall Wallis test: Chi29 =27, P <0.0001 for bottom grubbing;
Chi29 = 20, P <0.0001 for fishing and Chi29 = 23, P <0.0001 for wall).
The percentage of time inverted in bottom grubbing by the mother was not
significantly different from the percentage of bottom grubbing by the calf. But both
mother alone and calf alone spend a significantly bigger proportion of time bottom
grubbing than mother-calf together (Wilcoxon Two-Sample Test: z= 5.1, P’<0.0003
mother alone vs. mother-calf together; z= 3.1, P’<0.0063 calf alone vs. mother-calf
together). Figure 9 shows this in the sectors with turquoise blue colour.
The percentage of time inverted in fishing by the mother was not significantly
different from the percentage of time fishing by the calf. But both mother alone and calf
alone spend a significantly larger proportion of time fishing than mother-calf together
(Wilcoxon Two-Sample Test: z= 3.0, P’<0.007 mother alone vs. mother-calf together;
z= 4.1, P’<0.0003 calf alone vs mother-calf together). Figure 9 shows this in the sectors
with dark blue colour.
And as in bottom grubbing and fishing occurs in wall; the percentage of time
inverted in wall by the mother was not significantly different from the percentage of
wall by the calf. But both mother alone and calf alone spend a significantly bigger
proportion of time in the wall than mother-calf together (Wilcoxon Two-Sample Test:
z= 3.1, P’<0.006 mother alone vs. mother-calf together; z= 4.7, P’<0.0003 calf alone vs.
mother-calf together). See Figure 9 shows this in the sectors with pink colour.
Object handling was performed only by mother or calf alone, but never observed
in mother with the calf underneath. The proportion of time inverted by the calf during

21
all the period studied in handling an object was significantly higher than the proportion
of time that the mother had an object on her mouth (Wilcoxon Two-Sample Test: z = -
6.9, P’<0.0001). Figure 9 shows this in the sectors with dark purple colour.

Differences along time among animals or positions


The percentage of time for the different behaviours as a function of time by the
mother, the calf and the mother with the calf underneath is presented below. As changes
during the 10 months in the calf’s behaviours could not only be developmental, but the
environment will probably influence them the behaviour of the mother could be
interpreted as a “control”, therefore calf’s behaviours differing form the mother pattern,
are more likely that changes are due to age-related reasons. In the following plots, an *
is present when there are significant differences along time in that behaviour performed
by that actor.

Mother alone fishing


40 *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 Calf6alone fishing
7 8 9 10 11
40 Months *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 6 7 8 9 10 11
Mother-calf underneath fishing
Months
40
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 6 7 8 9 10 11
Months

Figure 10. The percentage of time fishing per month during the studied period by the calf alone, the
mother alone and the mother with the calf underneath are shown. Plots with an asterisk on the upper
right side have significant differences in the behaviour among months for the actor of the plot.

22
In Figure 10 the percentage of time fishing per month during the studied period
by the calf alone, the mother alone and the mother with the calf underneath are shown.
The percentage of time the calf was fishing alone was tested for differences
among months and significant differences were found between at least two months
(Kruskall Wallis test, Chi29 =24, P = 0.002). Also the percentage of time the mother was
fishing alone was tested for differences among months and significant differences were
found between at least two months (Kruskall Wallis test, Chi29 = 22, P = 0.006). In
contrast, no significant differences were found among months in the percent of time
fishing together.
As it has been detailed above (Figure 9) there were no significant differences in
the proportion of time the mother and the calf spend fishing alone considering all the
studied period together. However we see that there are significant differences among
months and therefore to conclude the mother and the calf spend fishing the same
percentage of time, we need to see if there are significant differences between mother
and calf fishing at diverse periods. As observed in the mother plot (Figure 10) there is
an increase in the 7th month of calf age, and differences among mother and calf could
arise at that point, but the period before could have diminished the differences, as
almost no fishing occurs (i.e. divide the studied period in parts to test for differences
among periods instead of only testing all the study period as a whole).
The study period was divided between the month 6th and 7th and the percentage
of time the calf spends fishing was tested for differences between this two periods. After
the 6th month the calf significantly fishes more than before the 7th month (Wilcoxon
Two Sample test z= -3.8, P= 0.0001). Significant differences were found at the same
point for the mother fishing alone (Wilcoxon Two Sample test z=-2.3, P= 0.02), being
fishing behaviour greater after the 6th month. Therefore, the periods defined by the 7th
month were tested for differences between mother and calf. The period pre-seventh
month was not significantly different between mother and calf (Wilcoxon Two Sample
test z=0.0, P= 1.0); and the period post-seventh month was significantly different
(Wilcoxon Two Sample test z=-3.1, P= 0.002).
This shows there are not overall differences in the percentage of fishing between
mother and calf, but there are significant differences after the 7th month, fishing
increasing in both animals, but significantly more in the calf than in the mother.

23
Mother alone object
45

Percentage of observed
40
35
30

time
25
20
15
10
5
0
Month 3 4 5 Calf6 alone object
7 8 9 10 11
Months
45 *
40
observed time
Percentage of

35
30
25
20
15
10
5
0
3 4 5 6 7 8 9 10 11
Months

Figure 11. The percentage of time object handling per month during the studied period by the calf
alone and the mother alone are shown. Plots with an asterisk on the upper right side have significant
differences in the behaviour among months for the actor of the plot.

In figure 11 the proportion of object handling time per month during the period
studied by the calf alone or the mother alone are shown.
The percentage of time the calf was handling an object alone was tested for
differences among months, and significant differences were found between at least two
months (Kruskall Wallis test, Chi29 = 34, p<0.0001). Also the percentage of time the
mother was handling and object alone was tested for differences among months and no
significant differences were found. When testing for differences among mother and calf
object handling considering the study period as whole (Figure 9) the calf spend a
significantly bigger amount of time handling an object than the mother did. This shows
(considering the mother as a control) that the calf spends much more time
playing/manipulating an object than the mother does, and this behaviour changes along
time in the calf are probably due to developmental causes.
If the study period is divided in two between the 6th and the 7th month, as done
with fishing, significant differences are found between the two periods (Wilcoxon Two
Sample test z=-3.8, P= 0.0001), being the percentage time of object handling greater
from the 3rd to the 6th month, than from the 7th to the 11th. The opposite is seen for the
prevalence of fishing by the calf.

24
In Figure 12 the proportion of bottom grubbing time per month during the period
studied by the calf alone, the mother alone and the mother with the calf underneath are
shown.

Mother alone bottom grubbing


40
35
observed time
Percentage of

30
25
20
15
10
5
0
3 4 5 Calf alone
6 bottom
7 grubbing
8 9 10 11
40 Months
35
observed time
Percentage of

30
25
20
15
10
5
0
3 4 Mother-calf
5 underneath
6 bottom
7 grubbing
8 9 10 11
40 Months *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 6 7 8 9 10 11
Months

Figure 12. The percentage of time bottom grubbing per month during the study period by the calf alone, the
mother alone and the mother with the calf underneath are shown. Plots with an asterisk on the upper right side
have significant differences in the behaviour among months for the actor of the plot.

The percentage of time the calf was bottom grubbing alone was tested for
differences among months and no significant differences were found. The percentage of
time the mother was bottom grubbing did also not provide any significant difference
among months. In contrast significant differences were found among months in the
percent of time mother-calf underneath bottom grubbed (Kruskall Wallis test, Chi29
=32, P <0.0001). As we have detailed above (Figure 9) there were no significant
differences in the proportion of time the mother and the calf spend bottom grubbing
alone considering all the studied period together (Figure 9). Which shows bottom

25
grubbing is a behaviour performed similarly during all the studied months by the
animals alone.
In contrast, the prevalence of this behaviour by the mother with the calf
underneath differs from the prevalence of the behaviour by her when alone, and from
the one of the calf alone (Figure 9). In addition it changes along time. This could be
explained by a developmental process. In Figure 12 it can be seen that from the 8th
month bottom grubbing underneath becomes almost absent. The studied time was
divided between the 7th and 8th month and the two period tested for differences. In the
period form the 3rd to the 7th month the percentage of mother-calf underneath bottom
grubbing was significantly higher than the percentage of bottom grubbing by the
mother-calf in the period from the 8th to the 11th month (Wilcoxon Two Sample test z=-
4.2, P<0.0001)

Mother alone wall


40 *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 6 7
Calf alone wall 8 9 10 *11
Months
40 *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 Mother-calf
6 underneath
7 8 9 10 11
Monthswall
40 *
Percentage of observed

35
30
25
time

20
15
10
5
0
3 4 5 6 7 8 9 10 11
Months

Figure 13. The percentage of time wall scanning per month during the studied period by the calf alone,
the mother alone and the mother with the calf underneath are shown. Plots with an asterisk on the upper
right side have significant differences in the behaviour among months for the actor of the plot.

26
In Figure 13 the proportion of wall scanning per month during the period studied
by the calf alone, the mother alone and the mother with the calf underneath are shown.
The percentage of time the calf was wall scanning alone was tested for
differences among months and significant differences were found in at least two of the
months (Kruskall Wallis test, Chi29 =30, P =0.0002). The percentage of time the mother
alone was wall scanning did also provide significant difference among months (Kruskall
Wallis test, Chi29 =26, P =0.001). Significant differences were found as well among
months in the percent of time mother-calf underneath wall scanned (Kruskall Wallis
test, Chi29 =31, P <.0001). As detailed above (Figure 9) there were no significant
differences in the proportion of time the mother and the calf spend wall scanning alone
considering all the studied period together (Figure 9), as occurred with bottom
grubbing. This shows wall scanning is a behaviour performed similarly in all the
studied months (considered all the time together) by the animals alone, however it needs
to be contrasted if the differences along time between mother and calf wall scanning
alone, change similarly (then they are probably due to environmental changes, but not
developmental ones), to be able to conclude both animals alone behave in the same way
for this specific behaviour, differently from the conclusion about the bottom grubbing
behaviour (see above).
The performance of wall scanning by the mother with the calf underneath, that
differs form her own behaviour alone, and from the one of the calf alone (Figure 9) also
changes along time, but it needs to be tested if it does it in a different way as mother and
calf along time, and then it could be concluded that changes in wall scanning
underneath are developmental, as it was concluded with bottom grubbing.
Not a specific pattern was seen between percentage of wall scanning in mother
and calf alone; each month was tested against the same month for the other animal or
both together, and some of the months were significantly different depending on the
animal (Wilcoxon Two Sample tests), which shows that the three conditions (mother
alone, calf alone, and mother-calf underneath) do not vary along time in the same way,
do not follow the same pattern.
A breakpoint for the percentage of mother-calf underneath wall scanning seemed
to stand out as in bottom grubbing. The studied time was divided between the 7th and 8th
month and the two periods tested for differences. In the period form the 3rd to the 7th
month the percentage of mother-calf underneath wall scanning was significantly higher

27
than the percentage of wall scanning by the mother-calf in the period from the 8th to the
11th month (Wilcoxon Two Sample test z=-4.3, P<0.0001)

In Figure 14 the proportion of resting time per month during the period studied
by the calf alone, the mother alone and the mother with the calf underneath are shown.

Mother alone resting


40
35
30
Percentage of
observed time

25
20
15
10
5
0
3 4 5 Calf
6 alone resting
7 8 9 10 11
Months
40
Percentage of observed

35
30
25
time

20
15
10
5
0
Mother-calf underneath resting
3 4 5 6 7 8 9 10
40 Months *11
35
observed time
Percentage of

30
25
20
15
10
5
0
3 4 5 6 7 8 9 10 11
Months

Figure 14. The percentage of time resting per month during the studied period by the calf
alone, the mother alone and the mother with the calf underneath are shown. Plots with an
asterisk on the upper right side have significant differences in the behaviour among months for
the actor of the plot.

The percentage of time the calf alone and the mother resting alone were tested
for differences among months and among any of them significant differences were
found. In contrast significant differences were found in the percent of time mother-calf
underneath rested (Kruskall Wallis test, Chi29 = 18, P = 0.02).

28
As detailed above (Figure 9) there were significant differences if considering
resting over all the studied period, in the percentage of the calf resting alone, the mother
resting alone and the mother-calf underneath resting.
The variation of resting percentage for the calf alone during the period is not
significant, which defines the normal variation for the calf. The same occurs with the
mother, being the variation along time within the normal range of variation for her.
Being the percentage of time inverted in resting over all the period greater for the calf
alone than for the mother alone, but not changing significantly in any of them during
this time, which could reflect individual differences, but most probably age-related
differences, not varying for the calf to the mother level during the observed period.
On the other hand the percentage of mother-calf underneath resting does vary
along time, being significantly a higher proportion of time than for mother and calf
alone (Figure 9). It is necessary to take into account that it is the proportion of resting
for mother-calf underneath, from the total mother-calf underneath time what is being
considered here.
The percentage of time together was divided in two periods (at all the months)
and tested for a break point, or a increasing or decreasing difference but not significant
differences were found (Wilcoxon Two-sample tests, with Bonferroni correction for
multiple comparisons). No clear pattern was found in the percentage of resting mother-
calf underneath along time.
In Figure 15 the proportion of swimming time per month during the period
studied by the calf alone, the mother alone and the mother with the calf underneath are
shown.
The percentage of time the calf was swimming alone was tested for differences
among months and significant differences were found (One Way ANOVA test, F= 3.4,
P= 0.004). The percentage of time the mother alone was swimming did also provide
significant difference among months (One Way ANOVA test, F= 2.8, P= 0.01).
Significant differences were found as well among months in the percent of time mother-
calf underneath swam (One Way ANOVA test, F=5.1, P= 0.0002).

29
Mother alone swimming
100
*
90

Percentage of observed
80
70
60

time
50
40
30
20
10
0
3 4 5 6 7 8 9 10 11
Calf alone Months
swimming
100 *
Percentage of observed

90
80
70
60
time

50
40
30
20
10
0
3 4 5
Mother-calf 6underneath
7 swimming
8 9 10 11

100
Months *
Percentage of observed

90
80
70
60
time

50
40
30
20
10
0
3 4 5 6 7 8 9 10 11
Months

Figure 15. The percentage of time swimming per month during the studied period by the
calf alone, the mother alone and the mother with the calf underneath are shown. Plots with
an asterisk on the upper right side have significant differences in the behaviour among
months for the actor of the plot.

As explained above significant differences were found considering the period


overall among mother alone, calf alone and mother-calf underneath. It is necessary to
test if they vary similarly along time, or not, and specific periods account for the
variation among actors.
In the case of the mother the Ryan-Einot-Gabriel-Welsch Multiple Range Test
showed significant differences between the 3rd and 5th month in the percent of
swimming, when testing for differences between each month. The period was divided in
two to search for breakpoints and no significant differences were found dividing it in
two in any month.
In the case of the calf the Ryan-Einot-Gabriel-Welsch Multiple Range Test
showed significant differences between the 3rd and 4th month in the percent of

30
swimming, when testing for differences between each month. The period was divided in
two to search for breakpoints, only one significant result was found, when setting the
division between the 7th and 8th month (One-Way ANOVA test, with Bonferroni
correction, F=11.60, P’=0.0104). This shows that there are not strong differences along
time for the calf, however it can be seen a decrease of the swimming alone percentage
in the 4th month, and then an increase again in the 8th month. It is necessary to take into
account that it is the proportion of swimming for calf alone, from the total time spent
alone what is being considered here. Therefore Figure 15 shows the percentage of time
from the total spend alone, that the calf was involved in swimming, is slightly bigger in
the 3rd month, then decreasing and from the 8th month onwards slowly increasing again.
As in the mother there is a decrease in the first and second month of study, this change
in both animals similarly is probably related with environmental facts but not
developmental changes in the calf, as occur as well in the mother. The decrease exactly
in the 8th month could be due to the change in the environment (the nets were removed),
as the time the animal was visible could have been swimming, and performing other
behaviours under pontoons.
In the case of mother-calf underneath the Ryan-Einot-Gabriel-Welsch Multiple
Range Test showed significant differences between the 10rd and 7th, and 8th and 9th (the
late three not differing among them) months in the percent of swimming, when testing
for differences between each month. The period was divided in two to search for
breakpoints, only one significant result was found, when setting the division between
the 9th and 10th month (One-Way ANOVA test, with Bonferroni correction, F= 20, P’
<0.0008). It needs to be highlighted again, that this swimming mother-calf corresponds
to the percentage of swimming form the total time mother-calf were together. From the
absolute values of time together-alone (Figure 1) it is possible to see that the total time
together decreases at the 10th month, this increases in the percentage of swimming is
therefore not absolute. And means that the mother-calf underneath used less time doing
other behaviours than swimming, and swimming percent increased.

MEAN DURATION OF BEHAVIOURS AND INTERVALS


In Table 3 the mean, maximum and median durations of swimming, resting and
bumping are shown, performed by mother and calf in different spatial conditions, giving
some basis for further studies. And in Figure 16 the means of swimming, resting and

31
bumping are plotted, and statistical differences among the median duration in each
behaviour depending on the actors and their relative position are shown.

Tabe 3. Duration of swimming and resting when performed by the mother, the calf, or both as a unit in different positions. Duration
of the bumping behaviour. SD=Standard Deviation. SEM=Standard Error of the Mean. Max=Maximum. N=number of observed
behaviour events.
Alone Mother-calf Mean/Median duration
Calf Mother Underneath Beside 35 B Sw imming (n)
Swimming 30 A
C
(N) 1528 1255 3630 153 25
Mean 14.6 30.6 24.3 23.1 20
A
Median 10 20 17 10 15
SD 14.0 35.4 24.8 38.4 10
SEM 0.36 1 0.41 3.1 5
Max 134 300 254 308 0
40 Calf Mother D
Underneath Resting
Beside
Resting (N) 616 457 1980 66 35 Bumping
Mean 17.1 18.8 26.2 16.6 30 Alone C Mother-calf
Median 14 17 25 13 25 B
20 A AB
SD 14.6 11.7 13.5 11.4
15
SEM 0.59 0.55 0.3 1.4 10
Max 178 71 108 47 5
0
Mother-calf Calf Mother Underneath Beside
Underneath Alone Mother-calf
Bumping (n) 45
Mean 37.4 Figure 16. The mean duration of swimming and resting are
shown. Statistical differences depending on the actor or
Median 40.0
position were tested by behaviour and shown in the plot. The
SD 24 actor/positions with different letter significantly differ from
SEM 3.6 each other within behaviour.
Max 139

In Figure 16 the mean duration of swimming and resting by the calf alone,
mother alone, and mother-calf underneath or mother-calf beside, are presented.
Significant differences were found in the median duration of swimming
behaviour when the calf was swimming alone, the mother was swimming alone, or the
calf was swimming underneath the mother (Wilcoxon Mann-Whitney test, with
Bonferroni correction, P<0.001 in all comparisons). Additionally swimming duration
significantly differ when mother-calf were swimming beside each other, underneath, or
the mother alone (Wilcoxon Mann-Whitney test, with Bonferroni correction, P’<0.006
in all comparisons), but the calf swimming alone and mother-calf beside duration had
no significantly different duration. (Figure 16)
The duration of resting significantly differs when the actor is the calf, the mother
or both swim underneath (Wilcoxon Mann-Whitney test, with Bonferroni correction,
P’<0.002 in all comparisons). However the duration of resting when mother-calf rest

32
beside each other does not differ from when the calf rests alone or the mother rests
alone. The duration of resting mother-calf beside each other does significantly differ
from the duration of resting of calf underneath the mother (Wilcoxon Mann-Whitney
test, with Bonferroni correction, P’<0.001 in all comparisons; Figure 16).
The bumping behaviour (the calf rubs her head with her mother’s belly, see
methods for description) was only observed in occasions during resting. The median
duration of resting when the calf was rubbing her head towards the mother, bumping,
seemed to have a longer duration than normal resting. Bumping had a significantly
longer duration than resting and any of the other resting conditions — calf or mother
alone, beside or underneath mother-calf — (Wilcoxon Man-Whitney test, with
Bonferroni correction, P’<0,001 in all comparisons).

Table 4. Duration of bottom grubbing, wall scanning, fishing and object handling when performed by the mother, the calf,
or both as a unit in different positions. Duration of the bumping behaviour. SD=Standard Deviation. SEM=Standard Error
of the Mean. Max=Maximum. N=number of observed behaviour events. All the durations are in seconds.
Mean/Median duration
Alone Mother-calf 25 B
Calf Mother Underneath 20 Bottom grub
A
Seconds

C
Bottom grub (n) 278 367 356 15
Mean 17,05 21,51 14,6 10
Median 14.0 18.0 12.0
5
SD 11,83 15,15 10,06
SEM 0,71 0,79 0,53 0
30 Calf Mother Underneath
Max A
99 102 64 25 A Alone Together
Net
20
Seconds

Wall (n) 317 203 172 B


Mean 15
20,81 24,25 15,67
10
Median 17.0 18.0 12.5
5
SD 16,69 20,1 15,09
SEM 0,94 1,41 1,15 0
30 Calf Mother Underneath
A
Max 129 126 155 25 Fishing
Alone Together
Fishing (n) 259 74 45
Seconds

20
B
Mean 15
22,27 12,38 8,47 B
10
Median 15.0 8.5 6.0
5
SD 35,19 10,64 7,81
0
SEM 2,19 1,24 1,16 120 Calf Mother Underneath
Max 453 49 43 100 Alone Object
Together
Seconds

Object (n) 624 8 80 A


Mean 60
21,96 61,13 40 A
Median 16.0 13.5 20
SD 21,11 134,05 0
SEM 0,84 47,39 Calf Mother Underneath
Max
201 392 Alone Together

Figure 17. The mean duration of swimming and resting are


shown. Statistical differences depending on the actor or
position were tested by behaviour and shown in the plot.
The actor/positions with different letter significantly differ
33 from each other within behaviour.
In Table 4 the mean, maximum and median durations of bottom grubbing, wall
scanning, fishing and object handling are shown, performed by mother alone, calf alone
and mother-calf underneath, giving some basis for further studies. And in Figure 17 the
mean durations of behaviours are plotted, and statistical differences among the median
duration in each behaviour depending on the actors and their relative position are
shown.
The median duration of wall scanning does not significantly differ for mother
alone and calf alone. But the median durations for both (mother alone and calf alone) do
differ form the median duration of mother-calf underneath (Wilcoxon Mann-Whitney
test, with Bonferroni correction, P’< 0.0002 both comparisons; Figure 17, beige).
The median duration of bottom grubbing significantly differ when the actor is
the calf, the mother or mother-calf underneath, all differ from each other (Wilcoxon
Mann-Whitney test, with Bonferroni correction, P’< 0.0027 in all comparisons; Figure
17, green).
The median of the bottom grubbing events duration, significantly differs when is
performed by the calf or the mother (Mann-Whitney Wilcoxon test: z=-3.7 and
P’=0.0002) the mother or both as a unit (underneath), or in close proximity.
The median duration of fishing significantly differ when the actor is the calf and
the mother. And the mean duration of the calf fishing alone and mother-calf underneath,
also differ from each other (Wilcoxon Man-Whitney test, with Bonferroni correction,
P’< 0,02 in all comparisons; Figure 17, blue).
The median duration of object handling significantly does not differ significantly
when the actor is the calf alone or the mother alone (Figure 17, grey).
Table 5. Duration of nursing, turning, grooming and escaping-following. All the durations are in
seconds.
Together
Grooming (N) 362
Underneath
Mean 52.5
Nursing (N) 734
Median 32.5
Mean 3.72 SD 62.39
Median 3
SEM 3.28
SD 1.67
Max 415
SEM 0.06
Max 15
Turning (N) 88 Escaping-following (N) 61
Mean 5.14 Mean 10.16
Median 1 Median 8
SD 9.62 SD 10.46
SEM 1.03 SEM 1.34
Max 60 Max 55

34
The mean duration of the nursing, turning, grooming and escaping-following
behaviours only performed in specific positions. Nursing and turnings occurring when
the calf was underneath the mother. Nursing occurring in bouts, here were present only
the mean duration of each attachment to the mammary glands. Turning occurring when
the calf was underneath the mother. Its duration as presented here was usually short.
Escaping-following The mean duration of escaping-following had a longer
duration than turns, and it’s by its description a more intense form of avoidance.
Grooming is in contrast, and affiliative behaviour. Here we present the duration
of grooming bouts, which is one of the behaviour with the longest duration.

The time mother and calf stayed together, was distributed in bouts, the mean
duration of the intervals. The duration of the intervals together (in infant position) stays
constant along time. This indicates that the duration spent together is biologically pre-
defined for this species.
In Figure 18 the increasing maximum durations of intervals alone are plotted
along time. It can be seen that they progressively increases along time, varying from a
maximum duration of time separated of 353 seconds (around 6 minutes) at 3 months of
calf age, to up to a maximum of 2085 seconds (around 35 minutes) at 8 months of age.

Maximum duration of intevals alone

2500
2085
2000
1637
Seconds

1500 1335
1130

1000 812 849 902


706 791
504
500 353 451

0
3 4 5 6 7 8 9 10 11 12

Months

Figure 18. Increasing maximum duration of intervals spent alone by the harbour porpoise calf as a
function of age. The increasing maximum intervals per observation day are plotted at the time they
appear.

35
DISCUSSION

DESCRIPTION OF BEHAVIOURS
Swimming occurred in different relative positions: the mother alone, the calf
alone, the mother and the calf beside each other, and the calf underneath the mother.
The last position, mother-calf underneath, is known as infant position (in dolphins e.g.
McBride & Kritzler 1951; Tavolga & Essapian 1957; Amundin 1986) and has been
hypothesized to be related with protection from predators, nursing facilitation,
phylogenetic conservation or/and having social significance (Gubbins et al. 1999).
In contrast with other species where the calf stays and swims near to the mother
in many varied positions, above her, beside, over the tail, etc. (such as belugas e.g.
Krasnova et al. 2009; or dolphins e.g. Tavolga & Essapian 1957; Gubbins et al. 1999)
the harbour porpoises calf spend the greatest majority of the time swimming in the
infant position, and in fewer occasions beside the mother. The other mother-calf
swimming positions seen in other marine mammals were never observed in the harbour
porpoise mother-calf pair studied here. This suggests that they are probably not typical
for this species. Beside swimming was not a very common behaviour not observed all
the months. It was correlated with the time the mother and the calf spent in the infant
position (either swimming or resting), and the affiliative (social positive interaction)
behaviour called grooming; which also suggests a behaviour showing an association
between mother and calf, but having a less intense nature and performed less often. It
also showed a tendency to decrease along time.
Resting as described in the methods refers to the act of logging on the surface
(Andersen & Dziedzic 1964) in the case of the mother alone, the calf alone or mother-
calf beside each other. The calf was considered to probably rest underneath the mother
while she logged at the surface. It was never observed that the animals were resting in
any other way, as dolphins do, for instance staying immobile under the water (e.g.
Sekiguchi & Hohshima 2003). But the speed and changes in swimming patter were not
studied, as done in Oleksenko & Lyamin (1996) study, where it was shown that a
harbour porpoise mother-calf pair was probably in a non-completely active state when
swimming following the same pattern at a lower speed. We probably could have
underestimated the resting time by considering only the logging time; however, to be

36
able to conclude when animals are certainly resting, electroencephalogram (EEG)
analyses should be made. It has been demonstrated in dolphins that when they sleep a
slower wave activity in the electroencephalogram (EEG) of one hemisphere at a time is
exhibited (Lyamin et al. 2007). And therefore, a similar study in porpoises will allow
firmer conclusions on resting states.
A similar pattern as beside swimming was observed in beside resting. Beside
resting decreased with time, and it was not a very frequent behaviours as it was not even
observed all the months. It was positively correlated with beside swimming, time
together, and grooming. Therefore, showing affiliation, but as beside swimming in a
less intense way than infant position or grooming. Therefore, both beside activities were
seen to be present when the relation was stronger among mother-calf.
Grooming is described in many marine mammals and primates (for instance in
dolphins e.g.; Tamaki et al. 2006) and has been described as an affiliative behaviour. As
described in the methods the behaviour is a succession of rubbing events between two
animals that occur in bouts, one animal strokes the other rubbing its body towards the
other, having the active part, and usually the other responds in the next event rubbing
itself against the other (Amundin & Amundin 1971). Actor and receiver usually switch
several times in the same grooming boat. However, sometimes one individual takes the
active part most or all the times. In this study it was quantified the duration of the
grooming boat. It was the behaviour with the longest mean duration, 52.5 seconds for
each bout. It was not seen every observation day, and sometimes there could be months
without observing any grooming. It was correlated with the time the mother-calf spent
together and with beside resting and swimming.
Evasive and aggressive behaviours were only observed from the mother towards
the calf. One of the evasive behaviours was turning, which occurred when the calf was
in infant position (i.e. underneath her), the mother reacted turning sometimes staying
belly up, or twisting several times. Another one, escaping-following, always occurred in
this way: mother with calf underneath her, the mother reacting actively escaping from
the baby, swimming away from the calf. Immediately, as a response, the calf followed
her trying to swim below the mother again. Sometimes when this happened, the mother
allowed the calf to attain the infant position again, whereas at other times the calf
stopped following her and stayed by herself. In some occasions the turnings were
followed by the mother escaping from the calf. Sometimes only turnings occurred and
some times only escape-follows were seen without a previous turning. As escaping-

37
following most of the time ended in a separation of the mother and a calf, it is of a more
intense nature than turning that usually always ended in the calf staying in infant
position. Aggressive behaviours were observed always in connexion with the calf in the
infant position. All these aggressive and evasive behaviours are probably related with
the beginning of weaning and the parent-offspring conflict seen in many animals
(Triver’s 1974), the parent trying to decrease care, and the offspring demanding extra
care. Avoidances were not correlated (not even negatively correlated) with other
behaviours.
The bumping behaviour was defined as the calf rubbing her head against the
mother’s belly while being underneath her. It was only observed in occasions during
resting and was not a very commonly observed behaviour. Probably it was easier for the
calf to rub the immobile belly during resting than when swimming. If more discrete
rubbing also occurred while swimming this would not be possible to observe from the 5
meter observation height. During the most intense bumpings the calf even turned
horizontally and pushed the mother from below even slightly lifting her tail. The mother
never avoided that performance, on the contrary. The median duration of resting when
the calf was rubbing her head against the mother’s belly had a significantly longer
duration than resting in any of the other resting conditions — calf or mother alone,
beside or underneath mother-calf without bumping — which shows that bumping was a
positive behaviour increasing the duration of resting when occurring. It has been
observed in other marine mammals, and has been related with nursing stimulation (in
dolphins: Cockroft and Ross 1990; belugas: Recchia 1994). In this study we cannot
confirm or reject its relation with nursing stimulation as nursing amounts were constant,
in the presence and absence of bumping. It can however be concluded that a positive
answer was observed from the mother when the calf performed it, which gives it the
positive function of an affiliative grubbing.
Nursing always occurred in infant position. The calf always nursed underneath
the mother while swimming, it was never observed when the mother was resting. Its
duration is very brief, each suckling lasting a mean of 3.7 seconds. A more detailed
nursing description will be presented in another report (Delgado et al. in prep a). The
calf nursed in infant position rolling to get attached to a mammary and staying belly up
while nursing. This has been described as the most common way of nursing in dolphins
(e.g. Gubbins et al. 1999), it has however also been reported in dolphins that the mother
“rolled over one side and glided slowly” during the first two weeks of the calf, probably

38
to facilitate nursing to the calf with still not well developed motor skills (McBride &
Kritzler 1951). It was never observed form the 3rd to the 12th month of the calf’s age in
this study.
Fishing was potential fish ingestion, as it was not possible to distinguish whether
the animals swallowed the fish or not. The general energetic requirements were fulfilled
by the dead fish provided by the trainers during the day. This fishing observations
where not very common and the size of the fishes that were able to pass through the net
was quite reduced.
Object handling was a really playful behaviour, as even though the object (fauna
and flora) could be finally eaten, it was manipulated for some time. As it was almost
absent in the mother, but very common in the calf. The calf inverted the highest
proportion of time chewing on a rope tied to one end to the pool, being this the
preferred object. It was also seen to play with sea stars, catching them and dropping
them and re-catching them again, with algae carrying them around on the mouth,
chasing ducks, etc. it was however, never seen interested in playing with balls left for
the trainers for that purpose. Porpoise’s calves seem to be very playful, as in several
occasions young animals were kept in captivity and performed commonly playful
behaviours such as the ones described above with: algae, pieces of rocks, sea stars,
insects on the surface, swimming games between young animals, etc. balls were not
preferred objects. (Andersen & Dziedzic 1964; Amundin & Amnundin 1971; Wilson
1999). In agreement with Amundin (1974a) playing was never aggressive, in
concordance with the less aggressive behaviour of porpoises in comparison with
dolphins (Andersen & Dziedzic 1964; Amundin & Amnundin 1971; Amundin (1974a)
and Delgado et al. in prep.b)
Bottom grubbing was described by Amundin and Amundin (1974) in wild an
captive conditions, the animal scans the sandy bottom of the pen/sea and between rocks,
algae in what is probably the search of food (crabs, sea stars, mussels, hidden fishes)
using echolocation mostly (Amundin, personal communication). It is a typical
behaviour for the species, as it is performed by the animals also in captive situations
even though the energetic requirements are fulfilled with the dead fish diet provided by
the trainers.
Wall scanning was a behaviour similar to bottom grubbing. The animals were
probably scanning the nets with algae in the search of food. It could also be acoustical
communication with the other animals in the contiguous pen.

39
Fear-stress reaction was in agreement of that described by Andersen & Dziedzic
(1964) and Amundin & Amundin (1971) the animals swimming at very high speed,
slightly leaping out of the water, and sometimes being submerged for a longer time until
the animal jumps, all at high speed, breathing violently. In this study this reactions were
only seen with the pass of motor-boat by the harbour beside the pen. This demonstrates
that sounds made by boats can scare the animals, maybe due to pain or startle caused by
the sound. Not other reasons scared the animals as observations were done when the
centre was closed and no external factors disturbed them.
Coughing. Being the production of a deep hoarse sound, was only observed
when the mother and the calf were showing a fearful-stressful reaction always observed
with the pass of motor-boat; which links both behaviour, being coughing a fearful-
stressful response. It was demonstrated by Amundin (1974b) that underwater
expirations were a way of showing frustration. This could be a manifestation of the
same behaviour, not described in this way previously.
Echelon swimming. It has been observed in other marine mammals that echelon
swimming decreases with time, and it is present in the first stages of development
because of being a less energy consuming way of moving for the calf, taking profit of
the hydrodynamic wave of the mother’s body. Tavolga & Essapian (1957) reported
dolphin calves to swim mostly in echelon position during the first two months, and then
gradually switch to infant position (e.g. Tavolga & Essapian 1957). In this study period
echelon swimming was never seen. It was however observed to be present in an earlier
age, previous to the beginning of the study (personal observation). It can be concluded
that for this harbour porpoise calf the behaviour was not observed form the 3rd month
onwards.
Approaching-turning-spinning defined in the ethogram as a playing behaviour,
was included in the study because it was observed once at the beginning of the study,
however it was so uncommon that was not observed any more. It has been described in
other marine mammals as a common playful behaviour, and also in harbour porpoises
among several young animals (Amundin & Amundin 1971). As seen in the study
between mother and calf seems quite uncommon.

DIFFERENCES ALONG TIME

40
First of all it should be highlighted that differences along time that occur exactly
before and after the 5th month, or/and before and after the 8th month could be caused by
changes in the environment, as the enclosure the animals were kept in changed in its
structure at these two months. Therefore, possible behavioural changes occurring due to
the aging of the calf can be confounded with the environmental changes in these exact
time periods.
The time mother and calf spent together decreased along time in the studied
period. The longest accumulated time together per month was observed in the first
month studied, being the calf 3 months old at that time representing the 69% of the
observed time (41min in 1h). The rest of the period was seen to stay constant until the
10th month. At 10 month of calf’s age, they significantly decreased their time together.
The average time until the 10th month was around 33min in 1h, i.e. 55% of the time
observed, and from the 10th month was around 21min in 1h, i.e. 35% of the observed
time. In Borowska (2009) the same calf was observed in September 2008, two weeks
later to the end of this study, during two weeks, the levels of the mother-calf swimming
together, similar to time-together in this study but not including the time of grooming,
bumping, or suckling. The percentage of swimming together was around 52% of the
observed time (63,13min/2h), being this time greater than the time-together in the
previous three months, this could show that the time-together levels may no continue
decreasing, but vary; also different methodologies could have caused part of the
differences as well. The same calf was the subject of another study at one year and a
half and the time spent together with the mother was around 10% of the time observed
(3500seconds in 10 hours), which shows a greater achievement of independence at that
time (Delgado et al. in prep.b), the mother-calf were however in another social context,
being enclosed together with the other two porpoises in the centre.
When considering the duration of each interval of time alone, it was seen that the
maximum duration of the separation increased with time. Being the maximum time
separated of 353seconds (around 6 minutes) in the first studied month, when the calf
was 3 months old, to up to a maximum of 2085 seconds (around 35 minutes) at 8
months of age, the maximum time separated did however not increase from the 8th
month. This developmental sign is in agreement of what seen in other marine mammals;
intervals spent away from the mother tend to increase along time (in dolphins e.g.
Tavolga & Essapian 1957; Mann & Smuts 1999)

41
This shows important developmental tendencies. The calf achieved at 8 months
a bigger independence, probably due to a change in the calf. And the total time spent
with the mother decreased later on, at 10 months of age, this last decrease was probably
caused by a fall in maternal investment, as suggested from the increase of evasive and
aggressive reactions from the mother (discussed below). Both animals change their
interaction along time.
The evasive and aggressive behaviours (turning, escaping, aggression), that are
probably a way of decrease the mother-calf bond, showed a parent-offspring conflict,
and had a general tendency to increase with time. Considering all these behaviours
together, they increases in a great manner from around the 7th and 8th, and definitively
around the 9th month of calf age. Turnings were observed for the first time around the
5th month of calf age, and seem to be a least intense way of avoidance. When testing
only escaping-following and aggression along time, as seeming of a more intense nature
and appearing latter in time and could reflect a more conscientious way of showing an
active avoidance from the mother, are seen to increase as well around the 7th 8th and 9th
month, however using only this two behaviours the differences were highly significant,
greater differences than when considering turning as well. This shows that these two
behaviours define the period better than turnings. And as suggested above, increasing
avoidance form the mother result in a decrease in the time together at the 10 months of
the calf.
Bumping behaviour was not very common, it was however significantly greater
form the 8th and 9th months onwards than before this time. This could link the behaviour
with nursing stimulation, or with mother affiliation. As this positive behaviour is
observed more in the months with more avoidances (it is however not negatively
correlated with them, probably due the few it was days observed). It could be a rubbing
stimulation performed when the calf is in infant position to favour the mother’s
predisposition to allow the calf to stay underneath her.
Nursing accumulated duration did not vary along the studied period, keeping a
constant amount. Therefore, the data show that the avoidance and the decrease of time-
together did affect the mother-calf bond but not the amount of nursing. The results of
nursing give the first empirical information on this aspect on the species, as the weaning
time was expected to occur around 8 to 10 months, by the analysis of stranded
specimens and the presence of fish in their diet (Lockyer 2003). The presence of fish in

42
the stomach of a calf does not mean complete weaning, nor independence from the
mother. These results suggest that nursing is probably prolonged than expected.
From the results of this mother-calf pair a decrease in the mother-calf bond
occurred from the 8th month onwards, that did not reflect a decrease in nursing, which
indicates that weaning will probably take longer for this observed animal. Until which
extent captivity influences this facts, or how much variation is accounted for individual
differences, cannot be known with only one calf observed, but still the present data sets
a basis for comparisons of other individuals in other conditions, if ever possible also
wild animals.
As explained above, fishing behaviour is referred to potential fishing, as it could
not be distinguished if the fish was eaten or it was just a playful behaviour. The calf was
first seen to chase fishes on the 17-12-2007, when being four and a half months old. The
percentage of time fishing by the calf was seen to significantly increase after the 7th
month of calf age. It did also increase for the mother around that time, but the
percentage was significantly higher in the calf than in the mother. Even if the calf was
not eating the fishes and just playing with them, there is a significant increase in fish
chasing around the 7th month, and this increase is greater than the same observed in the
mother. It can be argued that the increase in the mother around the same time of the calf
could have been due to an increase in the availability of fish. As, fish abundance would
be very limited in winter compared to in the spring, which may explain why fishing
increases at around 7 months after birth, ie in February. Even if this is the case, the calf
inverts a greater percent of time fishing than the mother, which reflects a learning
process. No changes were seen along time in the amount of fishing for the mother with
the calf underneath her, as fishing was almost non-existing in that mother-calf spatial
position. The first confirmation of fish eating by the calf occurred at around one year
and a half, when the animal accepted dead fish from the trainers, it was before tried with
live fishes given by the trainers but the calf was never seen to eat them (Wahlberg
personal communication). The calf was found to be very thin at the time of the first
confirmed fish ate (Wahlberg personal communication). This suggests that weaning is
progressive as seen in other mammals, in this calf was seen to start with a mother
attention decrease around the 10th month of age, when nursing was still occurring at
great mounts and was not seen to decrease. Around that time as expected. One year and
a half was the time the calf started eating fish, and nursing will have decreased, or
stayed constant, not being enough for the growing baby, as deducted from the reduced

43
weight of the calf. This results delay the expected time of fish eating, and even more the
complete weaning time. Undoubtedly, many factors affect the development of calves
and even in almost the same conditions some animals are more precocious than others
(e.g. Reid et a. 1995). Therefore, many observations on animals in controlled conditions
will permit a slow approach to a better knowledge of the species.
Object manipulation was almost only performed by the calf (the calf played with
an object 624 times against the 8 times observed in the mother). Object handling
decreased with time and it was found a breaking point around the 7th month, where
fishing decreased. This tendency is the inverse observed in fishing, which was not really
abundant until the 7th month. This match in timing between decrease in object
manipulation and onset of fishing could explain a developmental process, object
manipulating increasing at 3 months up to 6, and then decreasing; while fishing
increasing at the time of object playing decreases. There was no pattern in object
handling in the mother, nor specific differences along time, which shows (considering
the mother as a control) that the calf spends much more time playing/manipulating an
object than the mother does, and this behaviour changes along time in the calf are
probably due to developmental causes.
Bottom grubbing does not change along time in mother and calf. This shows that
bottom grubbing is a behaviour performed similarly during all the studied months by the
animals alone. No developmental changes are seen in the calf, as she behaves in the
same way as the mother does. In contrast the amount of bottom grubbing performed by
the mother with the calf underneath her does change along time: in the 8th month a
significant decrease is seen. It could be argued that as that decrease occurs in the 8th
month, simultaneously with the change in the environment (removing of the nets), the
mother-calf could have been bottom grubbing underneath the pontoons in the not visible
time and that explain the decrease; however on that assumption the percentage of
bottom grubbing should have decreased similarly for the mother alone and for the calf
alone, and that did not occur. This suggests that the decrease around the 8th month on
the percentage of time bottom grubbing mother with calf underneath is indeed due to a
developmental process. This decrease of mother-calf underneath bottom grubbing could
be explained either by the calf separating from the mother when she starts bottom
grubbing and the calf is in infant position, as the only one performing bottom grubbing
in that position is the mother, the calf just tries to keep with her momentum underneath.
Or, the mother may avoid the calf when she wants to bottom grub, as the calf’s presence

44
may interrupt her in this behaviour. In any case there is a developmental change, due to
an increase in the independence of the calf, or a decrease in the maternal investment
when interfering with other priorities such as food search.
Wall scanning changed along time for both the mother and calf, but no clear
conclusions or any developmental trend can be observed. The data appear as random
changes different from mother and calf (maybe influenced by several unaccounted
factors). The percentage of mother-calf underneath wall scanning did change along time
coinciding with the pattern seen in bottom grubbing, a significant decrease at the 8th
month of calf age, being almost absent afterwards. As not an exact same decrease was
seen for the mother and calf alone performing wall scanning, the developmental
hypothesis suggested for bottom grubbing applies for wall scanning, reinforcing the
theory as two mother-calf underneath behaviours behave in a similar way, decreasing
around the 8th month.
The resting percentage of time did not vary along time neither for mother or calf
alone, nor for mother-calf underneath. This shows that neither developmental nor
environmental changes occurred within the studied months. This indicates that resting
percentage is a behaviour not much altered during this studied developmental period. It
has been seen in dolphins that calves rest during the first weeks in echelon position, and
then switch to sleeping in infant position (McBride & Kritzler 1951).
Swimming percentage of time changed along time for the calf, the mother and
the mother-calf underneath. For calf and mother alone both decreased the percentage of
swimming from the 3rd or 4th month, probably due to environmental changes as the
decrease happened in parallel. A breakpoint was found in the 8th month for the calf
alone increasing the swimming percent after the 8th month. Again, the simultaneous
change in the environment (the net was removed) can probably explain this. (It is
necessary to take into account that it is the proportion of swimming for calf alone from
the total time spent alone what is being considered here, and the increase in swimming
could be explained because the time the animal was visible could have been swimming,
whereas it would perform other behaviours under pontoons). For the percentage of time
mother-calf underneath swam an increase in the percentage of swimming was seen
between the 9th and 10th month. From the absolute values of time together-alone
(accumulated time) it is possible to see that the total time together decreased around the
10th month, therefore this increase in the percent of mother-calf underneath swimming
increases supposes that the mother-calf underneath invest less time doing other

45
behaviours than swimming, and swimming percent increases. This is probably a
developmental change: the mother-calf underneath stay together only for swimming and
other behaviours performed with mother-calf underneath decrease along time, as also
does the total time together.

Even though, no clear significant differences were found in the accumulated


time together between the months there was a decrease after the 4th month and an
unexpected increase the from the 6th to the 9th, this increase in time together could be
due to the fact that from the 6th month the mother was released during the day to the
contiguous pen to interact with the other animals. The calf never acceded to abandon the
pen she was born in, and therefore stayed alone while the mother was in the other pen
(Personal observation). The separation from the mother during the day could probably
have increased the amount of time the calf stayed underneath the mother when
observation were made in the afternoon when the mother was back with the infant, to
compensate the lack of this possibility during the day (This analysis will be include
previous to publication, when checked if there is enough information in the record of
the trainers to know which days the mother was with the others animals; the
consideration of this variable came out when obtaining the results presented here).

DIFFERENCES BETWEEN MOTHER AND CALF ALONE OR TOGETHER

Some differences among mother alone and calf alone behaviours or when the
calf was underneath the mother stood out, and some behaviours and reactions were
similar for both, and in the different spatial conditions. All the studied period was
considered as a whole, not tested for differences along time (done and discussed above).
The duration changes due to the actors or positions were analysed in two ways: The
percentage duration of behaviours per observation (the above data but not considering
differences along time, the percentage of time per observation averaged per all the
studied period), and the mean (also median and maximum) duration of the behaviour
events. Both types of data will be discussed together.

Mother vs. calf


In the first environment there was a floating pool inside the pen, the mother
swam inside it, but the calf never did so. This reflects some developmental differences,

46
maybe also related to individually distinguishable traits. Even though the trainers tried
with positive reinforcement (i.e. toys) to teach her to swim inside the pool, they did not
achieve it at that time, but the calf did it later on (trainers’ personal communication and
personal observation). Showing the calf was reluctant to new situations, not as bolt as
could have been expected for a young. For the mother was not a new situation, she had
been familiarized and trained to go inside the floating pool before. This is in agreement
with the reluctance of the calf to abandon the pen she was born in: When the calf was
one year old the gates communicating the two pens were left open so the calf and the
mother could pass to the other pen or the other animals go in freely to the calf pen. The
calf however refused to pass to the other pen. The calf preferred to stay alone, as the
mother passed to the other pen to interact with the other and swim in the bigger pen
(trainers’ personal communication and personal observation).
It has been seen in other marine mammals that after some weeks after birth,
others not before two months of age, the vigilance from the mother decreases and
allows the calf to interact with other individuals, mostly other females or young animals
(dolphins e.g. McBride & Kritzler 1951; Chirighin, 1987; Mann & Smuts 1998; Mann
& Smuts 1999; Mello et al. 2005; belugas e.g. Krasnova et al. 2009). The studied calf as
kept with the mother alone in one pen only interacted with the other animals in a few
occasions (dates will be checked prior to publication) for a short time (hours). This
could have influenced her development, and therefore generalizations of boldness at
species level need more animals to be studied in different social conditions.
The total percent time the calf inverted on swimming is shorter than the total
percent the mother inverted swimming. The calf inverts less time swimming than the
mother when both are alone, and the duration of each swimming event is shorter in the
calf than in the mother. In contrast to resting, the total percent time invested by the calf
in resting is longer than the time invested for the mother, which means the calf rests in
general more than the mother, but in shorter intervals than her. The total percent of time
invested in bottom grubbing is equal to the percent of time spent on grooming by the
mother, but the duration of the event is shorter for the calf than for the mother.
It can be seen that the behaviour events performed by the calf alone have a
significant shorter duration than the same behaviour performed by the mother alone
(resting, swimming, bottom grubbing). This reflects a more active behaviour of the calf,
changing activities more often.

47
Fishing percentage considering the overall studied period was not significantly
different between calf and mother. This is due to the limited overall amount of fishing,
as not many fishes entered the pen through the net holes. It was however an important
indicator when the study period was divided in two (with a significant difference
percent of fishing before and after the 7th month for both mother and calf, fishing
increasing after the 7th month for both and being almost negligible before the 7th
month). In the period from the 7th month onwards there were significant differences in
the percent of fishing for mother and calf. The median duration of fishing events had a
longer duration in the calf than in the mother, contrasting with the usual shorter
durations of events for the calf than for the mother. This, probably reflects the learning
process and the inexperience of the calf in catching fishes that makes the duration of
fishing events longer than the one from the mother, as well as that probably the calf is
as well playing with them.
The percentage of time invested in object handling by the calf is much longer
than the percentage of time invested by the mother. The calf was seen to play with an
object 624 times along the studied period, whereas there were only 8 events observed
with the mother. The duration of object handling events was not significant different
between the mother and calf.

Mother-calf underneath vs. alone or beside spatial positions


The percentages of bottom grubbing and wall scanning behaviours have a
shorter duration when the calf is underneath the mother than when mother or calf
performs them alone. In addition, the duration of bottom grubbing and wall scanning
events have a shorter duration when the calf is underneath the mother while she
performs them, than when mother and calf do them alone as well. This reflects either
that the mother-calf as a unit is not the preferred moment to perform these behaviours
by the mother, or that the calf and mother separate when that activity is started by the
mother, it was commonly observed that the calf tended to leave infant position to do
something else while the mother was busy scanning for food. As discussed below it was
also seen that along time the percentage of both behaviours of the mother with the calf
underneath decreased, supporting that is not the preferred time by the mother to do
them, and that tendency becomes more obvious with time because of the calf being
more independent or the mother less tolerant with the calf being close. This also shows
that it was probably not done by the mother with the intention of teaching. Although

48
this contrasts with the mean duration of fishing events when the calf is underneath the
mother, which does not differ from the duration of the mother fishing alone. However,
there is a significantly smaller percent of time the mother inverts fishing with the calf
underneath than the mother alone, which shows is more probable the not-teaching
hypothesis. As if the mother was trying to teach the calf when it is underneath, a greater
event duration and the percentage of time inverted on fishing would be excepted when
the calf is in infant position than when the mother is lone, to suggest that the mother is
actively teaching the calf. And together with the much longer time of the calf fishing
alone, and its greater fishing frequency compared to the mother, trial and error, and
perhaps observation of the mother, seem to be the present.
The percentage of swimming had a longer duration when the calf is underneath
the mother than when the calf is alone, but a shorter duration than when the mother is
alone. The duration of swimming events when the calf is underneath the mother had a
longer duration than when the calf is alone, but shorter than when the mother is alone.
When mother and calf swam beside each other the median duration of the
behaviour had a similar duration than when the calf swam alone, but differ form the
other longer durations, mother alone, and mother-calf in infant position.
The percentage of resting when the calf is underneath the mother is significantly
greater than the percentage of the calf and the mother resting alone. Accordingly its
really interesting to notice that resting events are significantly longer when the calf is
underneath the mother than when both of them are alone or beside each other. This
longer duration is probably an important adaptative behaviour, as when the calf is under
the mother protection, both mother and calf rest for a longer time. This seems
meaningful, as the mother does not need to be as vigilant as when the calf stays alone,
and the same for the calf.

Perspectives
From this study it is clear that the more variables and factors can be controlled
for a behavioural study, the clearer results will be and the fewer alternative hypotheses
will remain. As it occurs in any scientific study. Therefore the changes in the
environment reported during this study have diminished the possibilities to draw
conclusions from the results presented here by introducing many new variables to take
into account. Thus, diverse possible explanations were possible for the observations
made here (they could either be caused by developmental or environmental changes, or

49
a combination of those). On the other hand when keeping alive beings in captivity,
animal welfare should have the highest priority. Changes in the environment are known
to be positive and enriching for the animals and decreasing the prevalence of
stereotyped behaviours (Kastelein & Staal 1997) (always maintaining the required
spatial dimensions, big enough, for the species).
In addition, social influence highly probably shapes development in the species,
as occurs with social animals, among which are harbour porpoises (Andersen &
Dziedzic 1964; Amundin 1974a; Delgado et al. in prep). In this study, the mother-calf
pair being kept alone for a year will conceivably have determined the calf development,
it is however not known how and when cow-calf pairs interact with other individuals of
the species in the wild, even though, this timing could be pointed to be from few months
after birth as seen in other marine mammals: after some weeks after birth, others not
before two months of age, the vigilance from the mother decreases and allows the calf
to interact with other individuals, mostly other females or young animals (dolphins e.g.
McBride & Kritzler 1951; Chirighin, 1987; Mann & Smuts 1998; Mann & Smuts 1999;
Mello et al. 2005; belugas e.g. Krasnova et al. 2009). Therefore, the way this mother-
calf isolation could have shaped the calf development can not be drown from this study.
And should be accounted for comparisons in further studies.
Covering all the developmental process from birth to adultness will as well
provide a more complete understanding of the process.
Future behavioural studies in this species will help to resolve the uncertainties
created by the changes in the environment; as well as the social and environmental
conditions the animals were conditioned by in the study.

50
REFERENCE LIST

Altman, J. (1974). Observational study of behaviour: Sampling methods. Behaviour 49,


pp. 227-265.

Amundin, M. (1986). Breeding the bottle-nosed dolphin (Tursiops truncatus) at the


Kolmarden dolphinarium. Int. Zoo. Yb. 24/25, pp. 263-271.

Amundin, M. (1974a). Occupational therapy for Harbour porpoises, Phocoena


phocoena. Aquatic Mammals, 2 (3), pp. 6-11.

Amundin, M. (1974b). Some evidence for a displacement behaviour in the harbour


porpoise, Phocoena phocoena (L): A causal analysis of a sudden underwater expiration
Through the blow hole. Rev. Comp. Animal, 8, pp. 39-45.

Amundin, B. & Amundin, M. (1971). Nagra etologiska iakttagelser över tumlaren,


Phocoena phocoena (L.), i fangenskap. Zool. Revy., 33, pp.51-59.

Amundin M. & Amundin B. (1974). On the behaviour and study of the harbour
porpoise, Phocoena phocoena, in the wild. Investigation on Cetacea, 5, pp. 317-328.

Andersen S. & Dziedzic, A. (1964). Behaviour patterns of captive harbour porpoise


Phocaena phocaena (L.) Bull. Inst. océanogr. Monaco, 63 (1316).

Blanchet, M-A., Wahlberg, M., Kristensen, J., Hansen, S., Jensen A-L & Elk V.N.
(2008). First birth of a harbour porpoise (Phocoena phocoena) under human care.
Soundings, 33 (2) pp. 6-9.

Borowska, E. 2009. The relationships between mother and calf in harbour porpoise
(Phocoena phocoena), at the basis observations in Fjord&Baelt Centre in Denmark. ).
M.Sc thesis, Warsaw University of Life Sciences. 58pp.

51
Chirighin, L. (1987). Mother-calf relationships and calf development in the captive
bottlenose dolphin (Tursiops truncatus). Aquatic Mammals, 13 (1), pp. 5-15.

Cockroft, V.G. and Ross, G.J.B. (1990). Observations on the early development of a
captive bottlenose dolphin calf. In: Leatherwood S, Reeves R, editors. The bottlenose
dolphin. San Diego: Academic Press. Pp 461-78.

Delgado, L., Wahlberg, M. & Norum, U. In prep.a. Social interactions in captive


harbour porpoises (Phocoena phocoena). M.Sc thesis, University of Southern Denmark.
95pp.

Delgado, L., Wahlberg, M., Blanchet, M-A., et al. In prep.b. Nursing behaviour in a
harbour porpoise (Phocoena phocoena).

Dytham, C. 2003. Choosing and using statistics: A biologist guide. Blackwell Science.
Blackwell Publishing, Oxford.

Fowler, J., Cohen, L & Jarvis, P. 1998. Practical statistics for field biology. Ed.
Whiley.

Gubbins C., McCowan B., Lynn, S.K., Hooper, S. & Reiss, D. 1999. Mother-infant
spatial relationships in captive bottlenose dolphins, Tursiops truncatus. Marine
mammals science, 15(3), pp. 751-765.

Kastelein, R.A. & Staal, C. 1997. Swimming behaviour of a harbour porpoise


(Phocoena phocoena) under different conditions in human care. In The Biology of the
Harbour Porpoise. (Eds. A.J. Read, P.R. Wiepkema & P.E. Nachtigall), pp.235-253. De
Spil Publishers, Woerden, The Netherlands.

Lyamin, O., Pryaslova, J. Kosenko, P. & Siegel, J. 2007. Behavioural aspects of sleep
in bottlenose dolphin mothers and their calves. Physiology & Behaviour, in press.

52
Lockyer, C. 2003. Harbour porpoises (Phocoena phocoena) in the North Atlantic:
Biological parameters. In Harbour porpoises in the North Atlantic. NAMMCO Sci.
Publ. 5. (Eds T. Haugh, G. Desportes, G.A. Víkingsson & L. Witting), pp. 71-90.

Mann, J. & Smuts, B. 1998. Natal attraction: allomaternal care and mother-infant
separation in the wild bottlenose dolphins. Animal Behaviour, 55, pp. 1097-1113.

Mann, J. & Smuts, B. 1999. Behavioural development in wild bottlenose dolphin


newborns (Tursiops sp.) Behaviour, 136, pp. 529-566.

McBride, A.F. & Kritzler, H. 1951. Observations on pregnancy, parturition, and pos-
natal behavior in the bottlenose dolphin. Journal of Mammalogy, 32 (3). 251-266.

Mello, I., Nordensten, L. & Amundin, M. 2005. Reactions of three bottlenose dolphin
dams with calves to other members of the group in connection with nursing. Zoo
Biology., 24, pp. 543-555.

Oleksenko, A.I. & Lyamin, O.I. 1996. Rest and activity states in female and baby of
harbour porpoise (Phocoena phocoena). J. Sleep Res, 5, supplement 1.

Perelberg A. & Schuster R. 2008. Coordinated breathing in bottlenose dolphin


(Tursiops truncatus) as cooperation: Integrating proximate and ultimate explanations.
Journal of comparative Pysichology, 122(2),pp. 109-120.

Recchia, C. A. 1994. Social behaviour of captive Belugas, Delphinapterus leucas.


Ph.D. dissertation, MIT/WHOI. WHOI-94-03.

Reid, K., Mann, J., Weiner, J.R. & Hecker, N. 1975. Infant development in two
aquarium bottlenose dolphins. Zoo biology, 14, pp. 135-147.

Sekiguchi, Y., & Hohshima, S. 2003. Resting behaviours of captive bottlenose


dolphins (Tursiops truncatus). Physiology & Behaviour, 79, pp. 643-653.

53
Tamaki, N., Morisaka, T. & Taki, M. 2006. Does body contact contribute towards
reparing relationships? The association between flipper-rubbing and aggressive
behaviour in captive bottlenose dolphins. Behavioural processes, 73, pp. 209-215.

Tavolga, M.C. & Essapian, F.S. 1957. The behaviour of the bottlenose dolphin
(Tursiops truncatus): mating, pregnancy, parturition and mother-infant behaviour.

Teilmann, J., Larsen, F. & Desportes, G. 2007. Time allocation and diving behaviour
of harbour porpoises (Phocoena phocoena) in Danish and adjacent waters. J. Cetacean
Res. Manage., 9 (3), pp. 201-210.

Trivers, R. L. 1974. Parent-offspring conflict. American Zoologist, 14, pp. 249-264.

Wahlberg, M. et al. In prep. Age estimates in captive harbour porpoises based on their
growth curves.
Wilson, S.D. 1999. Preliminary Investigations into the social behaviour and
development of a juvenile harbour porpoise Phocoena phocoena (Linnaeus, 1758) in
captivity. M.Sc. Thesis, School of Ocean Sciences, University of Wales, Bangor, UK.
134pp.

Zar, J.H. 1999. Biostatistical analysis. Prentice Hall, In., New Yersey.

54