maturation and development of social behaviour during early ontogeny in free-ranging dog puppies in...

Maturation and development of social behaviour

during early ontogeny in free-ranging dog

puppies in West Bengal, India

Sunil Kumar Pal *

Katwa Bharati Bhaban, Katwa, Burdwan 713130, West Bengal, India

Accepted 21 May 2007

Available online 22 June 2007

Abstract

Six bitches and their offspring were observed in the natural environment to study the early ontogeny of

certain processes like eye opening, nipple preference, play, sexual activities and locomotion. Thirteen puppies

out of 35 survived at the age of 3 months indicating 63% mortality. Eye slits were observed on day 12 (median)

of life, and the eyes completely opened on 17 day (median) of life. The rearmost (fourth) nipple pair

presumably containing more milk was suckled preferably by the puppies showing no sex bias. Social

communication among the littermates was developed through playful interactions. Play behaviour developed

in three subsequent stages—social play, agonistic play, and pseudo-sexual activity; at 3, 5 and 6 weeks,

respectively. Playful interactions increased with the age of the puppies to establish a stable social hierarchy, and

suddenly decreased between 8 and 10 weeks of life. Aggressive play tended to be initiated by males and specific

individuals within a litter, suggesting individual variation in aggressiveness. Development of pseudo-sexual

behaviour especially in male puppies by week 6 was an interesting feature of this study. The puppies were

mobile at the age of week 3 when the eyes completely opened, and they were highly mobile when the suckling

bouts disappeared and they traveled independently for food collection (10–11 weeks of life).

# 2007 Elsevier B.V. All rights reserved.

Keywords: Canis familiaris; Vision; Nipple preference; Play; Sexual activity; Locomotion

1. Introduction

From birth to about 2 weeks of age, puppies are relatively helpless with nearly all effort

devoted to eating and sleeping. Their eyes and ears are not yet open and body temperature is

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Applied Animal Behaviour Science 111 (2008) 95–107

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E-mail address: [email protected].

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maintained by their mother, siblings and environment. This is often called the neonatal period.

The period from 14 to 21 days seems to be a transition period when pups begin to develop their

motor skills as well as develop their vision and hearing. Social behaviour in dog puppies starts to

develop from week 3 of life (the transitional period, Markwell and Thorne, 1986), when the

puppies begin to contact littermates, and continues during the socialization period (4–10) when

they start to communicate with each other using many gestures and vocalizations (Scott and

Fuller, 1965; Campbell, 1975; Fox and Bekoff, 1975; Fox, 1978). A number of studies have

previously been done on the development of social behaviour, especially on dominance and

dominance-related behaviour (Scott and Marston, 1950; James, 1952; Scott and Fuller, 1965;

Fox, 1971; Ewer, 1973; Bekoff, 1974a,b, 1977, 1995, 2001; Campbell, 1975; Fox and Bekoff,

1975; Bradshaw and Nott, 1995; Macdonald and Carr, 1995; Serpell and Jagoe, 1995). Very few

studies have been conducted on the physical development of puppies (Rheingold, 1963; Lund

and Vestergaard, 1998). However, most previous studies have been conducted in American and

European countries, and on enclosed captive populations. Since the environment affects the rate

of development (DeHasse, 1994), there is a need for studies in a naturalistic setting for

comparative purposes.

2. Material and methods

2.1. Study area

The study was conducted in the town of Katwa (238140N, 87850E) in the state of West Bengal, India. The

study area (0.5 km2) with a population of approximately 4200 people, was on the outskirts of the town and

was linked with other parts of the town by a metalled road. The town’s main bus stand was situated in the

study area. The area comprised 20% low-income, 65% middle-class, and 15% higher-class neighbourhoods

(Pal, 2003, 2005). The study area was characterized by an abundance of open land with many private houses,

cowsheds and poultry farms. Interviews with the area residents confirmed that about 20% of the low-income

residents allowed female dogs to use their cowsheds or unused poultry farms for pup rearing. Garbage was

the main food resource for the dogs and was scattered throughout the study area. However, the restaurants

and butcheries adjacent to the bus stand were food resources for the studied dogs.

2.2. Methods

The study was carried out on free-ranging domestic dogs (Canis familiaris). Six adult bitches, whose home

range and prior reproductive history were known, were selected for this study. The animals were identified by

photographic mark-recapture (Beck, 1973) and also visually by coat colour and pattern. The sex of the dogs

was determined by observing the animals and their behaviour, especially their urination postures (Bekoff,

1979). Visual assessment was used to distinguish puppies (birth–4 months) and adults (1–7 years).

The study began in April 2005 and continued until the end of March 2006. Data were collected by using

ad libitum and focal-animal sampling (Altmann, 1974). The focal bitches were occasionally fed bread to

make them tame during the rearing period for collecting data on pup activities. The focal bitches were

observed once a week before gestation; and during gestation, the bitches were followed daily between 06:00

and 18:00 h. Direct observations and interviews with the area residents confirmed the date and time of

parturition.

After parturition, the litters were observed thrice a day (30 min per session) between 06:00 and 18:00 h.

Data were collected on a weekly basis; and data collection continued for the first 13 weeks of rearing or until

weaning.

Litter sizes were based on the first sighting of the puppies. Pup survival was estimated from the

difference between the numbers at the first sighting and the numbers observed over the following weeks.

S.K. Pal / Applied Animal Behaviour Science 111 (2008) 95–10796

In agreement with the results of previous studies (Pal, 2005), bitches in this study had four pairs of

nipples. During lactation, the size of the nipple pairs was estimated visually. The relative size of the different

nipple pairs was taken as an estimate of milk content.

Playful interactions were divided into three categories: (a) social play, (b) agonistic play (play-fighting),

and (c) pseudo-sexual play. Social play included mutual mouthing, biting, tumbling over each other’s body,

pouncing on and lunging into other, running and chasing among the puppies (Rheingold, 1963; Bekoff and

Byers, 1981, 1998; Fagen, 1981). Agonistic play included stalking and chasing, ambushing and attacking,

standing over a littermate; circling the littermate with a stiff wagging tail, raised hackles, baring teeth,

snarling or biting, direct stare with dilated pupil, standing with forepaws on littermate’s back, wagging tip of

erect tail, erect or completely flattened ears, and taking play fighting to extremes (Rheingold, 1963; Lund

and Vestergaard, 1998; Pal et al., 1998a). Pseudo-sexual play in puppies included mounting, clasping and

pelvic thrusting during playful interactions among the littermates.

The puppies were considered as mobile when the eyes were completely opened, and they were considered

as ‘highly mobile’ when they spontaneously and independently traveled the long distance from the nest site

during a 30 min-observation period after the ending of suckling bouts. The origin of locomotion (nest site) and

subsequent locations of each puppy were plotted on a scale map of the area. The distance between the nest site

and the final location was measured by a tape and by a map measurer along the routes by which the locations

were linked, and the estimated distance was considered as the distance of locomotion.

All observations were made on foot or by a bicycle with use of binoculars. A Yashika Fx-3 super (2000)

camera with 35–70 mm lens was used for documentation of different activities of dogs.

For statistical analysis a Pearsonian chi-square model was followed. A paired t-test was used to examine

the differences between male and female puppies in relation to playful interactions. Pearson’s coefficient of

correlation was used in this study to calculate the correlation between the age of puppies and the frequency

of suckling bouts, and also to calculate the correlation between the age of puppies and the frequency of

playful interactions. Analysis of variance (ANOVA) was used to test the differences of the puppies in

relation to the weekly mean distance covered by them. Probability level for rejection of the null hypothesis

was set at P > 0.05 for all tests.

3. Results

3.1. Litter size, sex ratio and mortality

Focal bitches (n = 6) produced six litters between November and December comprising a total

of 35 puppies (Table 1) with a male biased sex ratio of 1.69:1. Mean (�S.D.) litter size was 5.83

(�1.57) (range 4–9).

Of all puppies (n = 35), only 13 puppies survived at the age of 13 weeks (3 months) indicating

63% mortality.

S.K. Pal / Applied Animal Behaviour Science 111 (2008) 95–107 97

Table 1

Birth history and pup rearing for six bitches in West Bengal (India) from march 2005 to march 2006

Female

ID

Date of

parturition

Duration of

lactation (week)

Litter size Pups survived at 13 weeks of age

M F M F

PPF 7th October, 2005 11 5 4 1 1

SMF 11th October, 2005 11 3 2 1 1

TSF 1st December, 2005 11 2 3 2 1

RSF 3rd December, 2005 10 5 1 2 1

NPF 2nd December, 2005 10 4 2 1 1

KBF 18th December, 2005 11 3 1 1 –

3.2. Development of vision

Immediately after birth, the puppies were completely blind and the eyes opened gradually.

Eye slits in some puppies were first observed on day 10 (range = 10–13 days, median = 12 days).

The eyes completely opened between 15 and 18 days (median = 17 days).

3.3. Nipple preference and suckling bouts

Each of the six lactating bitches were observed to have four pairs of nipples. Among the four

nipple pairs, the fourth pair (rearmost) was the largest and the second pair was the smallest in all

bitches. In 67% cases, the third nipple pair was larger than the first nipple pair; and in 33% cases

the first nipple pair was larger than the third nipple pair.

A total of 2325 suckling bouts were recorded in this study, and the mean (�S.E.) number of

suckling bouts for 1st, 2nd, 3rd and 4th nipple pairs were 21.54 (�2.20), 8.85 (�1.30), 32.15

(�2.17) and 116.31 (�4.64) times, respectively. So, the 4th nipple pair (rearmost) was suckled in

a greater frequency than the other pairs (x2 = 158.96, d.f. = 3, P < 0.05). The rate of suckling for

the 4th nipple pair bouts was 125.38 times in the case of male puppies and it was 101.80 times in

the case of female puppies. So, there were no significant differences between the male and female

puppies in relation to the rate of suckling of the fourth nipple pair (x2 = 2.45, d.f. = 1, P > 0.05,

Fig. 1).

The puppies (n = 13) were observed to suckle up to 10–11 weeks of age. The mean

frequency of suckling bouts (�S.E.) for an individual puppy varied from 22.77 (�0.48)

(1st week of life) to 5.50 (�0.25) times (10–11 week of life). During the first 5 weeks, the

puppies were observed to suckle almost in every 30-min observation period (Fig. 2). From the

5th week, the frequency of suckling bouts gradually decreased (r = �0.9622, P < 0.05).

Average number of suckling bouts in case of male puppies varied from 0.51 to 2.17 times/h

and in case of female puppies it varied from 0.55 to 2.17 h�1 (Fig. 3), and there was no

difference between the male and female puppies in relation to their rate of suckling

(t = �1.00, d.f. = 10, P > 0.05).


Fig. 1. Average no. of suckling bouts for each nipple pair by 13 puppies recorded on the basis of thrice per day with 30-

min duration in each observation period from birth to 10–11 weeks of age.

3.4. Play behaviour

Social play as first playful interaction among the littermates was first observed by week 3.

Throughout the rearing period, a total of 6014 intra-litter playful interactions were recorded of

which 3549 were classified as social play. The weekly mean (�S.E.) number of social play varied

from 12.83 (�1.98) (3rd week of life) to 81.33 (�5.90) (7th week of life) to 1.44 (�0.18) times

(13th week of life (Table 2). The weekly mean number of social plays initially increased with age

(r = 0.9074, P < 0.05), but it suddenly decreased in week 8 and continued to decrease thereafter

(r = �0.9309, P < 0.05). Moreover, there was significant variation among individual puppies in

relation to the total number of social play recorded from birth to 13 weeks of age (x2 = 200.87,

d.f. = 11, P < 0.05, Table 3). The frequency of social play in the case of male puppies varied from

0.16 to 9.31 times/h/week, whereas in the case of female puppies it varied from 0.10 to 5.68


Fig. 2. Weekly mean (�S.E.) number of suckling bouts by 13 puppies recorded on the basis of thrice per day with 30-min

duration in each observation period from birth to 10–11 weeks of age.

Fig. 3. Number of suckling bouts per hour per puppy recorded on the basis of thrice per day with 30-min duration in each

observation period from birth to 10–11 weeks of age.

S.K

.P

al/A

pp

liedA

nim

al

Beh

avio

ur

Scien

ce1

11

(20

08

)9

5–

10

71

00

Table 2

Weekly mean (�S.E.) number of playful interactions recorded on the basis of thrice per day with 30-min duration in each observation period among 13 puppies

Playful

interactions

Age of the puppies (week)

3rd 4th 5th 6th 7th 8th 9th 10th 11th 12th 13th

Social play 12.83 � 1.98 23.42 � 1.40 26.67 � 1.60 39.75 � 2.91 81.83 � 5.90 52.92 � 3.93 30.25 � 2.45 15.42 � 1.81 7.42 � 0.89 3.17 � 0.47 1.44 � 0.18

Agonistic play – – 7.92 � 0.92 16.42 � 3.63 30.83 � 6.21 48.33 � 9.78 24.42 � 5.83 13.25 � 2.90 8.33 � 2.00 3.58 � 0.97 1.80 � 0.42

Pseudo-sexual play – – – 5.50 � 0.58 9.00 � 1.04 11.08 � 1.29 13.42 � 1.38 7.33 � 0.91 2.75 � 0.47 1.33 � 0.25 0.42 � 0.23

times/h/week (Fig. 4). The difference between the male and female puppies in relation to the

frequency of social play per hour was statistically significant (t = 3.83, d.f. = 10, P < 0.05).

Agonistic play was first observed by week 5, and a total of 1855 episodes were recorded

(Table 2). The weekly mean (�S.E.) number of agonistic play varied from 7.92 (�0.92) (5th

week of life) to 1.80 (�0.42) times (13th week of life). The weekly mean number of agonistic

plays increased with age (r = 0.9890, P < 0.05), but it suddenly decreased in week 9 and

continued to decrease thereafter (r = �0.9598, P < 0.05, Table 2). Moreover, one male in each

litter initiated play with agonistic interactions with a greater frequency than the others indicating

the highest ranking of aggression (Table 3). So, there were significant individual differences

among the puppies in relation to the total number of agonistic play recorded from birth to 13

weeks of age (x2 = 879.59, d.f. = 11, P < 0.05). The rate of agonistic play in the case of male

puppies varied from 0.19 to 5.93 times/h/week, whereas in the case of female puppies it varied


Table 3

Total number of playful interactions performed by each of 12 puppies recorded on the basis of thrice per day with 30-min

duration in each observation period during birth to 13 weeks of age

Puppy ID Playful interactions

Social play Agonistic play Sexual play

PPM1 (PP) 331 167 57

PPF2 (PP) 189 114 31

SMM3 (SM) 367 166 65

SMF1 (SM) 213 91 31

TSM1 (TS) 393 400 63

TSM2 (TS) 356 107 71

TSF2 (TS) 220 89 24

RSM1 (RS) 365 365 59

RSM3 (RS) 320 101 74

RSF1 (RS) 245 76 30

NPM2 (NP) 335 116 75

NPM2 (NP) 215 63 30

Total 3549 1855 610

Litter ID in parenthesis.

Fig. 4. Number of social play bouts per hour per puppy (n = 12) recorded on the basis of thrice per day with 30-min

duration in each observation period from birth to 13 weeks of age.

from 0.13 to 2.74 times/h/week (Fig. 5), and the difference was statistically significant (t = 3.54,

d.f. = 8, P < 0.05).

Mouthing and clasping among the littermates were common during playful interactions.

However, at the age of 6 weeks, puppies of both sexes were observed to mount, clasp and give

pelvic thrusts. Only one pup in the KB-litter survived at the age of 13 weeks, and so, no data on

sexual behaviour for this litter were obtained. A total of 610 sexual interactions were recorded in

this study. The weekly mean (�S.E.) number of sexual activities varied from 5.50 (�0.58) (6th

week of life) to 13.42 (�1.38) (9th week of life) to 0.42 (�0.23) (13th week of life). The weekly

mean number of sexual activities increased with age to week 9 (r = 0.9929, P < 0.05), but it

decreased from the age of week 10 (r = �0.9319, P < 0.05, Table 2). Moreover, there was

significant variation among individual puppies in relation to their sexual activities initiated from

birth to 13 weeks of age (x2 = 200.87, d.f. = 11, P < 0.05, Table 3). The rate of sexual activity in

the case of male puppies varied from 0.10 to 1.63 times/h/week, and in case of female puppies it

varied from 0.02 to 0.79 times/h/week (Fig. 6). So, male and female puppies were significantly


Fig. 5. Number of agonistic play bouts per hour per puppy (n = 12) recorded on the basis of thrice per day with 30-min


Fig. 6. Number of pseudo-sexual activities performed by 12 puppies recorded on the basis of thrice per day with 30-min


different from each other in relation to the frequency of sexual activity per hour (t = 4.34, d.f. = 7,

P < 0.05).

3.5. Locomotion

Throughout the rearing period, a total of 951 observations (30-min duration for each

observation) were made to estimate the distance covered by the puppies per week. From the first

day of litters’ life, the puppies were observed to move several centimetres in search of the nipples

of their mothers. In week 1, the puppies were observed to reach their mothers by crawling. In

week 2, the puppies were observed to cover a distance of about 1 m with belly dragging from the

nest site to their mothers. In week 3, the puppies were observed to leave the nest site. The weekly

mean (�S.E.) distance covered by the puppies was a minimum of 2.62 (�0.03) m during the 3rd

week, and a maximum of 61.20 (�0.34) m during the 11 weeks of litters’ life (Fig. 7). So, the


Fig. 7. Weekly mean (�S.E.) distance moved by 13 puppies recorded on the basis of thrice per day with 30-min duration

in each observation period from birth to 11 weeks of age.

Fig. 8. Weekly mean distance moved by eight male and five female puppies recorded on the basis of thrice per day with

30-min duration in each observation period from birth to 11 weeks of age.

distance covered by the puppies gradually increased with the age of the puppies, and there was a

positive correlation between age and the distance moved by the puppies (r = 0.9730, P < 0.05).

During the last week of suckling bouts (10–11 weeks), the average distance covered by the

puppies varied from 57.34 to 63.42 m; however, there were no significant variations among the

puppies in relation to the weekly mean distance covered by the puppies (F = 0.05; d.f. = 12, 113;

P > 0.05). The weekly mean distance covered by the male puppies varied from 2.62 m (during

the 3rd week) to 60.46 m (during the 10th or 11th week) and in case of female puppies it varied

from 2.61 m (during the 3rd week) to 60.22 m (during the 10th or 11th week, Fig. 8). There were

no significant differences between the male and female puppies in relation to the weekly distance

moved by the puppies (t = 1.73, d.f. = 8, P > 0.05).

4. Discussion

Kleiman (1968) reported that domestic dogs breed twice a year, and several subsequent

authors (Daniels and Bekoff, 1989; Boitani et al., 1995) have confirmed this. However,

observations conducted during this 1-year study indicate that the free-ranging dogs in West

Bengal breed only once a year (see also Ghosh et al., 1984; Pal et al., 1998b; Pal, 2001). The

mean litter size recorded in this study was similar to that reported by Macdonald and Carr (1995)

and Pal (2001). Moreover, litters had a male-biased sex ratio, as also reported by Beck (1973),

Daniels (1983), Daniels and Bekoff (1989), Macdonald and Carr (1995), and Pal (2001). The

mortality rate of 63% before the age of 4 month is similar to that reported in previous studies in

domestic dogs (e.g., Scott and Causey, 1973; Daniels and Bekoff, 1989; Boitani et al., 1995; Pal

et al., 1998b; Pal, 2001).

Domestic dogs (Canis familiaris) are born in a semi-altricial state like most carnivores (Ewer,

1973; Poole, 1985), and they are basically deaf, blind and immobile. In this study, eyes

completely opened between 15 and 18 days of life, a little earlier than reported by Rheingold

(1963) and Serpell and Jagoe (1995). From this study, it may be suggested that visual

communication in domestic dogs naturally develops by week 3 when the puppies are able to

respond to objects other than to mother and littermates.

The number of nipple pairs varies among species and reflects the litter size (Ewer, 1973). The

domestic dogs in this study had a rather low number of nipples as compared to other carnivores

and only the fourth pair (rearmost) was clearly preferred by puppies. In contrast, no nipple

preference in domestic dogs was found by Rheingold (1963), despite the suggestion that the

rearmost pair might contain more milk. During the first 5 weeks of a litter’s life, puppies were

observed to suckle almost in every 30-min observation period, which was also previously

reported by Pal (2005). The frequency of suckling decreased as puppies grow older (Pal, 2005).

Although evidence is scant, there was no difference between male and female puppies in relation

to the rate of suckling.

Although there are many important life stages, the key sensitive period in a puppy’s

development is generally accepted to be from 3 weeks to 3 months of age. This 9-week period is

the primary sensitive period of canine development. This period is referred to as the ‘‘critical

bonding period,’’ or ‘‘sensitive period’’ (Serpell and Jagoe, 1995). This is the period of rapid

development of social behaviour patterns. During this stage, the dog learns dominant behaviours,

submissive behaviours, and agonistic behaviours. Species identification is acquired during this

sensitive phase of development. In a previous study conducted in the town of Katwa (Pal, 2003),

the free-ranging dogs were described as ‘social’ animals (e.g., Fox et al., 1975; Carr, 1985; Font,

1987). Social play is the major form of social interaction among puppies. According to Spinka


et al. (2001), play is very important for social, cognitive, and, or, physical development, and may

also be important for training youngsters for unexpected circumstances. Therefore, it is important

for individuals to engage in at least some play (Bekoff and Byers, 1981). In this study, the puppies

were observed to start play with their littermates at the end of the 3rd week of age, which had

previously been observed by Lund and Vestergaard (1998). Agonistic play as second stage of

play behaviour developed by week 5; and sexual behaviour as the third stage of play behaviour

developed by week 6. From the developmental data collected in Katwa town, I suggest that play

behaviour in puppies develops through three consecutive stages: (i) social play, (ii) agonistic

play, and (iii) sexual play.

Aldis (1975) reported that play was not less frequent in female than in male domestic dogs. On

the contrary, in this study all the three types of playful interactions were more frequent in male

than in female puppies. This sex difference agrees with the findings of Baldwin and Baldwin

(1977) and Symons (1978) in many group living primates, Holmes (1995) in ground squirrels,

and Lund and Vestergaard (1998) in domestic dogs.

Blackshaw (1985) suggested that the formation of a dominance hierarchy is a necessary part

of achieving a stable social environment. Since dogs are social animals (Pal, 2003), they must

develop a dominance hierarchy. Play fighting allows puppies to establish social ranks (Fox and

Bekoff, 1975), which are necessary for dominance hierarchies. In this study, one male from each

litter initiated most of the intra-litter agonistic interactions. This may contribute to stability in the

group and also reduce intra-group aggression (Scott and Fuller, 1965; Bradshaw and Nott, 1995).

The developmental data in Katwa town suggest that, if there is a sensitive period for the

acquisition of dominance-related assertiveness or the consequences of aggression in domestic

dogs, it is likely to peak during the first 10 weeks of the litters’ life. This is also in agreement with

the views of Scott (1962), Fox et al. (1976), Bradshaw and Nott (1995), and Serpell and Jagoe

(1995).

An increase and sudden drop of different types of playful interactions especially the agonistic

plays with the age of the puppies were also previously noted by Lund and Vestergaard (1998).

Scott and Fuller (1965), and Bradshaw and Nott (1995) also suggested that if the pups remain

together, each pairwise relationship becomes stable by about the 11th week. In my study the

puppies were observed to live together throughout the study period (13 weeks of life). Group

stability among the dogs was also previously reported in Katwa town by Pal et al. (1998a). The

frequencies of agonistic play among the puppies increased with the age of the puppies and then

suddenly decreased at the age of 8 weeks of life. So, it may be presumed that in nature a stable

hierarchy within the litter is established at the age of 7 weeks of puppy’s life.

Development of pseudo-sexual behaviour especially in male puppies by week 6 was an

interesting feature of this study, and was also observed by Rheingold (1963). Male puppies as

young as 6 weeks old may show sexual mounting even with pelvic thrusting. This can be

considered a normal part of play behaviour which is necessary for development of sexual

response in adults (Hart, 1980b). According to DeHasse (1994) and Scott and Fuller (1965) the

first signs of humping (pre-imitation of future sexual behaviour) appear as early as 3–4 weeks.

Although evidence is scant, it may be suggested that sexual behaviour in free-ranging dogs

develops at an early stage of life as pseudo-sexual behaviour, and predominantly in males.

The puppies were mobile by week 3, although few puppies before complete opening of their

eye slits, were able to reach their mothers from a distance of several cm (Rheingold, 1963). By

about 3 weeks of age, the puppies began to walk outside their nest. It has been suggested that free-

ranging dog puppies defecate and urinate away from their sleeping place (DeHasse, 1994). The

distance covered by the puppies gradually increased with the age of the puppies as previously


reported by Rheingold (1963). The distance covered by the puppies at a particular age recorded in

this study were nonetheless greater than those reported by Rheingold (1963). Previously,

Rheingold (1963), Macdonald and Carr (1995) and Pal et al. (1998b) reported that free-ranging

dogs are highly mobile at the age of 10–11 weeks. After weaning (10–11 weeks of life), the

puppies spontaneously and independently traveled an average distance of above 50 m. So, from

this study, it may be suggested that free-ranging dogs are mobile when the eyes are completely

opened (3 week of life) and they leave the nest. They are highly mobile when the suckling bouts

disappear (10–11 weeks of life, Pal, 2005) and they travel independently a distance of above 50 m

perhaps to collect food.

The result of Katwa study seems to agree in many aspects with the studies of Rheingold (1963)

and Lund and Vestergaard (1998); however, the development of certain processes like eye

opening, sexual behaviour, and locomotion seems to take place a little earlier in free-ranging

dogs than in the captive dogs reported by Rheingold (1963). Although, the sample sizes in this

study were too small and further research is needed, however, this study suggests that

development of some physical and social behaviour takes place a little earlier in free-ranging

dogs than in captive dogs.

Acknowledgements

I am sincerely grateful to Prof. E. Font who read the earlier drafts of this paper and improved

its quality, and also for his kind and generous help. I wish to thank the editor and two anonymous

referees for advice and comments on the earlier drafts of this article. My son Mr. Subhadip Pal

and my wife Dipti Pal assisted with fieldwork.

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