Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers
Black-tailed prairie dogs Cynomys ludovicianus (Ord, 1815) are diurnal rodents that live in intricate cities. Their social complexity rivals that of some primates, and, in some respects, resembles the behavior of humans. Due to the rich variety of readily-observable, sophisticated behaviors such as...
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Інститут зоології ім. І.І. Шмальгаузена НАН України
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irk-123456789-657942014-07-03T03:01:31Z Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers Eltorai, A.E.M. Sussman, R.W. Экология Black-tailed prairie dogs Cynomys ludovicianus (Ord, 1815) are diurnal rodents that live in intricate cities. Their social complexity rivals that of some primates, and, in some respects, resembles the behavior of humans. Due to the rich variety of readily-observable, sophisticated behaviors such as coloniality, infanticide, anti-predator behaviors, “kin recognition”, cooperation, conflict, and reproductive success, the black-tailed prairie dog is a wonderful model species for the study of behavior. Using a captive population, we were able to quantify the effects of observation on key social behaviors. Чернохвостые луговые собачки – Cynomys ludovicianus (Ord, 1815) – дневные грызуны, которые обитают в малодоступных местах. Сложность их социального поведения соперничает с таковой у некоторых приматов и в некотором отношении сходна с поведением человека. Благодаря богатому разнообразию легко наблюдаемых особенностей поведения, таких как колониальность, детоубийство, защита от хищников, распознавание родичей, сотрудничество, конфликты и репродуктивная успешность – чернохвостые луговые собачки являются прекрасным модельным видом для изучения поведения. Используя популяцию, обитающую в неволе, мы смогли количественно измерить влияние наблюдения на ключевые особенности социального поведения. 2011 Article Социальное поведение чернохвостых луговых собачек (Mammalia, Rodentia) в неволе при сменяющемся количестве наблюдателей / A.E.M. Eltorai, R.W. Sussman // Вестник зоологии. — 2011. — Т. 45, № 2. — С. 145–152. — Бібліогр.: 10 назв. — англ. 0084-5604 http://dspace.nbuv.gov.ua/handle/123456789/65794 591.55(599.322.2) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Экология Экология Eltorai, A.E.M. Sussman, R.W. Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers Вестник зоологии |
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Black-tailed prairie dogs Cynomys ludovicianus (Ord, 1815) are diurnal rodents that live in intricate cities. Their social complexity rivals that of some primates, and, in some respects, resembles the behavior of humans. Due to the rich variety of readily-observable, sophisticated behaviors such as coloniality, infanticide, anti-predator behaviors, “kin recognition”, cooperation, conflict, and reproductive success, the black-tailed prairie dog is a wonderful model species for the study of behavior. Using a captive population, we were able to quantify the effects of observation on key social behaviors. |
format |
Article |
author |
Eltorai, A.E.M. Sussman, R.W. |
author_facet |
Eltorai, A.E.M. Sussman, R.W. |
author_sort |
Eltorai, A.E.M. |
title |
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers |
title_short |
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers |
title_full |
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers |
title_fullStr |
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers |
title_full_unstemmed |
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of Observers |
title_sort |
social behavior of captive black-tailed prairie dogs (mammalia, rodentia) with changing numbers of observers |
publisher |
Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2011 |
topic_facet |
Экология |
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http://dspace.nbuv.gov.ua/handle/123456789/65794 |
citation_txt |
Социальное поведение чернохвостых луговых собачек (Mammalia, Rodentia) в неволе при сменяющемся количестве наблюдателей / A.E.M. Eltorai, R.W. Sussman // Вестник зоологии. — 2011. — Т. 45, № 2. — С. 145–152. — Бібліогр.: 10 назв. — англ. |
series |
Вестник зоологии |
work_keys_str_mv |
AT eltoraiaem socialbehaviorofcaptiveblacktailedprairiedogsmammaliarodentiawithchangingnumbersofobservers AT sussmanrw socialbehaviorofcaptiveblacktailedprairiedogsmammaliarodentiawithchangingnumbersofobservers |
first_indexed |
2025-07-05T16:11:18Z |
last_indexed |
2025-07-05T16:11:18Z |
_version_ |
1836824007341506560 |
fulltext |
UDC 591.55(599.322.2)
SOCIAL BEHAVIOR OF CAPTIVE BLACK-TAILED
PRAIRIE DOGS (MAMMALIA, RODENTIA)
WITH CHANGING NUMBER OF OBSERVERS
A. E. M. Eltorai, R. W. Sussman
Department of Anthropology Washington University
Campus Box 1114
One Brookings Drive
Str. Louis, MO, USA 63130
E-mail: aeltorai@gmail.com
Received 4 October 2010
Accepted 30 March 2011
Social Behavior of Captive Black-Tailed Prairie Dogs (Mammalia, Rodentia) with Changing Numbers of
Observers. Eltorai A. E. M., Sussman R. W. – Black-tailed prairie dogs Cynomys ludovicianus (Ord, 1815)
are diurnal rodents that live in intricate cities. Their social complexity rivals that of some primates, and,
in some respects, resembles the behavior of humans. Due to the rich variety of readily-observable, sophis-
ticated behaviors such as coloniality, infanticide, anti-predator behaviors, “kin recognition”, coopera-
tion, conflict, and reproductive success, the black-tailed prairie dog is a wonderful model species for the
study of behavior. Using a captive population, we were able to quantify the effects of observation on key
social behaviors.
Ke y wo r d s: Cynomys ludovicianus, black-tailed prairie dogs, captive, social behavior, observation.
Ñîöèàëüíîå ïîâåäåíèå ÷åðíîõâîñòûõ ëóãîâûõ ñîáà÷åê (Mammalia, Rodentia) â íåâîëå ïðè ñìåíÿþ-
ùåìñÿ êîëè÷åñòâå íàáëþäàòåëåé. Ýëòîðàé À. Å. Ì., Ñàññìàí Ð. Â. – ×åðíîõâîñòûå ëóãîâûå
ñîáà÷êè – Cynomys ludovicianus (Ord, 1815) – äíåâíûå ãðûçóíû, êîòîðûå îáèòàþò â ìàëîäîñòóï-
íûõ ìåñòàõ. Ñëîæíîñòü èõ ñîöèàëüíîãî ïîâåäåíèÿ ñîïåðíè÷àåò ñ òàêîâîé ó íåêîòîðûõ ïðèìà-
òîâ è â íåêîòîðîì îòíîøåíèè ñõîäíà ñ ïîâåäåíèåì ÷åëîâåêà. Áëàãîäàðÿ áîãàòîìó ðàçíîîáðàçèþ
ëåãêî íàáëþäàåìûõ îñîáåííîñòåé ïîâåäåíèÿ, òàêèõ êàê êîëîíèàëüíîñòü, äåòîóáèéñòâî, çàùèòà
îò õèùíèêîâ, ðàñïîçíàâàíèå ðîäè÷åé, ñîòðóäíè÷åñòâî, êîíôëèêòû è ðåïðîäóêòèâíàÿ óñïåøíîñòü –
÷åðíîõâîñòûå ëóãîâûå ñîáà÷êè ÿâëÿþòñÿ ïðåêðàñíûì ìîäåëüíûì âèäîì äëÿ èçó÷åíèÿ ïîâåäåíèÿ.
Èñïîëüçóÿ ïîïóëÿöèþ, îáèòàþùóþ â íåâîëå, ìû ñìîãëè êîëè÷åñòâåííî èçìåðèòü âëèÿíèå
íàáëþäåíèÿ íà êëþ÷åâûå îñîáåííîñòè ñîöèàëüíîãî ïîâåäåíèÿ.
Êëþ÷åâûå ñëîâà: Cynomys ludovicianus, ëóãîâûå ñîáà÷êè, ñîäåðæàíèå â íåâîëå, ñîöèàëüíîå ïîâå-
äåíèå, íàáëþäåíèå.
Introduction
“Prairie dogs have a complex social system, rivaling that of some primates and in some respects resem-
bling the behavior of humans. Perhaps that is why prairie dog exhibits in zoos are very popular, with people
sometimes spending hours watching the behavior of the animals” (Slobodchikoff et al., 2009: 43).
Black-tailed prairie dogs Cynomys ludovicianus (Ord, 1815) belong to the order of rodents and the squir-
rel family, Sciuridae. They are diurnal and live in socially-intricate colonies, when undisturbed, stretch for kilo-
meters and can contain thousands of individuals (Hoogland, 1995).
As a social species, black-tailed prairie dogs frequently interact among themselves. Friendly interactions
occur between individuals of the same family group, coterie. Examples of such friendly behaviors include: “play,
allogrooming, and mouth-to-mouth contact that resemble kisses” (Hoogland, 1995: 2).
The “greet-kiss” (King, 1955), where the prairie dogs “open up their mouths, and press their tongues
together for a brief period of time”, can occur between any combination of individuals (Slobodchikoff et al.,
2009: 56). Steiner proposed the following self-explanatory hypotheses for why this kiss may occur: Food
Information Hypothesis (1975), Individual Recognition Hypothesis (1974), and Dominance Maintenance
Vestnik zoologii, 45(2): 145—152, 2011 Ýêîëîãèÿ
Hypothesis (1975). However “the picture is still not clear about the functions of a greet-kiss. Perhaps the behav-
ior originated for any of the reasons given by the above hypotheses, and has persisted as a mechanism of social
reassurance, somewhat like a human kiss» (Slobodchikoff et al., 2009: 57).
Allogrooming, or mutual grooming, occurs when one individual picks the fleas, lice, and ticks from anoth-
er individual, consequently reducing the number of parasites on the groomed individual (Slobodchikoff et al., 2009).
Variations in how much time individual black-tailed prairie dogs spend doing particular behaviors have
been studied. Individually-distinct use of time serves to suggest that black-tailed prairie dogs may have unique
personalities, which effectively add to the nuanced prairie dog social system (Loughry, Lazari, 1994).
Less amicable behaviors are also observed. February through April, females are defensive of their nursery
burrows and hostile interactions frequently occur. However, when the pups emerge from their subterranean bur-
rows in May, friendly interactions are the coterie norm (Hoogland, 1995). The other form of hostile behavior
occurs “when prairie dogs from different coteries meet, they engage in a flagrant territorial dispute that involves
staring, tooth chattering, flaring of the tail, bluff charges, and reciprocal anal sniffing. Territorial disputes com-
monly persist for more than 30 minutes and sometimes include fights and chases as well” (Hoogland, 1995: 2).
From past studies of this North American ground squirrel, interesting observations have been reported
regarding coloniality, infanticide, anti-predator behaviors, “kin recognition”, cooperation, conflict, and
reproductive success. Prairie dogs are known to “have a sophisticated communication system that might out-
strip monkeys and apes in its complexity, ” making them a compelling study subject (Slobodchikoff et al., 2009:
1). Readily observable examples prairie dog behaviors include: the “jump-yip display” (individual stretches ver-
tically, throwing their body as they call), rapid scratching to remove fleas, burrow-mound enhancement, and
nest-building by collecting leaves, grass and twigs in their mouths (Hoogland, 1995). Due to the rich variety
of salient behaviors, the black-tailed prairie dog is a wonderful model species for the study of behavior.
Prairie dogs are even more fascinating study subjects considering the following observations: non-
parental prairie dogs make antipredator calls to warn distant kin; lactating females commit infanticide against
the offspring of close kin (sister, daughters)–resulting in the major source of pup death; mothers suckle the
offspring of female kin (the same pups they tried to kill earlier); and prairie dogs avoid inbreeding with close
kin while copulating with cousins (Hoogland, 1995).
Although the black-tailed prairie dog has been extensively studied in the wild, little literature exists regard-
ing captive black-tailed prairie dogs (Smith et al., 1973 appears to be the only captive study). “Visitor effect”
studies have focused primarily on primates (i. e. Hosey, 2005; Mitchell et al., 1992) while few studies have
observed the black-tailed prairie dogs’ response to visitors. Our previous work (Eltorai and Sussman, 2010) appears
to be the only study investigating black-tailed prairie dogs’ response to visitors.
In a study of captive black-tailed prairie dogs, Smith et al. (1973) found that summer months for cap-
tive black-tailed prairie dogs are characterized by “considerable disorganization but contained the seed of divi-
sion into groups” (p. 213). Unlike in the wild, “individuals other than the dominant pair in each zoo coterie
probably defend boundaries” (p. 214). These differences are most likely to be due to the relatively small amount
of space in the enclosure, the high population density, and the inability to emigrate leading to a high density
of individuals. Smith et al. also found that “the social behavior of the zoo prairie dogs is broadly comparable
to that of their wild counterparts” (p. 214). Studying black-tailed prairie dogs in captivity offers a unique oppor-
tunity to research a popular study species in an uncommonly studied setting.
In this study, we investigated the relationship between human observer density and the prairie dogs’ social
behaviors. Moreover, we looked at how observer density affected the behavior of the different-aged prairie dogs.
Ultimately, we hope our findings will help to further our understanding of the broader question: How does den-
sity of observers affect social behavior?
Methods
Subjects. The Saint Louis Zoo is home to approximately 25 black-tailed prairie dogs. The zoo exhibit
consists of mostly adult (determined by large size) prairie dogs and seven 1.5 month-old pups. The adults are
likely to range from three to five years of age. The sex of the individuals was not able to be determined. The
initial group of individuals was introduced to the exhibit in 1940. The present day zoo population is descen-
dent from the original group.
Data collection. Instantaneous scan samples were taken every 5 minutes to determine the number of indi-
viduals that were participating in a given behavioral category at the time. Using this sampling method, we were
able to observe the animals’ distribution within the enclosure at various points during the day; the number of
individuals physically touching another individual at a given time; the percentage of time the group spends on
a particular activity; and the visitor density at a given time. To account for normal changes in behavior due to
time of day, samples were taken at various, evenly-distributed points throughout the day (9AM-7PM).
Focal scan samples of an adult and a pup were taken every 5 minutes to see if there were any activity
and distributional differences with respect to age. Focal individuals were followed for as long as possible. When
the focal individual disappeared into one of the holes, another focal individual, of the same category (adult,
pup) was chosen to be followed. The replacement focal individual was chosen by being the most representa-
tive of the focal individual category at the time (i. e. resting adult in Region 4). Visitor number was found by
averaging the number of visitors measured every five minutes. The number of physically touching prairie dogs
is the sum of the every-five minute regional sampling measurements.
To further examine the focal individual categories, we used the all occurrence sampling method to focus
on specific social behaviors. Using this sampling method, we observed the duration and frequency of specific
behaviors of interest. When an individual went underground, the next focal categorized individual was chosen
146 A. E. M. Eltorai, Robert W. Sussman
by being the most representative of the current behavioral trends. The number of “socially close” individuals
to the focal individual was determined by being within a radius of two feet from the focal individual.
A digital watch was utilized to properly record the activities to the minute. All eighty-one hours of obser-
vation were collected from June through August, 2009 and detailed by hand using an appropriate data sheet.
All observations of the prairie dogs were above-ground and outside the burrows.
Enclosure. A map of the enclosure can be found in figure 1. The enclosure is approximately 75 � 40 feet.
There are relatively uniformly distributed hole-openings throughout the enclosure; the most frequently used
burrow entrances appear to be located in Regions 1 and 3. The ground is covered in wood chips. Food (gen-
erally: kale, Purina rat chow, sweet potatoes, and carrots; periodically: apples, bananas, and bamboo) is dis-
persed evenly throughout the enclosure each morning. The surrounding walls are textured rock; there is a climb-
ing path on the back wall that allows for the animals to reach elevated vantage points. There are several logs
of various shapes and sizes located in Regions 3, 4, and 5 that offer varied climbing, hiding and resting oppor-
tunities.
Statistics. Microsoft Excel was used to generate all of the plots and to determine the lines of best fit through
linear regression. The regions where there are overlapping visitor numbers were used when comparing adults
to pups on the same behaviors. To determine if adults vs. pups differed significantly (p < 0.05) from one anoth-
er on the same behaviors with respect to increased visitor density, two-tailed, independent samples Student’s
t-tests were used for figures 2—5, 7, and 8. Within-subjects ANOVAs were used to see if there are significant
differences for figure 6. Two-tailed probability values of the Pearson correlation coefficient r were found to deter-
mine, with respect to increased visitor density, the strength of the linear relationships. By plotting the behav-
iors vs. visitor density in a linear fashion, we understand that some of the nuanced changes will be overlooked.
We justify plotting the relationships as linear for the following two reasons: 1) it simplifies the data to cleanly
illustrate the major trends that are occurring; and 2) reduces the differences in the effects of the various types
of observers/zoo-goers (i. e. 12 quiet photographers vs. 12 screaming, crying third graders).
Results
In figure 2, we show the percent of time adults and pups perform various social behav-
iors with respect to changing visitor density. With increasing visitor density, adults spend
more time being alone. They spend less time being socially close to others. And when
they are around others it involves increased touching. Pups, on the other hand, with increas-
ing numbers of observers, spend less time being alone and more time being socially close
to others without touching.
There is a significant difference between percent of each hour adults vs. pups spend
being socially close to other prairie dogs without physically touching any others (p = 0.0062;
t (16) = 3.1488) and the percent of each hour spent touching other prairie dogs (p <
0.0001; t (16) = 5.5936), but no significant difference for the percent of each hour spent
being alone or being socially close to some and touching others.
In figure 3, we show adults’ and pups’ average duration of performing various
social behaviors with respect to increasing visitor density. When adults are socially close
to others, they do so for shorter periods of time with increasing number of observers. The
bout duration of adults being alone and physically touching increase as the number of
147Social Behavior of Captive Black-Tailed Prairie Dogs...
Fig. 1. Schematic map of enclosure showing how the space was divided into Regions 1—5 and where visitors
observed the prairie dogs.This image was reproduced from Eltorai and Sussman, 2010.
Ðèñ. 1. Ñõåìàòè÷åñêàÿ êàðòà îãîðîæåííîãî ó÷àñòêà, êîòîðàÿ ïîêàçûâàåò, êàê ïðîñòðàíñòâî áûëî ðàç-
äåëåíî íà ïëîùàäêè 1—5, è ãäå ïîñåòèòåëè íàáëþäàëè çà ëóãîâûìè ñîáà÷êàìè.
visitors increases. The pups’ average duration increased for all of the examined types of
social behaviors with increased observer number.
There is a significant difference between the durations of adults vs. pups being alone
(p < 0.0001; t (88) = 9.3870), being socially close but not touching (p < 0.0001; t (36)
= 16.8067), but no significant difference for physically touching only nor for being social-
ly close to some and physically touching others at the same time with increasing densi-
ty of observers.
In figure 4, we show that the average number of other prairie dogs that adults and
pups are socially close to mildly decreases as the number of visitors increases.
There is no significant difference between the average number of other prairie dogs
adults vs. pups are socially close to with respect to increasing visitor density.
In figure 5, we show that as the number of visitors increases the number of other
prairie dogs the focal adult physically touches remains constant, but the number of other
prairie dogs the focal pup touches increases mildly. Moreover, when adults touch oth-
ers, they are in contact with a greater number of individuals than pups.
There is a significant difference between the average number of other individuals adults
vs. pups physically touch (p = 0.0004; t (16) = 4.4241).
In figure 6, we look at the number of physically touching prairie dogs in each region
as the average number of visitors increases. With increasing visitor numbers the number
of physically touching prairie dogs increases in Regions 1 and 3; and decreases in
Regions 2, 4, and 5. As mentioned earlier, the main burrow entrances are located in Regions
1 and 3. Thus, it appears that with heavier observation, touching takes place closer to
burrow entrances.
There is a significant difference between number of physically touching prairie dogs
in Region 1 vs. 4 (p =0.040; F (15) = 5.113), Region 1 vs. 5 (p = 0.039, F (15) = 5.176),
148 A. E. M. Eltorai, Robert W. Sussman
Fig. 2. Minutes per hour adults and pups perform various social behaviors with respect to increasing visitor density.
Ðèñ. 2. Âðåìÿ,â òå÷åíèå êîòîðîãî âçðîñëûå îñîáè è äåòåíûøè äåìîíñòðèðîâàëè ðàçíîîáðàçíîå ñîöè-
àëüíîå ïîâåäåíèå ïî îòíîøåíèþ ê âîçðàñòàþùåìó êîëè÷åñòâó ïîñåòèòåëåé.
0
5
10
15
20
25
30
35
40
45
50
1 3 5 7 9 11 13 15 17 19 21 23 25
Number of visitors
M
in
ut
es
/ h
ou
r
Alone (Adult) (r = 0.3397)
Alone (Pup) (r = -0.4836)
Socially close but no touching (Adult) (r = -0.9892)
Socially close but no touching (Pup) (r = 0.5197)
Touching only (Adult) (r = 0.9886)
Touching only (Pup) (r = -0.4334)
Socially close and touching (Adult) (r = -0.2731)
Socially close and touching (Pup) (r = 0.4865)
Region 2 vs. 4 (p =0.004; F (15) = 11.967), and Region 2 vs. 5 (p = 0.004, F (15) =
11.594). There are no significant differences between for the number of physically touch-
ing prairie dogs in each region for Region 1 vs. Region 2, Region 1 vs. region 3, Region
2 vs. Region 3, Region 3 vs. Region 4, Region 3 vs. Region 5, and Region 4 vs. Region
5. In sum, there is a significantly more physical touching occurring in the front regions
vs. the more distant Regions 4 and 5.
Kissing and perianal sniffs are used both as greetings and a method for identifica-
tion. In the wild, if an individual from another coterie is encountered a bout of agonis-
tic behavior is likely to follow. In figure 7, we show that with increasing visitor density
the adults’ average number of mouth-to-mouth kisses per hour increases; the average num-
ber of perianal sniffs per hour decreases slightly; and the number of adult skirmishes decreas-
es. The data are based on the number of recorded events in which adults were undoubt-
edly in dispute. The skirmish behavior was classified by any combination of the follow-
ing: physical attacks, biting, wrestling, and teeth-bearing. On the other hand, with
increasing visitor density, pups’ average number of kisses and average number of peri-
anal sniffs per hour decreases, and the number of occurrences of pup wrestling very mod-
estly increases. Two important notes should be noted: 1) pup wrestling can either be viewed
as agonistic or playful and 2) the total number of pup wrestling occurrences is quite a
bit higher than the total number of adult skirmishes (43 vs. 12, respectively).
149Social Behavior of Captive Black-Tailed Prairie Dogs...
Fig. 3. The average duration of adults and pups performing various degrees of social behaviors with respect to
increasing visitor density.
Ðèñ. 3. Ñðåäíÿÿ ïðîäîëæèòåëüíîñòü äåìîíñòðàöèè ðàçíîîáðàçíîãî ñîöèàëüíîãî ïîâåäåíèÿ âçðîñëû-
ìè îñîáÿìè è äåòåíûøàìè â çàâèñèìîñòè îò âîçðàñòàþùåé ïëîòíîñòè ïîñåòèòåëåé.
0
0,5
1
1,5
2
2,5
3
3,5
4
4,5
5
1 3 5 7 9 11 13 15 17 19 21 23 25
Number of visitors
M
in
ut
es
Alone (Adult) ( r = 0.9949)
Alone (Pup) (r = 0.9994)
Socially close but no touching (Adult)
(r = -0.7963)
Socially close but no touching (Pup)
(r = 0.4590)
Touching only (Adult) (r = 0.7231)
Touching only (Pup) (r = 0.0721)
Socially close and touching (Adult)
(r = -0.9994)
Socially close and touching (Pup)
(r = 0.8493)
150 A. E. M. Eltorai, Robert W. Sussman
Fig. 4. The average number of other prairie dogs adults and pups are socially close to with respect to increasing
visitor density.
Ðèñ. 4. Ñðåäíåå êîëè÷åñòâî äðóãèõ îñîáåé áëèçêèõ ê âçðîñëûì îñîáÿì è äåòåíûøàì ëóãîâûõ ñîáà÷åê
â çàâèñèìîñòè îò âîçðàñòàþùåé ïëîòíîñòè ïîñåòèòåëåé.
0
1
2
3
1 3 5 7 9 11 13 15 17 19 21 23 25
Number of visitors
N
um
be
r
of
p
ra
ir
ie
d
og
s
Adult (r = -0.6288)
Pup (r = -0.4377)
Fig. 5. The number of other individuals the focal adult and pup physically touches with respect to increasing
visitor density.
Ðèñ. 5. Êîëè÷åñòâî ôèçè÷åñêè ñîïðèêàñàþùèõñÿ ñî âçðîñëûìè è äåòåíûøàìè èç ôîêóñíîé ãðóïïû ïðî-
÷èõ îñîáåé â çàâèñèìîñòè îò âîçðàñòàþùåé ïëîòíîñòè ïîñåòèòåëåé.
0
1
2
3
4
5
6
1 3 5 7 9 11 13 15 17 19 21 23 25
Number of visitors
N
um
be
r
of
p
ra
ir
ie
d
og
s
Adult (r = -0.08)
Pup (r = 0.2578)
151Social Behavior of Captive Black-Tailed Prairie Dogs...
Fig. 6. The number of physically touching prairie dogs in each region with respect to increasing visitor density.
Ðèñ. 6. Êîëè÷åñòâî ôèçè÷åñêè ñîïðèêàñàþùèõñÿ ëîãîâûõ ñîáà÷åê íà êàæäîì ó÷àñòêå â çàâèñèìîñòè
îò óâåëè÷èâàþùåéñÿ ïëîòíîñòè ïîñåòèòåëåé.
Fig. 7. With respect to increasing visitor density, the adults’ and pups’ average number of mouth-to-mouth kiss-
es and number of perianal sniffs per hour; the number of adult skirmishes; and the number of pup wrestling
bouts.
Ðèñ. 7. Èçìåíåíèå îñîáåííîñòåé ïîâåäåíèÿ ëóãîâûõ ñîáà÷åê ïî îòíîøåíèþ ê óâåëè÷èâàþùåéñÿ ïëîò-
íîñòè ïîñåòèòåëåé â ÷àñ.
0
5
10
15
20
25
1 4 7 10 13 16 19 22 25
Number of visitors
N
u
m
be
r
of
p
ra
ir
ie
d
og
s
to
uc
h
in
g
in
e
ac
h
re
gi
on
Region 1 (r = 0.9815)
Region 2 (r = -0.8907)
Region 3 (r = 0.9642)
Region 4 (r = -0.8245)
Region 5 (r = -0.5209)
0
5
10
15
20
25
30
35
1 3 5 7 9 11 13 15 17 19 21 23 25
Number of visitors
N
u
m
be
r
of
o
cc
u
re
n
ce
s
Mouth-to-mouth kisses (Adult) (r = 0.8538)
Mouth-to-mouth kisses (Pup) (r = -0.5445)
Perianal sniffs (Adult) (r = -0.9012)
Perianal sniffs (Pup) (r = -0.7457)
Adult skirmishes (r = -0.8259)
Pup wrestling bouts (r = 0.5638)
There is a significant difference between number of adult skirmishes vs. number of
pup wrestling bouts (p < 0.0001; t (14) = 12.6557) with adults being “more social. ” There
are no significant differences between average number of mouth-to-mouth kisses per hour
or average number of perianal sniffs per hour adult vs. pup with respect to increasing vis-
itor density.
Discussion
With increasing observation, adult prairie dogs touch a relatively constant absolute
greater number of other individuals and more frequently than pups do. However, pups
touch an increasingly greater number with more observers. Additionally, we showed that
with increasing visitors the number of physically touching prairie dogs increases on the
left side of the enclosure (Regions 1 and 3) and decreases on the right side (Regions 2,
4, and 5). We hypothesize that this may have to do with the main burrow entrances being
located, in higher concentration, on the left side of the enclosure. Thus, the prairie dogs
tend to touch more in the presence of increasing visitor numbers, but they do so close
to home. This finding may indicate that touching relates to an altered state of vulnera-
bility. Consequently, the prairie dogs adjust by situating themselves closer to escape routes.
Increases in visitor numbers appears to impact adults and pups differently. Adults
spend more of their time being alone while pups spend more of their time being social-
ly close to others. When adults are around other prairie dogs it involves increased touch-
ing. Duration of being alone or touching others increases for both adults and pups when
visitor numbers increase. With increased observation, adults kiss more and fight less, where-
as pups kiss less and fight more.
A comparison of pup to adult social behavior reveals different trends in each age cat-
egory. We hypothesize that these differences are due to learning. In other words, the pups
have not completely learned how to properly behave socially. To draw a broader con-
clusion from the data, it appears that pups demonstrate social behaviors immediately, but
“proper” social behavior takes time and is consequently learned. To help to further inves-
tigate this question, data could be collected on intermediately-aged prairie dogs to see if
their social behavior demonstrates a transitional phase between pups and adults.
If the act of observation changes behavior, then shouldn’t all observational research
findings be viewed with increased skepticism?
We thank John Hoogland for his encouragement and suggestions for our research and the St. Louis zoo
keepers for information regarding the prairie dog exhibit’s history.
Eltorai A. E. M., Sussman R. W. The “Visitor Effect” and captive black-tailed prairie dog behavior // Der
Zoologische Garten. – 2010. – 79. – P. 109—120.
Hoogland J. L. The Black-Tailed Prairie Dog : Social Life of a Burrowing Mammal. – Chicago : University
of Chicago Press, 1995.
Hosey G. R. How does the zoo environment affect the behavior of captive primates? // Applied Animal Behavior
Science. – 2005. – 90. – P. 107—129.
King J. A. Social behavior, social organization, and population dynamics in a black-tailed prairie dog town in
the Black Hills of South Dakota // Contributions from the Laboratory of Vertebrate Biology, University
of Michigan. – 1955. – 67. – P. 1—123.
Loughry W. J., Lazari A. The ontogeny of individuality in the black-tailed prairie dogs, Cynomys ludovicians //
Canadian Journal of Zoology. – 1994. – 72. – P. 1280—1286.
Mitchell G., Tromborg C. T., Kaufman J. et al. More on the ‘influence’ of zoo visitors on the behaviour of cap-
tive primates // Applied Animal Behaviour Science. – 1992. – 35. – P. 189—198.
Slobodchikoff C. N., Perla B. S., Verdolin J. L. Prairie Dogs: Communication and Community in an Animal
Society. – Cambridge : Harvard University Press, 2009.
Smith W. J., Smith S. L., Oppenheimer E. C., de Villa J. G., Ulmer F. A. Behavior of a Captive Population of
Black-Tailed Prairie Dogs. Annual Cycle of Social Behavior // Behaviour. – 1973. – 46. – P. 189—220.
Steiner A. L. Body-rubbing, marking, and other scent-related behavior in some ground squirrels (Sciuridae):
A descriptive study // Canadian J. Zoology. – 1974. – 52. – P. – 889—906.
Steiner A. L. “Greeting” behavior in some sciuridae, from ontogenetic, evolutionary, and socio-behavioral per-
spective // Naturaliste Can. – 1975. – 102. – P. 737—751.
152 A. E. M. Eltorai, Robert W. Sussman
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/AddPageInfo false
/AddRegMarks false
/ConvertColors /ConvertToCMYK
/DestinationProfileName ()
/DestinationProfileSelector /DocumentCMYK
/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /MediumResolution
>>
/FormElements false
/GenerateStructure false
/IncludeBookmarks false
/IncludeHyperlinks false
/IncludeInteractive false
/IncludeLayers false
/IncludeProfiles false
/MultimediaHandling /UseObjectSettings
/Namespace [
(Adobe)
(CreativeSuite)
(2.0)
]
/PDFXOutputIntentProfileSelector /DocumentCMYK
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
/UntaggedRGBHandling /UseDocumentProfile
/UseDocumentBleed false
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice
|