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Differences in winter activity, courtship, and social behavior of two captive family groups of Mexican wolves (Canis lupus baileyi)

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The purpose of this study was to determine differences in activity patterns and social behavior of two groups of endangered Mexican wolves maintained at two quite different facilities and to determine some of the variables that should be considered when making specific behavioral comparisons among wolves in this binational captive breeding program. Quantitative measurements of an Activity Index and social behaviors were obtained for three individuals in each pack. Within each age/sex category, activity, aggression, and play were more frequent in the pack at a zoo facility, compared to the pack at a field station facility. Frequency of courtship interactions and scent marking were significantly higher in the field station pack. The packs were similar in the frequency of active submission, but differed significantly in the pattern of this behavior. Given the large number of interacting variables and small number of individuals in this study, we recommend caution in generalizing results to other packs or facilities. Zoo Biol 16:435–443, 1997. © 1997 Wiley-Liss, Inc.
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Zoo Biology 16:435–443 (1997)
ZOO 527
© 1997 Wiley-Liss, Inc.
Differences in Winter Activity, Courtship,
and Social Behavior of Two Captive
Family Groups of Mexican Wolves
(Canis lupus baileyi)
José F. Bernal* and Jane M. Packard
Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas
The purpose of this study was to determine differences in activity patterns and
social behavior of two groups of endangered Mexican wolves maintained at two
quite different facilities and to determine some of the variables that should be
considered when making specific behavioral comparisons among wolves in this
binational captive breeding program. Quantitative measurements of an Activity
Index and social behaviors were obtained for three individuals in each pack.
Within each age/sex category, activity, aggression, and play were more frequent
in the pack at a zoo facility, compared to the pack at a field station facility.
Frequency of courtship interactions and scent marking were significantly higher
in the field station pack. The packs were similar in the frequency of active sub-
mission, but differed significantly in the pattern of this behavior. Given the large
number of interacting variables and small number of individuals in this study,
we recommend caution in generalizing results to other packs or facilities. Zoo
Biol 16:435–443, 1997. © 1997 Wiley-Liss, Inc.
Key words: activity; social behavior; Species Survival Plan; canid, wolf
INTRODUCTION
Behavior is an important consideration in making management decisions to
implement the goals of a Species Survival Plan [Kleiman, 1994]. For Mexican wolves,
behavioral interactions in a naturalistic enclosure at one field site [Servin, 1991],
general behavior at one zoo, and physiological norms have been reported for Mexi-
can wolves [Reyna Medrano et al., 1993]. However, norms of behavior have not yet
been established for Mexican wolves as has been done for northern subspecies of
wolves [MacDonald, 1980; Packard, 1980; Kreeger et al., 1996].
Received for publication 6 February 1995; revision accepted 2 June 1997.
*Correspondence to José F. Bernal-Stoopen and Dr. Jane M. Packard, Department of Wildlife and
Fisheries Sciences, Texas A&M University, College Station, Texas 77843-2258.
436 Bernal and Packard
A binational program of captive propagation (Mexico and USA) has been quite
successful for Mexican wolves [Siminski, 1993]. At the time this study was con-
ducted (winter 1990), there were only two breeding packs in captivity in Mexico. By
July 1996, the population had increased to 149 Mexican wolves in 30 facilities in
Mexico and the United States [Siminski, P., personal communication], and plans were
in progress for reintroduction of surplus animals to at least one site in the United
States [Parsons and Nicholopolous, 1995].
Although the value of direct observation for assessing behavioral profiles has
been acknowledged, statistical comparisons of behavioral differences of animals at
different facilities present certain difficulties. The purpose of this study was to deter-
mine differences in activity patterns and social behavior of one family group of Mexi-
can wolves maintained on-exhibit at San Juan de Aragón Zoo (SJA Zoo) and another
of similar social structure maintained off-exhibit in a naturalistic enclosure at San
Cayetano (SC Field Station). The zoo exhibit represents minimally enriched design,
whereas the field station enclosure represents a style thought to be suitable for adapting
potential candidates for release from captivity. Our objectives were to assess indi-
vidual variation in wolves at each facility related to age/sex category, time of day,
and location.
METHODS
Each pack consisted of a breeding male, breeding female, and one juvenile
offspring. The history of each pack differed prior to and after the study (Fig. 1).
The exhibit at SJA Zoo (approximately 170 m2) was located in Mexico City
and was open to the public daily between the hours of 0900 and 1700. Its completely
concrete substrate included a dry moat and an upper platform with two indoor pens
(3 m × 2 m). The pack was fed 6 days a week (primarily horse meat supplemented
with chicken parts). Food was provided after the enclosure was cleaned in the morn-
ing, and the wolves were kept during the night in off-exhibit holding pens. General
health care procedures have been described by Ramos Magaña et al. [1993].
In contrast, the SC Field Station was located within pine and oak woods in the
State of Mexico [Ferrusquia-Villafranca, 1993] and was not open to the general pub-
lic. The enclosure (1.3 ha.) provided a diverse, naturalistic environment with soil,
vegetation, a stream, and two holding pens for feeding. Feeding was 6 days per
week, with commercial dog chow supplemented occasionally with chicken parts.
During the study period, the animals were never captured and entry into the enclo-
sure was avoided as much as possible [Contreras, C., personal communication].
Both packs were observed by the senior author, using continuous focal group
sampling and a tape recorder [Martin and Bateson, 1993]. The pack at the SJA Zoo
was observed (62 hrs) for 2 days a week over 6 weeks during the months of January
to March (1990). The pack at the SC Field Station was observed (161 hrs) between 3
and 5 days a week from December (1989) to April (1990). Hours were distributed
unequally among the following three observation periods defined for analysis: morn-
ing (0700–0959), midday (1000–1259), and evening (1300–1900).
Standard definitions of behavioral categories [Packard, 1980; Zimen, 1981;
Servin, 1991; Derix, 1994] and observational techniques [Martin and Bateson, 1993]
were used. Activity was recorded by scan (sequential instantaneous) sampling with a
5-min sampling interval. The following behaviors were collapsed into the category
Activity Patterns and Social Behavior of Wolves 437
Fig. 1. The histories of the Mexican wolf packs at SJA Zoo and SC Field Station. Symbols represent birth (B), death (D with age in days, weeks
or months), transfers (T), genetic relationships (dashed lines), copulations (dots), presence in pack (solid lines), sex of individuals (male, female or
undetermined), and studbook identities (numbers in circles).
438 Bernal and Packard
of activity: walking, pacing, running, eating, drinking, scent-marking, and interact-
ing. The occurrence of double scent-marking [Mertl-Millhollen et al., 1986] was re-
corded as an index to evaluate the strength of pair-bonding. Double scent-marking
was defined as raised-leg urinations that were oriented on the same spot by both
members of the breeding pair within an observation period. The frequency of court-
ship interactions, scent-marking, active submission, play interactions, and aggressive
interactions were continuously recorded as all-occurrence event sampling. Courtship
behaviors included the following mutually exclusive categories: dance, genital in-
spection, tail avert, mount attempt, and copulatory-tie. Scent-marking included raised-
leg urination, scraping, scent-rolling, and defecation. Social interactions were
categorized as aggression, active submission, and play.
The Activity Index was based on the ratio of instants recorded as active, di-
vided by the total instants an animal was observed. To determine significant differ-
ences in the Activity Index of age/sex categories in each pack, the data were partitioned
by time periods (morning, midday, evening), and the Friedman Test [Lehner, 1979]
was applied, using a computer program [Abacus Concepts, Inc., 1992]. The effect of
time period on activity was analyzed separately for each individual, using the Kruskal-
Wallis Test of Mean Rank [Lehner, 1979]. To examine the effect of location on ac-
tivity, pairwise comparisons were made only between adults in each pack, using the
Mann-Whitney U Test [Lehner, 1979].
To compare the frequency of social behaviors within age/sex categories be-
tween packs, counts for each behavioral category were compared across individuals
using the Chi-square Goodness of Fit Test [Conover, 1980], adjusting expected values to
account for the unequal distribution of observation hrs between packs. The Log-likeli-
hood Ratio Test (G2) and Binomial Test (z) were calculated for more detailed analysis of
behavioral contingencies [Gottman et al., 1990], using a spreadsheet.
RESULTS
The two packs differed in terms of the relative activity patterns of individuals
(Fig. 2; SC Field Station, χ2 = 35.74, P = 0.0001; SJA Zoo, χ2 = 2.27, P = 0.3).
Individuals were not equally likely to be active at each of the three time periods
(Table 1). Activity levels of males differed significantly across packs for morning (U
= 175, P = 0.001) and midday (U = 58.5, P = 0.04), but not for evening periods (U =
74, P = 0.7), due to high activity of the male at the zoo. In a similar pattern, the
activity levels of adult females differed significantly for morning (U = 21, P = 0.001)
and midday (U = 46, P = 0.006), but not for evening periods (U = 65.5, P = 0.5).
Courtship interactions were more frequent at the SC Field Station than at the
SJA Zoo (Table 2). In the SJA Zoo, dance was lower than expected (z = –3.70) and
genital inspection was higher than expected (z = 3.0). The total frequency of mount
attempts was not different than expected by chance in both packs: SJA Zoo (z = –0.22)
and SC Field Station (z = 0.19). The reproductive period was unusually delayed at
the SC Field Station and two peaks of courtship occurred (Fig. 3). The highest peak
was recorded 2 weeks before estrus when the female was not fully receptive to the
male, yet received genital inspection and mounting attempts. During the estrus pe-
riod that resulted in conception (March 29–April 2), distinctive action patterns in-
cluded tail avert and copulation.
In contrast to similarities between the breeding females in each pack, the breed-
Activity Patterns and Social Behavior of Wolves 439
Fig. 2. The relative activity of individuals at the SC Field Station (shaded bars) and SJA Zoo (diago-
nally hatched bars). Horizontal bars indicate means, vertical bars indicate 95% confidence intervals.
TABLE 1. Relative activity patterns differed significantly among different periods of the day for
individuals in Mexican wolf packs at the SJA Zoo and SC Field Station
Activity index
(mean ± 95% confidence interval) Kruskal-Wallis
Pack Morning Midday Evening test of mean ranka
Individual (n = 8) (n = 13) (n = 5) HaP
SJA Zoo
Breeding male 96.9 ± 3.6 64.8 ± 21.9 63.6 ± 37.0 8.46 0.0146
Breeding female 92.1 ± 6.8 68.8 ± 19.5 60.9 ± 35.8 11.07 0.0039
Juvenile 98.2 ± 2.7 74.0 ± 18.2 80.0 ± 36.4 7.56 0.0283
SC Field Station
Breeding male 56.9 ± 12.0 22.4 ± 17.7 50.6 ± 9.3 10.87 0.0044
Breeding female 33.5 ± 15.6 8.1 ± 14.7 37.5 ± 13.1 18.52 0.0001
Juvenile 74.8 ± 14.6 35.0 ± 23.7 76.4 ± 9.2 11.08 0.0039
aThe statistic was corrected for ties.
ing males differed significantly in frequency of scent-marking (Table 2). In SJA Zoo,
raised-leg urination was higher than expected (z = 7.09) and scraping was lower than
expected (z = –5.11). Relative to her partner, the female at San Cayetano did more
raised-leg urination (z = 2.29) and scent-rolling (z = 2.77) and less scraping (z = –3.46).
Between January 28, 1990, and March 25, 1990, 10 double scent-marks were ob-
served at the SJA Zoo, compared to 14 double scent-marks observed from December
29, 1989, to April 4, 1990, at the SC Field Station.
The packs differed significantly in total aggression and the pattern of aggres-
440 Bernal and Packard
TABLE 2. Frequency of social interactions and scent-marking in packs at SJA Zoo (SJA) and SC
Field Station (SC)
Comparison ObservedaChi-square Significance
Behavior Pack Individuals (expected) goodness of fit (P value)
Courtship SJA Male/female 122 (81) 29.4 <0.001
SC Male/female 166 (207)
Scent-marking SJA Male/female 118 (102) 3.4 >0.05
SC Male/female 247 (263)
Raised-leg SJA Male 112 (63) 52.9 <0.001
urination SC Male 113 (162)
Aggression SJA Male/female 60 (30) 41.8 <0.001
SC Male/female 47 (77)
Active submission SJA Juvenile 87 (55) 26.7 <0.001
SC Juvenile 108 (141)
Play SJA Juvenile 182 (72) 235.7 <0.001
SC Juvenile 74 (184)
aExpected values were calculated by multiplying the total number of events by the proportion of total
hours that each pack was observed.
Fig. 3. The courtship activity of Mexican wolf pairs at the SJA Zoo and at the SC Field Station
during the 1989–1990 breeding season. The thick horizontal bars indicate periods of estrus as deter-
mined by the occurrence of the tail-avert action.
Activity Patterns and Social Behavior of Wolves 441
sion (Table 2). In the SJA Zoo, male-to-juvenile aggression was higher than ex-
pected (z = 2.39) and female-to-male aggression was lower than expected (z = –4.5).
At the SC Field Station, the male was not aggressive to the female; aggression of the
male to the female at the SJA Zoo occurred, but infrequently.
The packs did not differ significantly in total frequency of active submission
(Table 2), but differed significantly in the pattern of active submission (G2 = 8.18; df
= 1; P < 0.01). In the SJA Zoo, active submission from juvenile to male was lower
than expected (z = –2.12), but was higher than expected (z = 2.12) between juvenile
and female. At the SC Field Station, the juvenile was equally submissive to the adult
male and female. Play behavior was significantly more frequent at the SJA Zoo than
at the SC Field Station (Table 2). At the SC Field Station, play between the male and
juvenile was higher than expected (z = 8.02).
DISCUSSION
The direct-observation techniques employed in this study were useful for com-
paring activity and frequency of specific behaviors across packs for individuals in
similar age/sex categories. In making statistical comparisons, we found that it was
necessary to account for differences in activity relative to time of day and differ-
ences in the total time each pack was observed.
Initially, we were surprised that activity was greater at the zoo than the field
station, because we expected the natural vegetation to function as behavioral enrich-
ment. However, considering the higher pattern of morning activity in the zoo, we
hypothesize high activity could have been associated with the daily pattern of clean-
ing, occasional separation at night, and/or stimulation by vocalization of coyotes
(Canis latrans) in an adjacent pen. In addition, observation conditions also varied
between both enclosures; visibility at the Field Station Pack was obscured by veg-
etation and wolves may have been active when they were out of view.
We interpret the differences in courtship behavior in the two packs as more
related to individual differences (and possible pathology) than to the physical environ-
ment at the two facilities. The pattern of courtship and estrus shown by the pair at the zoo
was more like that described for other subspecies [Packard, 1980]. Although the higher
frequency of scent-marking, rolling, and scraping at the Field Station might have been
related to the soft substrate and vegetation, it was not indicative of the pattern usually
associated with a strong pair-bond. Double scent-marking is usually considered indica-
tive of a strong bond in wolves. Although it occurred in both packs, it was low relative to
the prolonged courtship and delayed estrus at the field station. The relatively low fre-
quency of raised-leg urination and high frequency of scraping by the male at the field
station also deviated from the norm. Since the policy was “hands off” at the field station,
no physiological data were available to assess possible pathology.
The relationships between the juvenile and breeding male differed greatly be-
tween packs. The breeding male at the zoo was more likely to direct aggression and
less likely to receive active submission (derived from food begging) and play from
the juvenile. In the wild, both the male and female actively participate in the rearing
of cubs [Packard et al., 1992]. We interpret this difference in juvenile/adult relation-
ships as related to the practice of isolating the female at the SJA Zoo prior to parturition
and during the first 3 months postpartum. At the SC Field Station, both the breeding
male and female were present and participated actively in the rearing of the pup.
442 Bernal and Packard
RECOMMENDATIONS
Direct behavioral observations are useful in making comparisons between indi-
vidual wolves living in different facilities. Comparisons at the two sites indicate the
zoo facility is more appropriate for intensive husbandry and production of pups.
However, the practice of separating the male and female near parturition should be
considered carefully, since the social bond between juvenile and adult male is impor-
tant to the goals outlined by the Species Survival Plan. In these two packs, the quiet,
undisturbed, naturalistic environment of the larger enclosure did not stimulate the
activity that may be required for physical conditioning of wolves selected for rein-
troduction. Similar techniques of comparing activity and social behavior might be
used in choosing between individuals as candidates for reintroduction and assessing
the suitability of individuals and pairs for reintroduction.
ACKNOWLEDGMENTS
We thank the Secretaría de Desarrollo Social (SEDESOL) and the San Juan de
Aragón Zoo for their support during the study. Specifically we thank the following
persons for their enthusiasm during the study: Luis Gerardo López Lemus, Carlos
Contreras, Gerardo Tapia, Ariel Alarcon, José Maria Reyes, Eleazar Loa, Mauro
Reyna, Patricia Reyes, Gerardo López Islas, Bernardo Manriquez, Guillermo Islas,
and Fernando Camacho. We also thank Gerardo López Islas, Fernando Cervantes,
Gerardo Ceballos, Mauro Ivan Reyna, Carmen Vazquez, Wendy Brown, Peter
Siminski, and Ed Spevak for reviewing earlier versions of the manuscript. This pa-
per is dedicated to the memory of Josefa, a very special Mexican wolf.
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... In a study comparing the amount and frequency of activity in gray wolves housed in small artificial enclosures against large natural enclosures, show no significant differences (Kreeger et al. 1996). In contrast, another study, during the winter of 1989-1990, evaluated the differences in the social behavior of two family groups of Mexican wolves, comparing a typical zoo facility and a field station facility, reported that for each of age and sex category, activity, aggression, and play are more frequent in the zoo group than in the field station group, while the frequency Downloaded by [Universidad Autonoma Metropolitana] at 14:23 31 March 2016 of interactions of courtship and scent marking are significantly higher in the field station group that in the zoo group (Bernal & Packard 1997). In both studies, given the small size of the sample can not be generalized the results (Kreeger et al. 1996;Bernal & Packard 1997). ...
... In contrast, another study, during the winter of 1989-1990, evaluated the differences in the social behavior of two family groups of Mexican wolves, comparing a typical zoo facility and a field station facility, reported that for each of age and sex category, activity, aggression, and play are more frequent in the zoo group than in the field station group, while the frequency Downloaded by [Universidad Autonoma Metropolitana] at 14:23 31 March 2016 of interactions of courtship and scent marking are significantly higher in the field station group that in the zoo group (Bernal & Packard 1997). In both studies, given the small size of the sample can not be generalized the results (Kreeger et al. 1996;Bernal & Packard 1997). ...
... mil/data) based on the geographical coordinates and the posterior adjustment in time to the real hour in both locations. Also we consider the period of daylight (daylight, twilight and nocturnal) and the age of individuals (juveniles, non-breeding adults, breeding adults and old wolves) because previous studies have shown to be factors influencing the activity of canids (Bernal & Packard 1997;Jedrzejewski et al. 2001;Merrill & Mech 2003;Siwak et al. 2003;Theuerkauf et al. 2003;eggermann et al. 2009). Astronomical parameters considered to determine the period of daylight were: times of sunrise and sunset, begin and end of astronomical twilight, and daylight and darkness periods, as well as twilight duration. ...
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Thesis
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En la actualidad, los cuatro objetivos de los zoos modernos son la educación, la investigación, la conservación y el ocio. Esta tesis se centró en la evaluación de algunos indicadores de bienestar animal en cuatro especies de mamíferos en condiciones de cautividad (el patrón de actividad diario, la ocurrencia de conductas aberrantes, el uso del espacio y las interacciones sociales). Los principales resultados obtenidos en este estudio pueden resumirse en: 1) Las conductas aberrantes en dos hembras de oso pardo (Ursus arctos) en cautividad mostraron variaciones estacionales al igual que lo observado en las conductas típicas de la especie en condiciones de libertad; 2) En el oso pardo (U. arctos) la presencia de público provocó un aumento en la ocurrencia de estereotipias y un uso del espacio más homogéneo, mientras que en el oso panda gigante (Ailuropoda melanoleuca) no se observaron estereotipias ni una influencia del público en el uso del espacio; 3) La seminaturalización de la instalación de dos individuos de oso pardo (U. arctos), un macho y una hembra, provocó que únicamente el macho alcanzara un nivel de actividad similar al observado en sus conspecíficos en libertad para la misma época del año (verano), haciendo un uso más homogéneo del espacio disponible. Los dos ejemplares de oso pardo estudiados respondieron de forma diferente al enriquecimiento estructural; 4) El estudio de la eficacia de un programa de enriquecimiento alimentario, sensorial y ocupacional en tres individuos de oso pardo (U. arctos), puso de manifiesto cuatro modelos de evaluación de dichos programas: el modelo de la ganancia, el la habituación, el la continuidad y el de la fluctuación de la eficacia del ítem; 5) La muerte del macho alfa en una manada de lobos ibéricos (Canis lupus signatus) provocó un aumento de las conductas de “vigilancia” y “no visible”, así como un uso del espacio menos homogéneo; 6) Las interacciones maternofiliales en tres madres y tres crías de león marino de California (Zalophus californianus) en cautividad mostraron varias semejanzas con las que presenta la especie los en libertad (los individuos se mantuvieron inactivos durante la mayor parte del tiempo y las crías macho presentaron mayores episodios lúdicos que las hembras). Los factores que influyen en el bienestar de los animales de este estudio deberían tenerse en cuenta para garantizar que las condiciones de cautividad son adecuadas para los animales. Estos factores son las condiciones sociales de la especie, el tamaño y el diseño idóneo de la instalación, los requerimientos climatológicos de la especie, la composición de la dieta, la estimulación física y/o psicológica, la influencia del público y los cuidados veterinarios. La clave para garantizar el bienestar de los animales radica en estudiar minuciosamente las necesidades físicas y psicológicas de los mismos a través del conocimiento tanto de la historia natural de las especies como de los trabajos realizados en otros zoos.
... This, paired with the fact that wolves engage in social play both as puppies and as adults, within and between age groups, suggests they may be a particularly good species to study equity within the play context. However, only a few studies on wolf play behavior have been carried out, with most of these studies focusing on adult animals, and none looking specifically at inequity aversion or equality within the play context [25][26][27][28][29]. In the current study, we investigate play behavior in wolves as puppies in relation to both the '50:50' rule, and in relation to their dominance relationship outside of play. ...
... This, paired with the fact that wolves engage in social play both as puppies and as adults, within and between age groups, suggests they may be a particularly good species to study equity within the play context. However, only a few studies on wolf play behavior have been carried out, with most of these studies focusing on adult animals, and none looking specifically at inequity aversion or equality within the play context [25][26][27][28][29]. In the current study, we investigate play behavior in wolves as puppies in relation to both the '50:50' rule, and in relation to their dominance relationship outside of play. ...
Article
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Social play is known as a cooperative interaction between individuals involving multiple mechanisms. However, the extent to which the equality of individuals' play styles affects the interaction has not been studied in many species. Dyadic play between wolf puppies, as well as between puppies and adults, was studied to investigate both self-handicapping and offensive behaviors to determine the extent to which wolves engage in play styles where one individual does not dominate the play. Our results did not support the hypothesized '50:50' rule, which suggests that more advantaged individuals should show higher rates of self-handicapping behaviors in order to facilitate play with others. Adult wolves performed significantly less self-handicapping behaviors than their puppy partners, and they performed significantly more offensive behaviors than their puppy partners. While the '50:50' rule was not supported at any time during our study period, dyads consisting of two puppies had significantly more equal play than dyads consisting of one puppy and one adult. These results suggest that wolf puppies are more likely to play on equal terms with similarly-aged play partners, while the dominance status of the partners dictates offensive and self-handicapping behaviors between animals of different ages.
... En el año 1991 Servín estudia los cambios comportamentales de una manada de esta especie a lo largo de un año, teniendo en cuenta la época de reproducción. Bernal y Packard (1997) comparan las frecuencias de las conductas de una manada de lobo mejicano en condiciones de cautividad y otra mantenida en condiciones de semilibertad. ...
Article
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Social dynamics is one of the most widely studied fields in the Canidae family in captivity. This study aims to determine how the death of the dominant male in a pack of wolves at Barcelona Zoo affects the pattern of daily activity and the use of space of the rest of the members. The individuals under study are two males and two females of the species Canis lupus signatus in naturalized surroundings. The observations were made using an instantaneous multifocal recording every 15 minutes. Each individual was observed for a total of 55.5 hours both prior to and after the death of the alpha male. The results for daily activity patterns show an increase in the behaviours of Vigilance, Maintenance and Lack of Visibility and a reduction in Inactivity. As regards the use of space, the use of the left central area increased, the use of the front central area decreased, while there were no differences in the use of the rear left area. The location of the animals was less homogeneous after the death of the alpha male.
... Its natural distribution is the most southerly and it is genetically different to the other four (García-Moreno et al., 1996;Parsons, 1996), but now considered extinct in the wild. Most remaining individuals live in zoos and are part of a Binational Conservation Programme between Mexico and the US (Bernal and Packard, 1997;List, 2005). ...
Article
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The effect of zoo visitors on the behaviour and physiological responses of the Mexican wolf has not been documented but is worthy of investigation since it is a critically endangered native species and most of the population live in zoos. The effect of the number of visitors was assessed in 12 wolves (five males and seven females) in three zoos of central Mexico. Each wolf was observed for a total of 72 h on Saturdays, Sundays, Mondays and Tuesdays (6. h/day) during a three-week period. Scan sampling was used to obtain individual time budgets. A faecal sample was collected on the following morning of each observation day from 10 wolves to determine cortisol level (RIA). A repeated measures analysis of variance revealed that there was an effect of day on the proportion of time lying (P<0.01), eating (P<0.01) and in locomotion (P<0.01). Wolves spent less time lying on Saturdays and Sundays (0.2980 ± 0.012 and 0.3266 ± 0.012, respectively) than on Mondays and Tuesdays (0.3821 ± 0.012 and 0.4075 ± 0.012, respectively), as well as less time eating on weekend days (Saturdays 0.1214 ± 0.015, Sundays 0.0556 ± 0.015) than in the other 2. days (Mondays 0.816 ± 0.015 and Tuesdays 0.0915 ± 0.015). Mean faecal cortisol was 203.7 ng/g of DM and ranged from 113.10 to 314.48. ng/g. There was an effect of day on the faecal cortisol response (P<0.01) with wolves having higher cortisol response on Sundays (242.50 ng/g ± 8.48) compared to the other 3. days (Saturdays 192.71 ± 8.48, Mondays 170.73 ng/g ± 7.80, Tuesdays 183.82 ng/g ± 8.48). This is the first study that measures faecal cortisol of Mexican wolves. These results indicate that the amount of visitors in zoos influence the behaviour and adrenal activity of these animals which could be undesirable for ex situ conservation efforts of this endangered species.
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Pair bonding is a psychological construct that we attempt to operationalize via behavioral and physiological measurements. Yet, pair bonding has been both defined differently in various taxonomic groups as well as used loosely to describe not just a psychological and affective phenomenon, but also a social structure or mating system (either social monogamy or just pair living). In this review, we ask the questions: What has been the historical definition of a pair bond? Has this definition differed across taxonomic groups? What behavioral evidence do we see of pair bonding in these groups? Does this observed evidence alter the definition of pair bonding? Does the observed neurobiology underlying these behaviors affect this definition as well? And finally, what are the upcoming directions in which the study of pair bonding needs to head?
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The aim of the study was to analyse the scent marking patterns in frequency, type and position throughout the reproductive period of two pairs of Canis lupus baileyi housed in two zoos (LZ = Leon and ZZ = Zacango), in terms of newly and established pairs. Focal behaviour sampling was used to register 1,211 scent markings. Data were grouped in three periods: before, during, after and posterior to the reproductive season. Between newly formed pairs no significant difference was found in the previous period (U = 6.50, p>0.065); also, no significant difference was observed in the double marking (U = 54.0, p>0.083) during the reproductive season, although it was greater in the LZ compared to the ZZ pair. After commingling together for one year the established LZ couple, showed an increase in double marking (U = 16.5, p<0.001) during the mating period in comparison with the prior year. Male wolves marked with greater frequency with the leg raised in both, the double and single marking, whereas females marked more in a squatting position. It is concluded that scent marking is different in recently formed pairs in captivity, which are found in a reduced space and have not free choice to elect its mate.
Article
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Se describen 37 conductas, en un grupo de cinco lobos mexicanos (Canis lupus baileyi) en cautiverio, que se integran en cinco grandes categorías; conductas amistosas, sumisas, juego, sexualesy agonísticas (agresión-defensa). Se cuantificó ia exhibición de cada una de las conductas, desde octubre de 1985 a diciembre de 1986. Se encontró un patrón anual básico de exhibición de cuatro de las cinco categorías de conductas estudiadas, ya que se presentó un aumento significativo de la exhibición de éstas categorías de conductas durante el período reproductivo que corresponden a los meses de diciembre a febrero, la única que no presentó este patrón fue el juego, este disminuyó durante este último período. las conductas amistosas se correlacionaron fuerte y significativamente con las conductas agonísticas (r=0.92), con la sumisión (r=0.90). y con las conductas sexuales (r=0.82), estos valores sugieren la gran importancia que tienen estas conductas amistosas en las relaciones sociales del grupo, ya que las demás responden fuertemente a los cambios que se presentan a lo largo del tiempo, excepto con el juego que no se encontró correlación significativa (r=-0,16). También se observó un período de intensa actividad agonística y amistosa durante noviembre de 1986, quizá debido al cambio del macho dominante del grupo. la importancia de las conductas sociales para la cría del lobo en cautiverio es discutida. Se plantea como hipótesis que las estrategias conductuales del lobo mexicano se han modificado y son diferentes a las conocidas para los lobos de regiones más norteñas.
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If behavioral mechanisms controlling suckling have been shaped by parent-offspring conflict in the ultimate sense, then proximate behavioral determinants of conflict should occur throughout lactation, with greatest intensity in the terminal phase, and offspring should have tactics for overcoming parental resistance. We observed the weaning process in a habituated wild wolf pack (Canis lupus) on Ellesmere Island, Canada, from estimated ages 5 through 10 weeks (including a continuous record for 192 h). The following variables declined with age: percentage of suckling bouts initiated by the nurser, persistence by pups, and mean duration of suckling bouts. Variables that increased with age were interbout interval, percentage of suckling bouts terminated by the nurser, and wincing or agonistic actions of the nurser. Behavioral conflict appeared in the developmental stage (estimated age 7 - 8 weeks) during which pups could feed on opened carcasses. Countertactics by pups to obtain milk were not apparent, although the pups developed diverse tactics for obtaining and sharing meat. In this group of wolves, weaning mechanisms were a complex function of food delivery by adults, discomfort of the nursing female as pups developed, and declining persistence of pups. If there is a conflict over what is optimal for pups and for the nurser in the ultimate sense, behavioral conflict is more likely to be expressed with regard to access to meat, or as conditional tactics dependent on food availability, rather than weaning conflict being controlled by fixed rules in this species.
Article
Raised-leg urinations were performed almost exclusively by dominant male and female wolves (Canis lupus). The alpha male and female in two captive groups showed full raised-leg urinations (RLUs) throughout the year. Each alpha male and female sniffed and marked on one another's marks frequently; however, the pair that bred did more marking and more double marking. They increased their marking frequency prior to and during the breeding season, decreased it at the time of parturition, while the female was in the den, and then increased to a moderate level when the female took part in pack activities again. Because the alpha pair was the principal sender and receiver of the message within the group, it is likely that double marks function to maintain and advertise the pair bond. Because the group that did not breed also did not double mark frequently, the presence of double marking in a captive group may be diagnostic of the viability of the group as a pack.
Article
Free-ranging gray wolves (Canis lupus) generally inhabit large home ranges, yet they are housed in a variety of restricted spaces when in captivity. There is continual debate as to whether space restrictions alter a wolf's behavior. The purpose of these studies was to remotely measure and then compare the amount and frequency of activity of gray wolves housed in small, artificial enclosures vs. large, more natural enclosures. Test animals comprised three adult wolves housed in kennels and three and four wolves housed in separate natural enclosures. Kenneled wolves had 2.8 m2 of surface area per wolf, and wolves in natural enclosures had 466.6 m2 (South Pack) and 349.9 m2 (North Pack) per wolf. Wolves were fitted with radiotelemetry collars containing activity sensors. Activity data were recorded every 20 min for 57 continuous hr. The amount of activity for each wolf was calculated using areas under the curve (AUCs), and the frequency of activity was analyzed by spectral analysis. There was no difference (P ≥ 0.22) in AUCs between kenneled wolves (1.399 ± 0.214 x 105 radians) and South Pack wolves (1.564 ± 0.139 X 105 radians) or North Pack wolves (1.617 ± 0.192 x 105 radians). All three groups had similar peak spectral values at frequencies that were close to daily cycles (i.e., ω = 0.12–0.17 cycles per unit time). Peaks in coherence near the dominant spectral frequency were most significant between the natural enclosures and the least significant between the kenneled wolves and the South Pack wolves. Based on these criteria of activity and under these circumstances, enclosure size appeared to have no effect on wolf activity. However, small sample sizes and variation in the data do not make these results definitive. © 1996 Wiley-Liss, Inc.
An update on the status of the Mexican wolf recovery pro-gram. Pp. 141–146 in ECOLOGY AND CON-SERVATION OF WOLVES IN A CHANGING WORLD
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Parsons, D.R.; Nicholopolous, J.E. An update on the status of the Mexican wolf recovery pro-gram. Pp. 141–146 in ECOLOGY AND CON-SERVATION OF WOLVES IN A CHANGING WORLD. L.N. Carbyn; S.H. Fritts; D.R. Seip; eds. Edmonton, University of Alberta Press, 1995
Canis lu-pus baileyi) aspecto zootecnico en el Zoológico San Juan de Aragón. Pp. 51–56 in MEMORIAS rActivity Patterns and Social Behavior of Wolves443 PRIMER
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Ramos Magaña, X.; Mata Castro, F.; López Islas, G.; Islas Dondé, G. Lobo Méxicano (Canis lu-pus baileyi) aspecto zootecnico en el Zoológico San Juan de Aragón. Pp. 51–56 in MEMORIAS rActivity Patterns and Social Behavior of Wolves443 PRIMER SIMPOSIUM NACIONAL SOBRE EL LOBO GRIS MEXICANO. M.I. Reyna Medrano; G. López Islas; B. Manrique Nobara; eds. Mexico City, Zoológico de San Juan de Aragón, 1993
Geology of Mexico: A synopsis. Pp. 3–108 in BIOLOGICAL DIVER-SITY OF MEXICO: ORIGINS AND DISTRI-BUTION
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Ferrusquia-Villafranca, I. Geology of Mexico: A synopsis. Pp. 3–108 in BIOLOGICAL DIVER-SITY OF MEXICO: ORIGINS AND DISTRI-BUTION. T.P. Ramamoorthy; R. Bye; A. Lot; J