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McIntyre, J. Theberge, M. Theberge, D. Long-distance vocal communication exists in many group-living carnivores. Understanding its behavioral and ecological significance suffers from few quantitative studies in undisturbed, wild populations.

Посетить страницу Yellowstone National Park, Wyoming, United States, we examined seasonal changes in occurrence of wolf howls and howling replies based on more than 11, unsolicited howls given over a year period.

Howling was 5-fold most frequent in the pre-breeding and breeding seasons. Pack howls primarily, but also single howls, were most common during these seasons. Answers during these seasons were predominately interpack howls. These howling peaks correlated with elevations in estradiol, testosterone, and luteinizing hormone reported elsewhere. Following the breeding season, overall howling abruptly decreased through March and April, although howling at den sites was frequent, particularly in April and May.

Howling frequency remained low all summer, during which time answers switched abruptly and almost exclusively from этом fifa 10 download tpb pc (перепутал to intrapack. Single howls stimulated distant pack members to answer with increasing frequency as the summer progressed. Although not independent, the frequency of both total howls and interpack howling rose howlwr the fall.

We relate these seasonal changes in total howling and interpack answers largely to breeding and spacing behavior in pre-breeding and breeding seasons, and intrapack answers to pack cohesion in other seasons.

Because our results may reflect a high-density, unexploited wolf population, dowjload studies under other conditions would be useful. Most group-living canids dingoes, Canis lupus dingo ; coyotes, Canis latrans ; red wolves, Canis rufus ; gray wolves, Canis lupus ; and eastern winter wolves howler exe download, Canis lycaonhyaenids spotted hyenas, Crocuta crocutaand felids African lions, Panthera leo engage in long-distance witer communication.

Detailed studies of their undisturbed behavior are few because they all are somewhat elusive and require large study areas. Our objective was to examine seasonal changes in the frequency of occurrence of wolf howls and howling replies to understand better their behavioral and ecological significance.

By relating the seasonality of howling to other seasonal changes in wolf behaviors, we hoped to contribute insights into both the triggers and the consequences of howling. The limited literature on seasonality in winter wolves howler exe download howling is inconsistent. Nonetheless, research on wild wolves in southern Europe Gazzola et al. The former study used recordings to stimulate howling.

In Minnesota, where wunter imitations were used to elicit howls, the peak also was not during winter wolves howler exe download breeding season but in October and November Harrington and Mechwith a secondary peak in March. These studies winter wolves howler exe download few dkwnload 2 to 3 in comparatively winter wolves howler exe download populations, with only 1 or 2 years of data. Small sample sizes winter wolves howler exe download not have reflected population-level patterns due to differences in pack age, sex ratio, and social composition, or a limited number of potential breeding partners.

Wolvee broad hiwler have been advanced for the long-distance vocalizations of wolves: inter- between pack howls consisting largely of territorial advertisement and defence spacing behavior and mate attraction; and intra- within pack howls involving coordination of movements, reunion, and bonding within social units summarized by Harrington and Asa The functions of howls of both categories are aided by the potential for recognition of individual wolves via their howls Theberge and Falls ; Palacios et al.

We hypothesized a peak in interpack howling during the pre-breeding and breeding seasons, largely based on an expected influence of reproductive hormones. Coyotes Gese and Ruffdingoes Corbett and Newsome ; Corbettand some hosler of wolves Peters and Mech ; Нажмите для продолжения and Laidlaw ; Servin winter wolves howler exe download marked peaks in long-distance vocalizations in the reproductive season, and all are monestrous.

In contrast, seasonality does not occur in the whoops of spotted hyenas K. Holekamp, Michigan Перейти University, pers. Seal et al. Estradiol surges during proestrus in February then drops precipitously at the end of proestrus and early metestrus pregnancy in late February or early Downloac, then drops more slowly until May and remains low all summer.

Testosterone remains at its страница high from December through March, then it, too, drops until June and remains low all summer.

Luteinizing hormone, involved in download assassin creed 4 black flag pc full version production, follows the same pattern as testosterone. Strengthening our prediction about a pre-breeding and breeding winter wolves howler exe download in interpack howling was a corresponding high level of territory-related interpack aggressive encounters at that time. In Yellowstone National Park, Wyoming, United States, a marked peak in winter wolves howler exe download distribution of such recorded encounters occurred in those seasons Quimby et al.

The literature gave us wilves basis for predicting the frequency of intrapack howling. Intrapack howling requires separation of pack members. In an Alaskan wolf population, pack cohesion dropped throughout the summer Peterson winter wolves howler exe download al.

In Yellowstone, Metz et al. Therefore, although reasons for these seasonal differences in pack cohesion were not defined, we predicted greater intrapack howling in summer. Nonetheless, any extrapolation from other studies about pack cohesion may be uncertain. A comparative нажмите чтобы прочитать больше of winter pack cohesion among 6 widely separated North American wolf populations showed little similarity Theberge and Theberge We studied the fully protected, high-density wolf population in the northern range of Yellowstone National Park using no artificial stimuli to winter wolves howler exe download howls.

In 28, h of field observations, one of us R. McIntyre recorded place, time, and winter wolves howler exe download information on all howls he heard. Winter wolves howler exe download day without exception between 1 January downlod 31 Decemberhe began field observations at daybreak, averaging 7. Often, wolves were inactive at midday, at which time he took a break wonter included in the above hours of field timeresuming in mid to late afternoon and through the crepuscular period until dark.

No data were collected after dark. The extensive, open, shrub-steppe ecosystem of the study area provided excellent opportunities for observation. Park roads provided access, and ongoing research on other aspects of wolf ecology and behavior made relevant information available on individual identities of wolves, pack structures, locations, and movements Smith et al.

McIntyre had wolves within sight the vast majority of his hours in the field. All observations were made with a spotting scope directly from park roads or pullouts winter wolves howler exe download from winter wolves howler exe download promontories. Wolves were normally 1 to 3 km away.

If closer, McIntyre, who often was in official park uniform, controlled visitors so that disturbance did not influence wolf behavior and downlkad rules against harassment of wildlife were followed. McIntyre recorded a running commentary of all wolf behavior with a voice recorder, and later transcribed his observations.

All data used in our analyses were extracted from the transcriptions. Most howls could be assigned to specific packs based on telemetry identification or visual markings.

Thus, our analysis frequently could distinguish howling wolves as pack or non-pack members. To avoid bias, only howls heard by McIntyre were used, and not those reported to him by others. We calculated monthly averages of howls heard per h of field time.

We assume constancy in this bias across types of howls. We classified answers as given by single wolves or packs, and by pack mate s or foreigner sor unknown. All howls or a sequence of howls were given a score of 1.

Winter wolves howler exe download also noted if howls in spring and summer were at den sites or rendezvous sites. Such sites were under direct observation at distances that did not disturb the wolves. Rendezvous sites were under-ranked because of our occasional uncertainty. We noted if здесь initiated pack howls or howled alone.

We examined departure from randomness in the pattern of seasonal howling jowler a 1-way analysis of variance ANOVA in which howls per field hours was the response variable and months the explanatory variable. The years were http://replace.me/3378.txt in the initial models, but had insignificant effects on the average howls per field hours and hence were excluded from the final model.

We used R for all statistical tests. We also examined departures from randomness in 7 paired comparisons in the yearly patterns of various subcategories of howling i.

To test for lg p705 pc download between and within these subcategories, we used 2-way ANOVAs with howls per field hours as the response variable winter wolves howler exe download both months and subcategories of howls as explanatory variables.

Again, the years were winter wolves howler exe download in the initial models, but had insignificant effects on the average howls per field hours and hence were excluded from the final models.

We used Shapiro—Wilk tests and residual winter wolves howler exe download to assure no model assumptions were violated. For simplicity, we refer to these 3 F -values as F1F2and F3respectively. The research was conducted under annual winter wolves howler exe download permits issued by the U. National Park Service, and met guidelines for the use of wild mammals in research approved by the American Society of Mammalogists Sikes et al. Of 11, howls, 5, were pack посетить страницу, 4, were single howls, and 1, were uncertain; 2, howls were answers.

As a year average, dowwnload was heard every 2. Seasonal variation in howling was considerable, following a consistently similar pattern in frequency of occurrence each year Fig.

The monthly average number of howls per h for the pre-breeding season December and Januaryand the breeding season February peaked at 69 and 72, respectively. Howling declined sharply through the post-breeding season Marchdenning Apriland when pups were at dens May. Yearly consistency in the monthly pattern of wolf Canis lupus howls on the northern range of Yellowstone National Park, Wyoming, over 10 years to Over the 10 years of the study, only 2 major inconsistencies broke this pattern.

Howls in April were abnormally frequent, represented by the highest data point shown for that month Wolvee. In that month, a trespassing pack invaded the den site of a resident pack and exhibited many days of aggressive behavior, preventing the resident pack from bringing food to the den site.

The pups eventually starved Smith et al. The other inconsistency was a low number of howls noted in September, November, and Decembershown as the lowest point for those months Fig. During this period, some packs dissolved and some new packs howleer to take their places.

Uncertainty winrer the population over trespassing wolves or pack affiliations may have influenced frequency of howling. Removal of ссылка на подробности 4 outlier points reduces the overall monthly variability but the overall pattern is obvious and statistically significant even with these points removed Fig.

Total howls consisted of both pack howls and single howls, which followed similar seasonal patterns of frequency of occurrence Fig. Pack howls were more frequent than single howls in November through January and again in March. A large increase occurred in single howls in December, January, and February. Removing answering howls did not affect the patterns, and patterns for the 2 categories still differed significantly.

Monthly variation in pack versus single wolf Canis lupus howls on the northern range of Yellowstone National Park, Wyoming, to

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