Evidence for territoriality is usually correlative or post hoc as we observe the results of past selection that are challenging to detect. Wolves (Canis lupus) are considered territorial because of competition for food (resource defense), yet they exhibit classic intrinsic behaviors of social regulation (protection against infanticide). This emphasis on prey and infrequent opportunity to observe wild wolf behavior has led to little investigation into the causes of or competitive underpinnings in the evolution of wolf territoriality. We report 6 cases of territorial wolf packs attacking neighboring packs at or near their den; 2 attacks were observed in detail. In all cases, except perhaps one, the attacking pack killed adult wolves either at the den or near it; in 4 cases, pups were probably lost. Loss of pups led to future loss of territory and in one case pack cessation. Intraspecific killing (measured in collared adults only) peaked in April, the month when pups were born and helpless in dens, even though aggressive interactions were at their seasonal low. Twelve of 13 (92%) of the wolves killed during the denning season (March, April, May) were reproductive (males and females), and 8 of 12 were dominant individuals (highest ranking wolf for that sex in the pack). Wolf—wolf killings were also high in October and December, the beginning and middle of the nomadic season, respectively. Aggressive interactions were more frequent during the nomadic season when wolves were roaming their territory as a group compared to the denning season when wolf activity was centered on the den and pack members less cohesive. We conclude that attacks on dens are a more effective form of interpack competition than interference during the breeding season, the current best-supported hypothesis, and that protected pup-rearing space is the primary cause of wolf territoriality.
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Vol. 96 • No. 6