a "Total" equals the total number of focal observations in which the focal animal associated with another species. "Bouts" equals the number of polyspecific associations.

a "Total" equals the total number of focal observations in which the focal animal associated with another species. "Bouts" equals the number of polyspecific associations.

Focal animals and associates readily approached one another, especially in the wet season. Associates refer to individuals with whom a focal animal associated; they do not include the focal animal's conspecifics. Both lemurs usually associated 10-20 m apart, but during the wet season they usually associated within 10 m (Table 2). When associations occurred within 10 m, associates were closer to the focal animal than was at least one of the focal animal's group. Associates were a focal animal's nearest neighbor in 30-43% of the time that associates and focal animals were within 10 m of each other.

Within associations, both lemurs responded to one another's vocalizations. For example, lemurs usually stopped their activities in response to one another's alarm calls or barks, especially if other vocalizations did not precede the alarms. Both species exchanged at least five forms of vocalizations. Much as they did within their own groups, lemurs responded with movements (e.g., head-turns) or similar vocalizations. Neither species reacted to three types of vocalizations that occurred within associations: those offered when conspecific groups met; vocalizations associated with intragroup agonism; and "echo" or lost calls.

Both species responded to one another's alarm vocalizations both within and outside association. Alarm vocalizations were so loud that lemurs more than 500 m away responded with their own alarm vocalizations. Eight or more groups of lemurs that shared parts of the same home range responded quickly to alarm vocalizations. Such widespread reactions occurred after lemurs detected either a raptor or a fossa (Cryptoproctaferox). In one nocturnal observation, the lemurs in the middle story spotted a terrestrial fossa and vocalized before I could spot it no more than 30 m from me.

Activity, Foraging, and Feeding Preferences within Associations

Crowned lemurs and Sanford's lemurs conducted a variety of activities while associating (Figure 3). The only behaviors they never conducted together were resting, grooming, and mating. The two species never huddled together or groomed

Figure 3. Average daily minutes spent in each activity within polyspecific associations.

each other. Otherwise, most behaviors were well-coordinated with one another. For example, during the wet season play comprised most "other" behavior. Juveniles and other subadults of both species wrestled, played tag, or jump-played with one another, while adults conducted other activities.

Within associations, both species spent nearly equal time conducting similar activities. Both lemurs foraged and fed slightly more than they rested; they rested more during dry season associations. Average hourly distance traveled did not differ much within and outside associations, except during the wet season. Within wet season associations, crowned lemurs traveled significantly less distance than they did alone (A-=30.97, p <0.05). Neither species appeared to follow the other within polyspecific associations.

Although less than one third of a species' wet season feeding time occurred within associations, the presence of associations had little effect on food item preferences (Table 3). In the wet season, lemurs within associations primarily fed on ripe fruit, much as they did outside of associations. No statistically significant differences between or within species occurred during this season. Crowned lemurs fed slightly more on midsized trees than they did outside of associations; Sanford's lemurs consumed slightly more treelets than they did outside of associations (Table 4). Again, lemurs selected similar food sources within and outside associations. No statistically significant differences occurred. During the rest of the year lemurs fed too infrequently within associations for statistical analysis.

Table 3. Average daily percentage of items consumed within associations and alone dur ing the wet season"
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