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Fr

a Fr: fruit; Lv: leaves; Nr: nectar; Br: bract (modified leaf in the inflorescence); Ex: exudates.

a Fr: fruit; Lv: leaves; Nr: nectar; Br: bract (modified leaf in the inflorescence); Ex: exudates.

leaves and 5% only nectar (if data collected in 1997 also included). Varecia were never observed to eat animal matter or soil, but they were seen licking exudates. Furthermore, they consumed only flower nectar, not sepals or petals. From general observations, there was a significant loss of both flowers and fruit, and even shrubs declined after the cyclone. Varecia drank water occasionally in the morning from tree holes when the temperature was very hot. Eighty-four percent of the Varecia food sources came from trees, 6% from lianas or vines, and the remaining 10% from epiphytes, shrubs, and shelf fungi.

Thirty-five percent of available food species were used as food sources on only one day during the entire study. More than half of the food sources of Varecia were eaten over fewer than 4 days during the entire study. In contrast, only three species of food sources were consumed more than one-fifth of the total number of the study days. The fruits of a nontree species, Clidemia hirta, were eaten most frequently by Varecia (80 days or 42% of the total days of study), followed by the leaves of two tree species, Polyalthia oligosperma (40%) and Cynometra cloiselii (23%).

The average percentage of each food type used was calculated for each individual using instantaneous sampling at 5-min intervals containing feeding data throughout the sample period (Redy-M n = 849, Grina-M n = 551, Boloa-F n = 449, Kolara-F n = 746, and Silvera-F n = 687). For all five focal animals, there were no consistent differences between seasons in time spent foraging or feeding. However, when considering the use of individual food species, there were consistent differences in intensity of use across focal animals from day to day and month to month. In other words, on different days of the month and different months of the year, Varecia devoted different amounts of time feeding on particular species. Overall the dietary diversity ranges from 1 to 11 species per day and 5-26 species per month. Generally, the number of species eaten was greater during the hot rainy season.

Fruit: At Manombo forest, fruit was a basic part of the Varecia diet and was eaten on nearly every sample day throughout the study. Varecia were observed eating fruit on 165/188 study days (or 88% of the total observations). In addition, a great proportion of the feeding time each day was devoted to fruit eating. When data from the five focal animals were combined, fruits constituted 75% of the amount of time spent feeding (Figure 2). Seven of the ten top foods during the total 18 months were fruits. Because of the low relative density of preferred food trees, most fruits eaten by Varecia were consumed while still unripe. Unripe fruits comprised 22.5% of all observations. Some fruit trees were depleted before fruits ripened. Although V. variegata are primarily frugivorous, they were not ripe fruit specialists, at least not at Manombo during this critical time.

Figure 2. Representation of food types in the diet of Varecia variegata editorium at Manombo forest.

fruits

m

leaves

s

nectar

mushroom

water

Figure 2. Representation of food types in the diet of Varecia variegata editorium at Manombo forest.

Seventy-eight percent of Varecia food trees consisted of berries and 22% were drupe. While Varecia fed on a wide range of fruit colors (seven different colors), most fruit consumed were green. Black-and-white ruffed lemurs at Manombo utilized fruits of all sizes as did the Varecia groups on Nosy Mangabe (Morland, 1991a).

During the study, the mean percentages of fruits in the diet of each individual were as follows: Redy-M male, 72% from 34 different species; Grina-M, 81% from 30 species; Boloa-F, 57% from 25 species; Kolara-F, 83% from 42 species; and Silvera-F, 77% from 33 species. Overall, Varecia fruit sources were obtained from 70 species of trees, shrubs, and vines.

Leaves: Combined data on all individuals revealed that 17% of Varecia's diet consisted of leaves, but only of a few tree species. A great proportion of time was spent eating the leaves of two plant species, Polyalthia oligosperma and Cynometra cloiselii. Young leaves were overwhelming preferred over mature leaves. Leaves were consumed on 45 of the study days (24% of total observations, n = 188). Only six food species were exclusively leaf sources. During the sample period, the mean percentages of folivorous material in individual diets were: Redy-M 20%, Grina-M 14%, Boloa-F 21%, Kolara-F 11%, and Silvera-F 17%.

Nectar: Although the mean average of nectivory was low overall (5%), it was an important food source from September 1999 to February 2000. Varecia did not feed on the nectar of all of the flowers available in the forest, but specialized on two species, Humbertia madagascariensis and Ravenala madagascariensis, which flowered at the end of the dry season. During one day, a single R. madagascariensis tree could be visited four to five times. It appears that R. madagascariensis trees were visited only to feed, as I never saw any individuals using those trees as resting places.

Shelf fungi: Shelf fungi comprised only a small percentage (1%) of the Varecia diet. The mean percentage of shelf fungi in the diet of each individual was as follows: 0.4% for Redy-M, 1.1% for Grina-M, 1.3 for Boloa-F, 2.1% for Kolara-F, and 1.5% for Silvera-F, and the focal animals ate only one species, Polyporus sp. (Polyporaceae), which occurred at low densities throughout the forest. Shelf fungi were eaten on 22/188 days, by both females and males.

Miscellaneous items: (bark and exudates): Bark and exudates played a very small role in the diet of Varecia. The total amount of those two items combined was insignificant (0.5%), perhaps because they were eaten opportunistically, and not by all animals. Grina-M was seen feeding on bark of Dypsis sp. in the wet and hot season. Kolara-F was observed consuming the bark of Garcinia aphanophlebia in a dry month (November 1999). Redy-M, Boloa-F, and Silvera-F were also seen feeding opportunistically on plant exudates around the pedicels in the moist, cool season (September-October 1999).

Water: Water was used especially during the hottest season, between the months of February and April of 1999. I did not observe Varecia drinking water directly from a stream, but they did drink water from holes within trees, either by leaning their heads into larger tree holes or by placing their hands into holes too small for the head to fit into, and licking the water off of their hands.

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