Ryan.qxd Activity patterns of African buffalo Syncerus caffer in the Lower Sabie Region, Kruger National Park, South Africa S.J. RYAN and W. JORDAAN Ryan, S.J. and W. Jordaan. 2005. Activity patterns of African buffalo Syncerus caffer in the Lower Sabie Region, Kruger National Park, South Africa. Koedoe 48(2): 117–124. Pretoria. ISSN 0075-6458. The activity budgets of three herds of African buffalo in the Lower Sabie region of Kruger National Park, recorded between 1991 and 1992 were quantified to examine both nocturnal and seasonal effects on feeding activity and 24-hr movement in an area of dense bushveld. We found that the average 24-hr distance traveled by herds (3.35 km) was shorter than that found in other studies and that there was no seasonal effect on this distance, which we attribute to ready availability of water in both seasons. We found that the buffalo spent a similar amount of time feeding (9.5 hrs) as in other studies, but that the proportional feeding and resting time was influenced by the time of day and the season. These herds spent proportionally more time feeding at night (44.5 % vs 32.0 %) and more time resting in the day (28.4 % vs 16.0 %). In addition, they appeared to rest more during the day and feed more at night during the wet season, when it was hotter. Keywords: African buffalo, Syncerus caffer, activity budgets, Kruger National Park S.J. Ryan, Department of Environmental Science, Policy & Management, 137 Mulford Hall, University of California at Berkeley, Berkeley, CA 94720, USA (sjryan@nature.berkeley.edu) (Present address: Museum of Vertebrate Zoology, 3101 Valley Life Sciences Building, University of California at Berkeley, Berkeley, CA 94720, USA); W. Jordaan, 3 Conservation Services, SANParks, Private Bag X402, Skukuza, 1350 Republic of South Africa. ISSN 0075-6458 117 Koedoe 48/2 (2005) Introduction Mammalian herbivores spend the majority of their time feeding; as Beekman & Prins (1989) pointed out, horses spend more than 70 % of their time grazing and cattle, 50–60 %. Ruminant ungulates must addition- ally allocate time to ruminating and resting. In savanna ecosystems, ruminants must not only obtain sufficient quality and quantity of food, but may also need to adjust their activ- ity budget as seasonal changes constrain when the activity can take place in a 24-hour period. African buffalo Syncerus caffer are large grazers, whose feeding ecology predicts greater selection of high biomass (Owen- Smith 1988; Redfern et al. 2003) than high quality grass swards. Additionally, Redfern et al. (2003) have shown empirically that buffalo select for areas close to water sources in Kruger National Park. Thus the feeding and ranging behavior of buffalo is subject to constraints such as the abundance of grass and the availability of water. In previous studies, seasonal changes in vegetation qual- ity and water availability have been shown to alter both ranging and feeding habits of buf- falo (Funston et al. 1994). In this study, we describe activities and behaviors of three breeding herds of buffalo in a semi-arid wooded savanna environment. This study contrasts with prior studies of buf- falo which tend to focus on diurnal behaviour in open savanna habitats (Funston et al. 1994; Prins 1996; Sinclair 1977; Winterbach & Bothma 1998), as herds were tracked in relatively dense bushveld. Moreover, this study comprises 24-hour samples, meaning that nocturnal behavior of buffalo was also quantified. In addition, this data was collect- ed during a severe drought period, such that there was less rainfall occurring than in prior studies at similar latitude (Funston et al. 1994). Buffalo groups were followed in the Lower Sabie region of Kruger National Park, South Africa. Tracking data from radio-collared individuals and visual observations were used to describe daily ranging activity, and scan sampling was used to describe daily herd activity between 1991 and 1992. Differ- ences in activity budgets, timing of activity during the day and ranging observed in buf- falo are hypothesised to arise due to season- al and habitat differences (Funston et al. 1994). We explore both diurnal and noctur- nal activities, seasonal differences and com- pare this study with others in different habi- tats. Study Site The data used in this study were collected in the Lower Sabie region of Kruger National Park (Fig. 1). The study area spans two eco- zones: Knobthorn/Marula Savanna and Lebombo Bushveld (Gertenbach 1983). The data was collected in an area bordered at (31.90°, -25.11°)–(32.00°, -25.35°); all data was recorded in the WGS84 datum. This region receives 400–500 mm of rain a year on average, and the rainfall and temperature follow a unimodal distribution, which leads to a warm wet season (Oct–Mar) and a cool dry season (Apr–Sep) (Gertenbach 1980). Methods Herd Activity Data Data was collected on three herds that were tracked using radio telemetry and visual observations. The herds were estimated at approximately 400, 200 and 250 animals, respectively. Fifteen separate focal samples of activity were taken between March 1991 and August 1992. Seven of these samples were ≥24 hrs in duration and eight were ≥11 hrs of day- time observations. Main herd activity was sampled at 5-minute intervals during the day and night, where possible. Herd activities recorded were: 1. Feeding - stand/walk and eat 2. Standing - standing with head up and ruminating 3. Resting - standing or lying with head bent down- wards, not eating 4. Moving - walking but not grazing 5. Drinking - drinking water or standing < 10 m from water. Koedoe 48/2 (2005) 118 ISSN 0075-6458 Fig. 1. The Lower Sabie region of Kruger National Park. Detail shows locations of observations (asterisks) on a coarse (5 km) grid; the ecozones and three perennial river courses are also shown. Herd Movement Data Using radio telemetry, locations of herds during 40 separate days were collected over the study period (dry season, n = 24 days; wet season, n = 16 days). These locations were used to calculate the distance traveled by the herd during each day’s sample. Due to sampling time differences (range: 13–29 hrs), the distance estimate from each data collection was mul- tiplied by 24÷(observation time (hrs)) to estimate a 24-hr distance. As this is simply a difference in loca- tion over an elapsed time, this represents a minimum distance estimate. We pooled all herds and types in the study to obtain an average daily movement esti- mate. These estimates were compared by season using a Welch modified t-test for unequal variances. Herd Activity To examine diurnal activity of buffalo herds, we cropped the 24-hr samples to 12 hrs and analysed all 15 samples together. Due to unequal observation times, we converted hours of activity into propor- tions of the total. An ANOVA with post-hoc means tests (Tukey-Kramer’s Honest Significant Differ- ence Test, α = 0.05, Table 1) was used to determine both the order of times allocated to each activity and which activity occupied significantly more of the time than others. We also used the 24-hr sample data to quantify the mean time spent by the herd in each activity during the full day (Table 2). We used the longer activity budgets to compare diur- nal and nocturnal activity budgets of buffalo. Due to differences in the hour of sunset in the datasets, we chose 19:00 (7 pm) as the division in the dataset. Welch modified ANOVAs for unequal variances were used to compare means. Significant results were further tested to examine whether differential activity might occur as a result of seasonality. Activity timing was quantified by pooling all herd activity budgets. The proportion of observations within each 5-minute scan corresponding to activi- ties 1–5 were plotted against time (Fig. 4). Activity periods were defined for activities 1–3 as periods of time when the proportion exceeded 20 %. For activ- ities 4 and 5, the activity periods were both of short duration and infrequent in occurrence, such that a single herd engaging in the activity might comprise the entire occurrence. Furthermore, this might appear as five separate observations within a few hours and thus appear to be a repeated low propor- tion activity. We took clusters of activity observa- tions, rather than the proportion to represent activity periods and descriptively reported the timing of these activities. Results Herd Movement Data The mean 24 hr estimated minimum distance traveled by all herds pooled was 3.35 ± 0.35 km (x ± SE). A Welch modified t-test did not detect a significant seasonal difference in ISSN 0075-6458 119 Koedoe 48/2 (2005) Table 1 An ANOVA and Tukey-Kramer’s Honest Significant Difference (HSD) post-hoc means test of the diur- nal activity budgets of buffalo herds; the percent of observation time in each sample with the standard error (SE) and the conversion of these means and errors into hours are included Activity HSD* % Observation Approx time (SE) hours (SE) Feeding A 37.47 (2.62) 4.50 (0.31) Standing B 28.78 (2.14) 3.45 (0.26) Resting B 25.86 (2.71) 3.10 (0.33) Moving C 5.22 (1.56) 0.63 (0.19) Drinking C 2.66 (0.64) 0.32 (0.08) *Activities with different letters have significantly different means ANOVA: F = 54.07, df = 4, p = 0.0001 Table 2 An ANOVA and Tukey-Kramer’s Honest Significant Difference (HSD) post-hoc means test of the 24-hr activity budgets of buffalo herds; the percent of observation time in each sample with the standard error (SE) and the conversion of these means and errors into hours are included Activity HSD* % Observation Approx time (SE) hours (SE) Feeding A 39.41 (2.21) 9.46 (0.53) Standing B 31.49 (1.69) 7.56 (0.40) Resting C 21.45 (2.36) 5.15 (0.57) Moving D 4.76 (0.90) 1.14 (0.22) Drinking D 2.88 (0.51) 0.69 (0.12) *Activities with different letters have significantly different means ANOVA: F = 90.21, df = 4, p = 0.0001 the 24-hr distance estimates (Wet: 3.14km, Dry: 3.49 km; t = 0.4547, df = 26.022, p = 0.6531). Herd Activity Analysis of the diurnal activity of all herds showed that significantly more time was devoted to activity 1, i.e., feeding, than other activities. The herds spent an average of 37.5 % or 4.5 hrs of the day actively feeding, and another 28.8 % (approx. 3.5 hrs) to standing and ruminating; and resting took another 25.9 % (approx. 3.1 hrs) of the day. This shows us that 75 % or 9 hrs of the day is devoted to obtaining and ruminating food (Table 1). Using the seven 24-hr samples, we found a similar breakdown of activity pat- tern, but the relative proportion of time spent feeding (1) and standing (2) increased and the other activities showed a corresponding decrease in proportions (see Table 2). When we compared diurnal and nocturnal components of the herd activity budget, we found that the buffalo spent a significantly greater proportion of time feeding (#1) dur- ing the night [44.5 % vs 32.0 % (F = 9.85, df = 1, p = 0.009)]; and a trend towards a greater proportion of time resting (#3) during the day [28.4 % vs 16.0 % (F = 3.61, df = 1, p = 0.08)]. We found no significant propor- tional differences in diurnal and nocturnal time budget for other activities (Fig. 2). Due to low sample size, we could not detect sea- sonal effects on overall activity budgets. However, in Fig. 3 there is a trend suggesting that buffalo spend more time feeding during the dry season than the wet, more time mov- ing around and less time resting. Although the sample size was too small for rigorous testing, we sought to find seasonal differ- ences in diurnal and nocturnal behaviour. The difference between day and night feed- ing and resting activities was not statistically significant, although the data suggest a trend toward a greater proportion of nocturnal feeding and resting in the wet season (Feed- ing (#1), Dry Season: F = 2.61, p = 0.16; Wet Season: F = 8.48, p = 0.07; Resting (#3), Dry Season F = 0.21, p = 0.67; Wet Season: Koedoe 48/2 (2005) 120 ISSN 0075-6458 Fig. 2. Relative proportions of time allocated by the buffalo herds to activities, diurnal (white) and noc- turnal (black) components, with their standard errors (SE). Fig. 3. Relative proportions of time allocated by the buffalo herds to activities, dry season (white) and wet season (black) components, with their standard errors (SE). F = 6.31, p = 0.07). All other potential sea- sonal effects were not statistically signifi- cant. Figure 4 shows the relative proportions of each activity in each 5-minute interval in the upper panel; in the lower panel we show the corresponding number of observations in each 5-minute interval. Due to differences in the start and end time of the diurnal obser- vation sessions, the number of observations within the 5-minute time intervals increase in the early morning and decrease towards evening. The nocturnal observation ses- ISSN 0075-6458 121 Koedoe 48/2 (2005) Fi g. 4 . R el at iv e pr op or tio ns o f ob se rv at io ns o f ea ch a ct iv ity ( se e m et ho ds ) in e ac h 5- m in ut e in te rv al ( up pe r pa ne l) a nd th e co rr es po nd in g nu m - be r of o bs er va tio ns in e ac h 5- m in ut e in te rv al ( lo w er p an el ). T he 2 4- ho ur c yc le r un s fr om 7 am -7 am ; w e m ar k 7p m b ot h as a r ef er en ce f or th e on se t o f no ct ur na l a ct iv ity a nd a s a no te o f th e re du ct io n in n um be r of o bs er va tio ns . sions, although fewer in number, have a con- sistent number of observations per scan. The 24-hour cycle runs from 7am-7am; we marked 7 pm both as an indicator of the onset of nocturnal activity and of the reduc- tion in number of observations. We found that the buffalo had two defined feeding (#1) periods during the day and into the night (approximately 5:30-10:30; peak: 9:15 and 15:00-21:00; peak: 17:10), and less defined but moderate continuous feeding during the night hours and into the dawn. We see that standing (#2), contributes a large proportion of the observations during and following the feeding bouts (8:15-13:50 and 18:00-24:00), and as feeding does, it continues at moderate levels throughout. Resting (#3) has a clear daytime peak, with a major bout occurring from 12:00-16:00, and another moderate bout occurring in the early morning (3:00- 6:00). Moving (#4) appears to occur in two clusters; one in the mid-morning to early afternoon between 10:00 and 13:00 and another cluster in the evening to night at 19:00-22:00; drinking (#5) occurs in an early morning cluster around 6:00-8:00 and anoth- er mid-morning cluster from 10:00-12:00; a couple of observations occur later in the day also and two toward the middle of the night. Discussion In this study, the estimated minimum dis- tance that the buffalo herds traveled in 24 hrs was 3–4 km. This did not change between seasons, suggesting that these buffalo have high site fidelity, perhaps due to a combina- tion of the constraint of artificial water avail- ability and the presence of reliable grazing resources. In Cameroon, Stark (1986) found that buffa- lo moved around 7 km in the wet season and 5.6 km in the dry; Grimsdell & Field (1976) reported 9.6 km on average in Rwenzori National Park, Uganda; Conybeare (1980) reported 6.1 km on average in the dry season in Sengwa Wildlife Research Area, Zimbab- we. Funston et al. (1994) found that in the early wet season the herds in the Sabie Sand reserve (on the western edge of Kruger National Park) tended to remain in an area around a watering hole for up to eight days and then move. They noted also that in the dry season, the herd would make long dis- tance (10 km) movements at night in search of better grazing. The present study took place in a more closed habitat than a majori- ty of these prior studies, and the high densi- ty of artificial water availability in KNP (Brits et al. 2002) might reduce the need for wide ranging in this habitat. Moreover, there are three perennial river courses running through this study area (see Fig. 1); and buf- falo herds tend to maintain smaller home ranges where there are perrennial rivers Hunter (1996). As this study was conducted during a drought period, these river courses may have dried out and served as areas retaining green grass swards, a reliable source of grazing, but not providing suffi- cient drinking areas. Thus, the buffalo may have been constrained by artificial water presence instead and behaved similarly to the study animals of Funston et al. (1994). We cannot compare our measure directly with the daily tracking data of prior studies, as it is not representative of a full grazing path route, rather the shortest distance between radio tracking locations. However, it provides us with a range approximation for the area. We found that feeding comprised around 9.5 hrs (39.41 %) on average of the 24-hr activity budget (Table 2). This falls within the range for grazing found by both Grims- dell & Field (1976) (9 hrs) and Sinclair’s (1977) bachelor bull study in the Serengeti (9.7-10.1 hrs). This compares with the 40 % estimated by Winterbach & Bothma (1998). We found that across both seasons, the buf- falo herds appeared to spend a greater pro- portion of time feeding at night than during the day, similar to the buffalo at Lake Man- yara, Tanzania (Beekman et al. 1989) and as seen by Sinclair (1977) in the Serengeti, although his observations occurred in the wet season, not the dry. Stark (1986) esti- mated that during dry season, his study herd of savanna buffalo (Syncerus caffer brachyceros) grazed for an average of 3.1 hrs (~26 %) in a 12-hr period. This is Koedoe 48/2 (2005) 122 ISSN 0075-6458 considerably less than the average for our study herds. In addition, as suggested by Sin- clair (1977), we saw a trend toward more time spent feeding in the dry season than the wet, perhaps in response to lower vegetation quality and quantity. We also saw a trend toward more time moving in the dry season, which may reflect searching for feeding patches; concomitant with this was less rest- ing, perhaps a repercussion of spending more time looking for food. We saw that the buffalo tended to spend time feeding and standing in succession (Fig. 4), with standing and ruminating occupying around 30 % of observed activity time, both diurnally and nocturnally (Tables 1 & 2). While this provided time for rumination, it is likely a combination of both rumination and vigilance behaviour. Buffalo exhibit stand- ing, raised head vigilance; it is plausible that they are not only using visual cues, but also olfactory. We see that this activity occurs in greater proportion at night (Fig. 2), although this is not statistically significant. We posit that with a reduction in visibility at night more vigilance will be necessary. As this data was collected in non-exclusive cate- gories, we can assume that the proportion of time spent being vigilant is higher than sim- ply the time allocated to the standing (#2) category. Buffalo were recorded as resting (#3) while standing, if they were not feeding. This suggests that vigilant behaviour may comprise a significant proportion of buffalo activity. We found that the buffalo grazed in two major daytime periods, which is consistent with the findings of Winterbach et al. (1998). We also found that they continued to graze moderately throughout the night. We saw that buffalo tended to rest more during the day than at night, particularly during the warm wet season. As found in other studies (Stark 1986; Winterbach et al. 1998), the herds had a long mid-day rest period, after the morning grazing peak as shown in Fig. 4. Sinclair (1977) showed that grazing activity ceased completely in buffalo at a certain ele- vated ambient temperature. Thus we would expect the buffalo to cease grazing and to rest when the temperature is hottest—mid- day in the warm wet season. We found that the buffalo herds had two main periods of diurnal drinking, which occurred in the early morning and mid- morning, after a grazing bout and while ruminating. There appears to be more time devoted to drinking during the day than at night, but we found no evidence to suggest that they spent more time drinking during the dry season than the wet. Previous studies have suggested that buffalo only need to drink once a day (e.g. Sinclair 1977). Win- terbach & Bothma (1998) found that buffalo drank in the early afternoon, whereas Grims- dell & Field (1976) found that they drank in the mid-morning. Since we did not distin- guish between drinking and standing near water, it is not clear how often these herds drank each day. This system has a high den- sity of available water sources, so buffalo may drink more than once in a day. Qualita- tive observations (WJ) suggest that buffalo will stand in the water, but only a few herd members will drink at one time. Future Directions This data represents only 15 sampling peri- ods in total, which was not sufficient to demonstrate seasonal differences in buffalo herd activity budgets, but provided a base- line to inform future data collection. In future studies, we would suggest additional observation sessions, spread more evenly across a year to encompass both seasons appropriately. This study presented an opportunity to refine activity categories; in particular, the overlap in potentially impor- tant behaviours between categories, such as vigilance, can inform category definitions. The categories of activity such as standing and resting were both likely during ongoing rumination, but this distinction made it hard- er to compare with prior studies. In the future, the addition of samples across the year will provide more detailed comparisons and will allow further exploration of the flexibility of activity budgets as buffalo adjust to seasonal changes. The interest in ISSN 0075-6458 123 Koedoe 48/2 (2005) nocturnal activity suggests that additional long observation sessions such as these will provide valuable information regarding buf- falo ecology. Furthermore, with the advent of more sophisticated tracking devices, such as GPS collars, the combination of geo- graphic locations and activity observations can be better used, in conjunction, to under- stand buffalo feeding ecology. Acknowledgements Thank you to Kruger National Park’s Scientific Ser- vices for permission to carry out this study. Thanks to A.G. Vaughan for help with data formatting, and to three reviewers for comments and edits. S.J. Ryan’s research was supported by EPA-STAR fellowship FP-916382. References BEEKMAN, J.H. & H.H.T. PRINS. 1989. Feeding strategies of sedentary large herbivores in East- Africa, with emphasis on the African buffalo, Syncerus caffer. African Journal of Ecology 27: 129–147. BRITS, J., M.W. VAN ROOYEN & N. VAN ROOYEN. 2002. Ecological impact of large herbivores on the woody vegetation at selected watering points on the eastern basaltic soils in the Kruger National Park. African Journal of Ecology 40: 53–60. CONYBEARE, A. 1980. Buffalo numbers, home range and daily movement in the Sengwa Wildlife Research Area, Zimbabwe. South African Jour- nal of Wildlife Research 10: 89–93. FUNSTON, P.J., J.D. SKINNER, & H.M. DOTT. 1994. Seasonal-variation in movement patterns, home- range and habitat selection of buffalos in a semi- arid habitat. African Journal of Ecology 32: 100–114. GERTENBACH, W.P.D. 1980. Rainfall patterns in the Kruger National Park. Koedoe 23: 35–44. GERTENBACH, W.P.D. 1983. Landscapes of the Kruger National Park. Koedoe 6: 9–122. GRIMSDELL, J.J.R. & C.R. FIELD. 1976. Grazing pat- terns of buffaloes in the Rwenzori National Park, Uganda. East African Wildlife Journal 14: 339–344. HUNTER, C.G. 1996. Land uses on the Botswana/ Zimbabwe border and their effects on buffalo. South African Journal of Wildlife Research 26: 136–150. OWEN-SMITH, R.N. 1988. Megaherbivores: the influ- ence of very large body size on ecology. Cam- bridge: University Press. PRINS, H.H.T. 1996 Ecology and behaviour of the African buffalo: social inequality and decision making. New York, NY.: Chapman & Hall. REDFERN, J.V., R. GRANT, H. BIGGS, & W.M. GETZ. 2003. Surface-water constraints on herbivore foraging in the Kruger National Park, South Africa. Ecology 84: 2092–2107. SINCLAIR, A.R.E. 1977. The African buffalo: a study of resource limitation of populations. Chicago: University of Chicago Press. STARK, M.A. 1986. Daily movement, grazing activity and diet of savanna buffalo, Syncerus caffer brachyceros, in Benoue-National-Park, Cameroon. African Journal of Ecology 24: 255–262. WINTERBACH, H.E.K. & J. DU P. BOTHMA. 1998. Activity patterns of the Cape buffalo Syncerus caffer caffer in the Willem Pretorius Game Reserve, Free State. South African Journal of Wildlife Research 28: 73–81. 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