A&?z. Behav., 1995, 50, 1325-1339

Communal and pack size in African wild dogs, Lycaon pictus

SCOTT CREEL*? & NANCY MARUSHA CREEL* *Selous Wild Dog Project, Frankfurt Zoological Society, TField Research Center for Ecology and Ethology, Rockefeller University

(Received 17 May 1994; initial acceptance 28 August 1994; final acceptance I4 March 1995; MS. number: ~7007)

Abstract. African wild dogs are 20-25 kg social carnivores whose major prey are ungulates ranging from 15 to 200 kg. In the Selous Game Reserve, Tanzania, wild dog pack size ranged from three to 20 adults (3-44 including yearlings and pups). Data from 905 hunts and 404 kills showed that hunting success,prey mass and the probability of multiple kills increased with number of adults. Chase distance decreasedwith number of adults. None the less, the distribution of per capita food intake across adult pack size was U-shaped, with a minimum close to the modal pack size. A similar result has been used to conclude that cooperative hunting does not favour in (Packer et al. 1990, Am. Nat., 136, l-19), and to argue that cooperative hunting is not responsible for group living in any carnivore (Car0 1994, of the Plains: Group Living in an Asocial Species). Daily per capita food intake only accounts for variation in the benefits to cooperative hunting, ignoring variation in costs. For Selouswild dogs, per capita food intake per km chased peaked close to the modal adult pack size (where per capita food intake per day was near its minimum). Thus, the energetics of cooperative hunting favour sociality in Selous wild dogs. Analyses that incorporate variation in both costs and benefits of hunting may show that cooperative hunting favours sociality in other specieswhere its influence has previously been rejected. 0 1995 TheAssociation for the Studyof AnimalBehaviour

In this paper, we present data on prey selection, carnivores. Influential early studies of carnivore hunting success, hunting effort and food intake ecology suggested that communal hunting might for African wild dogs in the Selous Game Reserve. favour sociality, either by increasing the size of We determine optimal hunting group size using prey that could be killed or by improving hunting the traditional evolutionary currency, daily per success(Schaller 1972; Kmuk 1975). For example, capita food intake. We then show that inclusion of Ngorongoro spotted hyaenas, Crocuta crocuta, hunting costs substantially affects predictions for typically hunted alone when pursuing Thomson’s optimal hunting group size. Wild dogs are well gazelles, Gazella thomsoni, but formed groups suited for a test of the effects of communal averaging 10.8 hyaenas when hunting , hunting on pack size (Packer & Ruttan 1988), burchelli (Kruuk 1972). Wolves, Canis because common prey are risky to attack (e.g. lupus, typically hunt Dal1 sheep, Ovis dalli, alone, warthogs, Phacochoerus aethiopicus; Fig. 1) or but tackle moose, Alces alces, in groups (Murie large relative to the dogs (e.g. yearling , 1944, Mech 1970). Associations between hunting Connochaetes taurinus, outweigh a wild dog by a group size and prey size are common among ratio of 6:l; Fig. 1). carnivores (Gittleman 1989), but are not univer- Communal hunting is one of the most con- sal. For example, , Panthera lea, hunting spicuous aspects of the behaviour of large social group size had no effect on species hunted or captured in (Stander Correspondence:S. Creel, RockefellerUniversity, Field 1992a). Research Center for Ecology and Ethology, Tyrrel ‘An association between hunting group size and Road, Box 38B RR2, Millbrook, NY 12545, U.S.A. prey size could result from a benefic’ial increase in (email:[email protected]). N. M. Creelis at the SelousWild Dog Project,Frankfurt ZoologicalSociety, the vulnerability of large prey to large hunting C/OSelous Conservation Programme, Box 1519, Dar es groups. An association between group size Salaam,Tanzania. and prey size, however, does not clarify whether

0003-3472/95/l11325+ 15 $12.00/O 8~)1995 The Associationfor the Study of Behaviour 1325 1326 Animal Behaviour, 50, 5

Figure 1. (a-c) Creel & Creel: Hunting group size in African wild dogs 1327

Figure 1. Sequel ice of hunting behaviour in communal hunts by African wild dogs. (a) Group travel slow1ly in search of preyf (compat ‘e gait with c). (b) Wildebeest herd in ‘pinwheel’ formation, attacked from several a Cc)/ Gait in full sptxd chase at 60 km/h (compare with a). (d) Dogs follow several lines of pursuit: should the hc ,d swiIng Ieft, the wild dc)g in the f‘oreground is positioned to intercept the wildebeest calf. (e) Several wild dogs distrac an aldult female wildeb test while : another attacks her calf. (f) Large or well-armed prey such as warthogs can only be tilled if 0 le head is restf aincd. 1328 Animal Behaviour, 50, 5 hunting large prey is a benefit or a necessity for intake: Caro 1994.) For another pack of wild dogs larger groups. Large hunting groups might also in the north of the Serengeti ecosystem, daily per take large prey because they must do so to meet capita food intake did not decreasewhen pack size their needs, regardless of costs (Caro 1994). decreased from 29 to 19 (Fuller & Kat 1990, Data on hunting success(defined as the percent- 1993). age of hunts that end in kills) measure the costs Using the currency of daily per capita food and benefits of communal hunting more directly. intake, only Etosha lions have been shown to hunt Studies relating hunting successto group size have preferentially in the predicted group size if com- had variable results. Ngorongoro spotted hyaenas munal hunting is the prime determinant of group succeededin 15% of solitary hunts of wildebeest size (Stander 1992a, b; Stander & Albon 1993). calves, but in 74% of group hunts (Knmk 1972). Although acknowledging that data are limited, Serengeti lions hunting Thomson’s gazelle suc- recent summaries of communal hunting have ceeded in 15% of solitary hunts, but in 32% of argued that communal hunting has little power to communal hunts (Schaller 1972). In Etosha explain grouping patterns in felids (Packer 1986; National Park, female lions’ hunting success Packer et al. 1990; cf. Stander 1992a) and across significantly increased across group sizes of one to social carnivores in general (Car0 1994). seven, for each of the five most common prey The above discussion has addressed only the (Stander & Albon 1993). Male Serengeti cheetahs effects of group size on the payoff to hunting, showed no clear effects of group size on hunting however, disregarding costs. For example, if indi- success for large or small prey (Car0 1994). viduals in groups of all sizes hunted sufficiently Kalahari spotted hyaenas showed no increase often to meet their needs an4 no more, daily per in hunting success across group sizes of one to capita food intake would not vary across group seven when challenging gemsbok or wildebeest, sizes. Groups of some sizesmight work harder to which comprise 68% of their prey (Mills 1985, attain this food intake, in terms of time or dis- 1990). tance travelled, but this variation in hunting effort Associations between group size and prey would not al&t the results discussed above. Such size or hunting successhave been widely used to results would be misleading. argue that communal hunting favours sociality, although these associations do not demonstrate that communal hunting has a net benefit. Sharing METHODS of prey in larger groups can lead to a decrease in daily per capita food intake even when hunting Study Area success or prey mass increases (Schaller 1972; Kruuk 1975; Packer et al. 1990; Caro 1994). The The Selous Game Reserve occupies 43 000 km2 relationship between daily per capita food intake in southern Tanzania. Our study area of 2600 km2 and group size has been measured for only three in the nbrthem sector of the reserve (7”35’S, social carnivores. In Serengeti lions, foraging 38”07’E) is a mosaic of miombo and chipya wood- group sizes did not match those that maximized land (dominated by Combretum, Brachystegial daily per capita food intake (Packer et al. 1990). Julbernardia, Pteleopsis, and Terminalia sericea), In contrast, Etosha lions typically foraged in the Terminalia spinosa woodland, thickets of Acacia group size that maximized food intake (Stander drepanolobium, riverine thickets along seasonal 1992a). For Serengeti cheetahs, changes in daily rivers, long grass (dominated by Andropogon, per capita food intake across group sizes were Sporobolus and Setaria) and wooded short equivocal (Caro 1994). For one pack of Serengeti grass. wild dogs, a -yield index was maximized at a group size of one for hunts of Thomson’s gazelles Study Population (the most common prey), and at four for hunts of wildebeest (Fanshawe & Fitzgibbon 1993). (The African wild dogs are considered endangered meat yield index was calculated by dividing prey (Ginsberg & Macdonald 1990) in part because mass between those dogs who participated in a they are rare under the best of conditions. Popu- kill, although packmates often shared the meat, lation density on our study site is one adult per so it was not equivalent to daily per capita food 25 km2, considerably above mean density in other Creel & Creel: Hunting group size in African \c,ilrl tlogs 1329 populations (Frame et al. 1979; Reich 1981; Fuller Definition of Terms et al. 1992; Maddock & Mills 1994). When year- lings and pups are considered, density reaches one Owing to variation in hunting techniques (e.g. dog per 17 km* at its annual peak following the stalking versus coursing) no single definition of a denning period of July-October (during the hunt applies to all large carnivores (Kruuk 1972; dry season). Pack size ranged from three to 20 Schaller 1972; Mills 1990; Stander 1992a). We adults (x& s~=7.7 & 0.23). Including yearlings defined a hunt as a pursuit of prey that either and pups, pack size ranged from three to 44 exceeded 50 m at a full run, ended with intense (18.3 & 0.67). The sex ratio was not significantly testing of prey at bay, or ended in a kill. In Selous, biased for pups or adults (Creel et al. 1995). Home wild dogs travel 12.3 km l 0.5 (x* SE through- range size averaged 438 f 87 km2 (N=6 restricted out) daily, and often pass near groups of potential polygons). prey that are ignored or tested briefly and with low effort. Our definition excludes low cost and Hunting Observations apparently casual interactions with prey. Hunting data came from six wild dog packs We have defined hunting group size as adult observed between November 1991 and March pack size. A priori, yearlings could plausibly be 1994 (11 pack-years of observation). Two mem- considered either dependants or hunters. Year- bers of each pack were radio-collared. All indi- lings (4.3 f 0.3 per pack in the 266 days analysed) viduals were identifiable by variations in the usually participate in hunts and sometimes patchwork of their black, tan and white coats provide obvious help, but sometimes cause obvi- (Fig. 1). We collected hunting data by direct ous hindrance. The number of yearlings was not observation during periods lasting from 1 to 14 associated with hunting success (simple regres- days. Radio-collared packs were initially located sion, t=O.66, df= 1,265, P=O.51) or with mean kill from a light plane or a hilltop. We observed 404 mass (t=0.22, P=O.83), but had a significant kills in 905 complete hunts. Of 310 observation association with increased chase distance (tx2.55, days (2210 h), we restricted our analyses to 266 P=O.O12). Together, these relationships suggest days on which we observed entire hunting periods that on average, yearlings have slight negative (Mills 1992). Following Packer et al. (1990), our effects on foraging successand should be consid- measures of hunting cost and benefit use daily ered dependents. When yearlings are considered means as data points, to remove a bias towards part of the hunting force, noise is added to the zeros that existed for some variables when analyses (i.e. r2-values drop) but the slopes and expressedper hunt. shapes of curves are not substantially altered. We observed wild dogs from a Land Rover at Although we measured hunting group size as distances of 2tl-400 m using binoculars and night the number of adults, daily mean food intake rates vision goggles. Most hunting occurred in two were estimated by dividing the mass of prey killed periods, 0500-0900 hours and 1730-1930 hours, by total pack size, including yearlings and pups. as in other populations (Kuhme 1965; Estes & Pups were weighted by a factor of 0.5 (following Goddard 1967; Fuller & Kat 1990). Probably M. G. L. Mills & M. L. Gorman, unpublished becauseSelous is wooded, wild dogs rarely hunted data; see also Packer et al. 1990). This measure at night (fewer than 10 kills, mostly during yielded an estimate of the food intake that a denning periods), although they often travelled pack’s hunting force provided for themselves and slowly on moonlit nights. During each hunt, we their dependants. We estimated mass of kills using attempted to record the habitat type, prey species, published figures (Sachs 1967; Blumenschine & prey number and herd composition, distance Caro 1986). Our analyses were based on prey mass chased (estimated using the vehicle odometer), killed (except where noted), which could be con- individuals leading the chase, individuals initiat- verted to edible mass using weighting factors, but ing the kill, characteristics of kill site, estimated similar percentages of the mass (60-80%) of most mass of prey killed, estimated mass of remains not carcasseswere edible. eaten, time on kill, and interactions with other Wild dog packs are highly cohesive, and all carnivores. In this paper we focus primarily on pack members normally moved together during relationships between group size and the costs and morning and evening hunting periods. (An excep- benefits of hunting. tion is the 2-3-month denning period, when the 1330 Animal Behaviour. 50, 5 dominant female usually does not hunt: Malcolm Snedecor & Cochran 1967; Draper & Smith & Marten 1982.) When a chase began, all pack 1981). members normally pursued and harassed prey. Social rallies almost invariably preceded hunting, and appeared to excite and coordinate the pack RESULTS for hunting (Kuhme 1965; Estes& Goddard 1967; Malcolm 1979). Excluding pack members from Mean Hunting and Foraging Success the hunting group based on behavioural data Selouswild dog packs made from 0 to 16 chases could be misleading for three reasons. (1) We per day (x* ~~~4.2f0.2, N=266 days), and often did not have all dogs continuously in view killed from 0 to 10 per day (1.8 f 0.1, through an entire hunt. (2) Multiple kills were N=266). Hunting success (kills/hunt) was 44% common, but we often detected multiple kills only (range O-100%) when calculated using only data after the hunt finished. Thus, dogs not in view at from complete days, and 45% (range t&1000/,) one kill were often pursuing another prey animal. using all observations. Estimated mass of prey (3) Participation in a hunt is diIhculr to define ranged from 0.5 to 208 kg (48.5 f 2.15 kg, operationally because the simple presence of an N=384). Feeding duration ranged from 1 to additional hunter may affect the prey’s behaviour 312 min (35.3 f 2.1 min, N=357). Chase distances or escape options (Reich 1981; Stander 1992a; see ranged from 50 m to 4.6 km (0.57 f 0.03 km, Results). N=775). Successful chases also ranged from No measures of hunting successor effort were 50 m to 4.6 km, but were generally longer affected by the within-pack adult sex ratio, which (0.84 f 0.05 km, N=304). ranged from 20 to 80% male. Packs killed 4.0 f 0.35 kg/dog/day (N=216), with a range of c37.5 kg. Clearly, a wild dog Statistical Methods cannot eat 37.5 kg in a day. Actual food consump tion averaged between 2.0 and 2.5 kg/dog/day, Data points were calculated on a daily basis based on two adjustments to the overall mass (e.g. daily mean mass of kills, daily mean chase killed. First, mass of prey was devalued to reflect distance) and variances were calculated using that 20-40% of it is usually not eaten (e.g. large daily means (N=266 days) as data points (follow- bones, stomach contents). Second, observations of ing Packer et al. 1990). We used temporal auto- feeding by wild dogs known not to have eaten for correlation to determine that successive days several days suggest that adult stomach capacity provided statistically independent points. is roughly 9 kg, so edible biomass in excess of Means are reported with standard errors 9 kg/dog was excluded. throughout. Linear regressions were fitted by ordinary least squares, using residual plots to test assumptions (Snedecor & Cochran 1967; Draper Prey Selection and Hunting Success & Smith 1981). For most regressions, overlapping In 817 hunts in which at least one prey and truncated points prevented clear display of species was identified, 17 species were hunted: scatter in the data, so 95% confidence limits on impala, Aepyceros melampus (N=293 hunts), blue linear regressions are shown as dotted lines. For wildebeest (N=266), warthog (N=88), African analyses of the relationship between foraging hare, Lepus capensis (N=32), (N=30), successand pack size, the form of the relationship duiker, Sylvicapra grimmia (N=27), Lichtenstein’s was of particular interest. Therefore, where hartebeest, Alcelaphus lichtensteini (N= 17), eland, non-linear regression models fitted significantly Taurotragus oryx, common reedbuck, Redunca better than linear models, we used polynomial arundinum, buffalo, Syncerus cafir, greater kudu, regressions. All polynomial regressions included Tragelaphus strepsiceros, bushbuck, Tragelaphus quadratic terms. No higher-order terms were sig- scriptus, sable antelope, Hippotragus niger, bush- nificant. We fitted non-linear regressions with pig, Potamochoerus porcus, waterbuck, Kobus initial values of the Marquardt parameter set so ellipsiprymnus, banded mongoose, Mungos mungo, that regression coefficients were estimated by a and yellow baboon, Papio cyanocephalus (NI 10 compromise between steepest-descentand Gauss- hunts each). In a sample of 368 identified kills, 10 Newton linearization methods (Marquardt 1963; prey species were killed: impala (N= 188 kills), Creel & Creel: Hunting group size in African wild dogs 1331

Table I. Profitability of common prey for African wild dogs in Selous Profitability Yo Mass Chase kg/km Species Hunts KillS Success (kg) (km) kg/hunt chased

Impala 293 188 64 31.9 1.19 20.4 17.1 Wildebeest 266 100 38 92.7 069 35.2 51.0 Warthog 88 31 35 33.8 0.31 11.8 38.1 African hare 32 10 31 2,o 0.13 0.6 4.8 Zebra 30 2 7 157.5 I.70 11.0 - Common duiker 21 16 59 17.6 0.53 10.4 19.6 Total 736 347 Weighed mean* 47 48.8 0.88 22.9 29.8

*Means were weighted using number of kills or chases (as appropriate) for each species. blue wildebeest (N= 100) warthog (N=31), com- Also, seasonal patterns of prey species’reproduc- mon duiker (N= 16), Lichtenstein’s hartebeest tion create asynchronous peaks in the availability (N= 15) African hare (N= IO), common reedbuck of vulnerable young (which are highly preferred (N=4), zebra (N=2), waterbuck (N= 1) and by wild dogs). More Xetailed analysis of profit, bushbuck (N= 1). prey availability and prey choice will be presented The four ungulate speciesthat were hunted but elsewhere. not killed were either much larger than the range of normal prey (eland and buffalo), had unusually dangerous horns or were uncommon in Selous Communal Hunting and Group Size (greater kudu and especially sable). Mongooses Cooperative hunting behaviour and yellow baboons were also not killed, but appeared to be hunted in . Coordination between the members of an Table I shows hunting success,chase distance pack is seen throughout a hunt and two measures of profitability (mass killed per (Fig. 1). At several stages, effectiveness appears to hunt, and per km chased) for prey specieshunted depend on the number of cooperating hunters. on more than 25 occasions. Impala were hunted Although its function for hunting is arguable, most often (40% of the total), killed most often the members of a pack almost invariably go (54% of the total) and yielded the highest hunting through an intense greeting ceremony or ‘rally’ success(64%). Zebra provided the most mass per just prior to a period of hunting. The rally appears kill, but were rarely killed, with a probability of to ensure that all pack members are awake, alert killing (7%) far lower than other species (mini- and ready to hunt simultaneously, prior to trot- mum of 31%). Excluding zebra, wildebeest were ting in search of prey (Estes & Goddard 1967; the heaviest prey killed (mean of 93 kg). African Malcolm 1979). Once on the move, pack members hares were killed with the shortest chases(mean of trot or canter together at 10 km/h, usually spread 130 m), but yielded little food (2 kg). over lo-100 m (Fig. la). Combining these relationships shows that Upon sighting prey, a pack often does not hunt. wildebeest yield the greatest food mass per hunt If the pack hunts, small prey (e.g. impala or and the greatest food mass per km chased (Table duiker) flee immediately, but large prey (e.g. I). Indeed, wildebeest were hunted three to 10 wildebeest) often stand in a defensive ‘pinwheel’, times more frequently than all prey speciesexcept facing outward, charging and using their horns to impala (Table I). Impala were hunted most fre- defend themselves (Fig. lb). Juveniles keep to the quently of all, despite ranking second in mass/ centre of the pinwheel. Well-armed prey (e.g. hunt and fourth in mass per km chased (Table I). warthog, greater kudu males) may also stand and The apparently sub-optimal preference for impala defend themselves rather than fleeing, even when is probably the result of different population solitary. When faced with a defensive formation, densities of prey species (impala are common). wild dogs encircle the herd and simultaneously 1332 Animal Behaviour, 50. 5

attack from severaldirections (Fig. I b). If the prey defend themselves well, the pack often departs after testing the prey for IO s to 5 min. The apparent goal of testing and attack is to force some or all of the herd to run, thus increasing their vulnerability. Simultaneous attacks appear to be effective because one wild dog can incite a charge, then packmates rush behind the charging prey to separate it from the herd. Once one or a few prey begin running, the entire herd often bolts, and a full-speed chase (at 40-60 km/h) ensues (Fig. Ic). L1u q u

Especially in woodland, prey do not run in a cl straight line. For example, they follow lines of 8 2oc on q nuOu q low resistance through trees, uneven ground, and “B waterholes. Prey with territories that are small q q

relative to the length of a chase (e.g. duiker) often 1 attempt to circle. Individual wild dogs pursuing a 0 4 8 12 16 20 prey animal do not all follow the same line (Fig. Adult pack size Id). Together, these patterns often result in one or Figure2. Hunting success(measured as kills/hunt) sig more wild dogs intercepting a prey animal after a nificantly increasedas the number of adult wild dam shortcut, whether intentional or not (Estes & increased.Dotted linesare 95O!confidence limits. Goddard 1967; cf. Fanshawe & Fitzgibbon 1993). Once a prey animal has been caught, pack sharp contrast to those seen in Serengeti Nationat members cooperate in pulling it to a halt, or in Park, where hyaena group sizesare larger (Hofer occupying the animal’s attention by feinting from & East 1993). There, hyaenas were present at 86Q the front, while others attack from behind and of wild dog kills (excluding gazelle fawns), and the begin disembowelling. Several dogs may attack duration that wild dogs retained their killp and distract a female while packmates attack its depended on the number of each species present dependent offspring (Fig. le). Cooperation is important in restraining the head of large (Fanshawe & Fitzgibbon 1993). In Selous, compe- tition at wild dogs’ kills also came occasionally prey (e.g. wildebeest) and dangerous prey (e.g. warthog: Fig. If). Becausekilling can take several from lions (four interactions, four kills lost;

m 2.5 =I a E

0 4 8 12 16 20 0 4 8 12 16 20 Adult pack size Adult pack size gfgare 3. Mean mass of prey killed significantly Fignre 5. Mean number of prey killed simultaneously increased as the number of adult wild dogs increased. increased significantly as the number of adult wild dogs Points are daily means. Dotted lines are 95% confidence increased. Points are daily means. Dotted lines are 95% limits. confidence limits.

The distance chased in a successfulhunt signifi- cantly decreased as group size increased (Fig. 4), 0 0 from 1.1 km in packs of three adults to 0.5 km in 0 0 Q lh 2.5 0 0 0 0 0 packs of 20 adults (b= - 33.7 f 9.37, ?=0.20, tz3.60, WO.001). The number of animals simul- Q taneously killed also increased with group size 0 0 0 (Fig. 5; bz0.02 f 0.005, r*=0.24, ~4.80, 0 00 cl0 Q E P-+001). 0 0 0 0 n In summary, larger packs were more likely to kill

0 in a given hunt, killed heavier prey with shorter chases,and killed more members of the herds they chased. None the less, larger packs made more chases per day (Fig. 6; b=0.23 f 0.05, ?=0.28, tz4.66, P

Hunting costs can be incorporated into forag- ing successin a more precise way by measuring I’ kg/dog/km chased per day, or kg/dog/km travelled 7 0 q per day. Both of these measures improve on kg/dog/hunt by incorporating variation in effort 6 on 0 q q 0 expended in a hunting bout. ‘Chasing’ refers only to focused, high-speed pursuit of prey (Fig. 1~). ‘Travelling’ refers to all movement, including both chases and slow-paced searching (Fig. la). Kilograms/dog/km chased has the advantage of narrowly focusing on effort that is unequivocally directed to hunting. Kilograms/dog/km travelled has the advantage of being a more inclusive 2 - x”O q q 00000 Cl q /’ measure of cost, but it might include costs of travel that were in fact directed to another pur- pose. In practice, wild dog packs rarely travel IIIIIIIII/, IIIII/II without hunting, and we consider all movement to 0 4 8 12 16 20 be hunting-related, although travel undoubtedly Adult pack size serves other functions simultaneously (e.g. terri- Figure 6. The number of hunts per day significantly torial defence, assessingdispersal opportunities). increasedas the number of adult wild dogs increased. As adult pack size increases, kg killed/dog/km Pointsare daily means.Dotted linesare 95%confidence travelled also increases, throughout the observed limits. range of pack sizes (Fig. 7c; Y=O.l5+0,06X -0.001X2, Fx,2M= 16.3, P

4 (b) q 0

4 8 12 16 20 0 4 8 12 16 20

Ll 0 0 4 8 12 16 20 Adult pack size Figure 7. The relationship of foraging success to pack size, shown by non-linear regressions. Points are daily means. Four measures of foraging success are shown as the dependent variable: (a) kg killed/dog/day; (b) kg killed/dog/hunt; (c) kg killed/dog/km travelled, including search; (d) kg killed/dog/km chased in full-speed pursuits. variance in foraging successis affected by group DISCUSSION size(Pulliam & Caraco 1984). For Selouswild dogs, variance in foraging successdid not correlate with Other Wild Dog Populations group size (NS for all measures of foraging success; e.g. for kg/dog/km travelled, r,=0.20, P=O.69), nor Wild dogs are generally regarded as efficient were non-linear associations apparent. Thus the hunters, and their high hunting successin Selous results of simple optimality and stochastic models (44%) parallels that recorded in other populations, are likely to coincide reasonably (as in Serengeti which ranges from 39 to 85% (Estes & Goddard lions: Packer et al. 1990). 1967; Kruuk & Turner 1967; Schaller 1972; 1336 Animal Behaviour. 50, 5

Effects of Group Size Unrelated to Hunting Our analyses of optimal group size address only the effects of communal hunting. Although our results confirm that communal hunting favours sociality in wild dogs, factors completely un- 120 related to hunting are also likely to affect pack 3 size. We do not suggest that other group-level 8 activities (e.g. group defence of territories or off- ? 90 spring: Packer et al. 1990) or patterns of related. b cl ness (Rodman 1981; Giraldeau & Gilhs 1988; Giraldeau & Caraco 1993) are unimportant. 60 Preliminary evidence suggests that there are other benefits of group living for Selous wild dogs. First, clashes between packs (which have included fatal fights) were won by the larger pack in 10 of 10 cases. In two clashes between packs of equal 0 4 8 12 16 20 size, each retreated once. Second, large groups Adult pack size produce large same-sexed cohorts, which may confer advantages in dispersal. Groups of tran- Figure 8. Frequency distribution of pack sizes in which sient females can take over existing packs by Selous wild dogs lived, based on individual-months of evicting resident females, and numbers are likely observation (following Rodman 1981). Dashed line shows normal approximation. to affect the outcome of take-over attempts, Finally, large packs may be better at defending their pups from predation. For example, a pack of Malcolm & van Lawick 1975; Fanshawe & Fitzgibbon 1993; Fuller & Kat 1993). Some of the 16 attacked an adult male lion that was stalking variation in hunting success between studies is their pups, and drove it away without casualties. Systematic data are needed to test whether these probably due to small sample sizes (the highest and lowest values reported were based on 130 and other potential benefits (and potential costs hunts). Pooling data from four wild dog studies in such as easier transmission of pathogens) are important. In addition, our data address only the the Serengeti ecosystem, hunting success was also 44% (N=666 hunts: Schaller 1972; Malcolm & energetics of hunting; selection on the risks of injury during hunting might not act in parallel. van Lawick 1975; Fanshawe & Fitzgibbon 1993; Fuller & Kat 1993). The accord with Selous is somewhat striking, given substantial differences in Hunting and Group Size the prey set available and the physical environ- ment. None the less, energetic returns might differ Analyses of optimal hunting group size are substantially between populations, owing to strongly dependent on the currency of foraging variation in prey size or hunting effort. success that is used (Fig. 7a-d). Hunting success The range of prey species hunted and killed is (kills/hunt) and daily per capita food intake (kg/ broader in Selous than has been reported for most individual/day) have been widely used in analyses other populations (e.g. Malcolm & van Lawick of optimal hunting group size in large carnivores 1975; Fuller & Kat 1990, 1993). This difference is (Mills 1985; Packer et al. 1990; Stander 1992a; probably partly due to sample size differences. All Fanshawe & Fitzgibbon 1993; Stander & Albon of the prey species killed in Selous have been 1993; Caro 1994). It is widely recognized that recorded in at least one other study (see especially hunting success will rarely be an appropriate de V. Pienaar 1969, who summarized results for currency, because it does not account for covari- 4406 carcasses eaten by wild dogs in Kruger ation between group size and mass of prey (Care National Park). Of the species hunted but not 1994). killed in Selous, three have not previously For wild dogs, daily per capita food intake is been reported (bushpig, baboon and banded also inappropriate for analysis of optimal group mongoose). size, because it fails to account for covariation Creel & Creel: Hunting group size in African wild dogs 1337

between hunting costs and pack size. Although rejection of the influence of communal hunting on kgldoglday showed an intermediate minimum sociality is premature. For stalkers, covariation when regressed on pack size (Fig. 7a), kg/dog/km between group size and the frequency of hunts chased showed an intermediate maximum (Fig. should be tested. For coursers, both the frequency 7d), and kg/dog/km travelled steadily increased and length of chases should be considered. For (Fig. 7~). Thus, both measures of foraging success both, the risks of injury should be assessed.Many that incorporate variation in costs show that large social carnivores display a range of costly communal hunting favours sociality in wild dogs. cooperative hunting behaviour patterns that logic The currency of kg/dog/km travelled suggests that suggests would not be maintained without an communal hunting provides directional selection offsetting benefit (Fig. 1; Mech 1970; Kruuk 1975; in favour of large groups, with other factors (e.g. Mills 1990; Stander 1992b). parasite loads or intra-group aggression) setting In principle, comparisons between speciescould an upper limit. The currency of kg/dog/km chased be standardized by measuring both benefits and suggeststhat communal hunting provides stabiliz- costs of hunting in kilojoules, rather than kilo- ing selection for packs slightly larger than the grams of meat or kilometres run. We did this observed modal pack size. using estimates of the energetic content of meat These results may be of widespread importance (6000 kJ/kg: Ulmer 1983), and an allometric when considering the role of communal hunting in regression of the costs of terrestrial locomotion the evolution of carnivore sociality. For example, (kJ/km= 10.7 x body mass in kg”.68: Tayler et al. lions in have been 1982; Calder 1984). Because these relationships examined in an influential series of papers on are not based on data specifia to wild dogs, the optimal group size (Schaller 1972; Caraco & Wolf results are speculative. None the less,it is interest- 1975; Rodman 1981; Packer 1986; Mange1 & ing that kJ killed/dog/kJ in chase was maximized Clark 1988; Packer et al. 1990). Packer et al. in packs of 12 adults, and kJ killed/dog/kJ in (1990) showed that kg/lion/day was minimized at travel was maximized in packs of 18. intermediate hunting group sizes.They concluded Finally, models of optimal group size often that ‘group-size-specific foraging success is assume that individuals’ fitness peaks at inter- insufficient to account for the observed distri- mediate group sizes (Rodman 1981; Giraldeau & bution of group sizes’ (page 2). Incorporating the Caraco 1993; Higashi & Yamamura 1993). As costs of hunting may alter this result (although Rannala & Brown (1994) noted, there is relatively lions in intermediate-sized hunting groups would little empirical support for this assumption. For still fail to meet their daily metabolic needs; the Selous wild dogs, the fitness effects of cooperative primary conclusion, that lions in small and hunting do peak at intermediate group sizes,using intermediate-sized prides should hunt alone, three of four logical currencies (kg killed/dog/km should therefore remain true: C. Packer, personal chased; kJ killed/dog/kJ in chase; and kJ killed/ communication). dog/kJ in travel; these currencies are collinear). Caro (1994) reviewed analyses of optimal hunt- Despite recent rejections of its importance, com- ing group size based on daily per capita foraging munal hunting may still prove to be a force in the success,and concluded: evolution of sociality and group size.

few studies report per capita foraging returns, but in the majority of those that do, per capita ACKNOWLEDGMENTS foraging success did not increase with group size. In populations in which it did, grouping We are grateful to the government of Tanzania for patterns did not reflect optimal foraging group permission to work in the Selous Game Reserve, sizes. Though limited, current evidence there- particularly the Wildlife Division of the Ministry fore suggests that cooperative hunting is not of Lands, Tourism and Natural Resources; the responsible for group living in any carnivore. Council for Science and Technology; and the [page 3421 Serengeti Wildlife Research Institute. Personal thanks to G. Bigurube, M. Borner, B. Kibonde, Until analyses of optimal hunting group size W. Minja, B. Mkosamali and J. Muhuri for their incorporate hunting costs, however, a broad logistic help, to G. Bigurube, M. Borner and 1338 Animal Behaviour, 50, 5

R. Jackson for radiotracking flights, and to F. Giraldeau, L.-A. & Gillis, D. 1988. Do lions hunt ii., Nottebohm and P. Tellerday at RUFRC. The group sizes that maximize hunters’ daily food returns7 Anim. Behav., 36, 611-613. manuscript benefited from comments by T. Caro, Gittleman, J. L. 1989. Carnivore group living: compara- F. Nottebohm, C. Packer and two anonymous tive trends. In: Carnivore Behavior, Ecology ad referees. This research was supported by the Evolution (Ed. by J. L. Gittleman), pp. 183-207. Frankfurt Zoological Society-Help for Threat- Ithaca, New York: Cornell University Press. ened Animals Project 1112/90, with additional Higashi, M. & Yamamura, N. 1993. What deternrures funding from the Scholarly Studies Program of animal group size: insider-outsider conflict and its resolution. Am. Nut., 142, 553-563. the Smithsonian Institution and an NSF-NATO Hofer, H. & East, M. L. 1993. The commuting system of postdoctoral fellowship. Serengeti spotted hyaenas: how a predator copes with migratory prey. I. Social organization. Anim. Behav,, 46,547-551. Houston, A., Clark, C. W., McNamara, J. & Mange], REFERENCES M. 1988. Dynamic models in behavioural and evol. utionary ecology. , Lond, 332, 29-34. Blumenschine, R. J. & Caro, T. M. 1986. Unit flesh Kruuk, H. 1972. The Spotted : A Study of pre. weights of some East African bovids. Afi. J. Ecol., 24, dation and Social Behavior. Chicago: University of 273-286. Chicago Press. Calder, W. A. 1984. Size, Function and Life History. Kruuk, H. 1975. Functional aspects of social hunting it, Cambridge, Massachusetts: Harvard University Press. carnivores. In: Function and Evolution in BehavioM Caraco, T. & Wolf, L. L. 1975. Ecological determinants (Ed. by G. Baerends, C. Beer & A. Manning), pp, of group sizes in foraging lions. Am. Nat., 109, 119-141. Oxford: . 343-352. Kruuk, H. & Turner, M. 1967. Comparative notes ou Caro, T. M. 1994. Cheetahs of the Serengeti Plains: predation by lion, , and wild dog in Group Living in an Asocial Species. Chicago: the Serengeti area, . Mammalia, 31, l-27. University of Chicago Press. Kuhme, W. 1965. Communal food distribution and Creel, S., Creel, N. M., Matovelo, J. A., Mtambo, M. division of labour in African hunting dogs. Nature, M. A., Batamuzi, E. K. & Cooper, J. E. 1995. The Land., 205,4434. effects of anthrax on endangered African wild dogs Maddock, A. H. & Mills, M. G. L. 1994. Population (Lycaon pictus). J. Zool., Lond., 236, 199-209. characteristics of African wild dogs Lycaon pictus in Draper, N. & Smith, H. 1981. Applied Regression the eastern Transvaal lowveld, , as Analysis. 2nd edn. New York: John Wiley. revealed through photographic records. Biol. Con- Estes, R. D. & Goddard, J. 1967. Prey selection and serv., 67, 57-62. hunting behavior of the African wild dog. J. Wildl. Malcolm, J. R. 1979. Social organization and the com- Mgmt, 31, 52-70. munal rearing of pups in African wild dogs (Lycuozz Fanshawe, J. H. & Fitzgibbon, C. D. 1993. Factors pictus). Ph.D. thesis, Harvard University. influencing the hunting success of an African wild dog Malcolm, J. R. & van Lawick, H. 1975. Notes on wild pack. Anim. Behav., 45,47!%490. dogs (Lycaon pictus) hunting zebras. Mammalia, 39, Frame, L. H., Malcolm, J. R., Frame, G. W. & van 231-240. Lawick, H. 1979. Social organization of African wild Malcolm, J. R. & Marten, K. 1982. Natural selection dogs Lycaon p’ictus on the Serengeti plains, Tanzania, and the communal rearing of pups in African wild 1967-1978. Z. Tierpsychol., 50, 225-249. dogs (Lycaon pictus). Behav. Ecoi., 10, I-13. Fuller, T. K. & Kat, P. W. 1990. Movements, activities, Mangel, M. & Clark, C. W. 1988. Dynamic Modeling in and prey relationships of African wild dogs (Lycaon Behavioral Ecology. Princeton, New Jersey: Princeton pictus) near Aitong, southwestern . Afr. J. University Press. Ecol., 28, 330-350. Fuller, T. K. & Kat, P. W. 1993. Hunting success of Marquardt, D. W. 1963. An algorithm for the least African wild dogs in southwestern Kenya. J. squares estimation of nonlinear parameters. J. Sot. ., 74,464467. ind. appl. Math., 2, 431441. Fuller, T. K., Kat, P. W., Bulger, J. B., Maddock, A. H., Mech, L. D. 1970. The Wolf The Ecology and Behavior Ginsberg, J. R., Burrows, R., McNutt, J. W. & Mills, of an . Minneapolis: University of M. G. L. 1992. Population dynamics of African wild Minnesota Press. dogs. Wildlife 2001: Populations (Ed. by D. R. Mills, M. G. L. 1985. Related spotted hyaenas forage McCullough & R. H. Barrett), pp. 1125-1139. together but do not cooperate in rearing young. London: Elsevier Applied Science. Nature, Lond., 316, 61-62. Ginsberg, J. R. & Macdonald, D. W. 1990. Foxes, Mills, M. G. L. 1990. Kalahari Hyaenas: The Compara- Wolves, Jackals and Dogs: an Action Plan for the tive Behavioural Ecology of Two Species. London: Conservation of Canids. Gland, Switzerland: IUCN. Unwin Hyman. Giraldeau, L.-A. & Caraco, T. 1993. Genetic relatedness Mills, M. G. L. 1992. A comparison of methods used to and group size in an aggregation economy. Evol. study food habits of large African carnivores. In: Ecol., 7, 429438. Wildlife 2001: Populations (Ed. by D. R. McCullough Creel & Creel: Hunting group size in African wild dogs 1339

& R. H. Barrett), pp. 1112 1124. London: Elsevier Rodman, P. S. 1981. Inclusive fitness and group size Applied Science. with a reconsideration of group sizes in lions and Murie, A. 1944. The Wolves o/’ Mounr McKinley. wolves. Am. NM., 118, 275283. Washington, D.C.: United States Government Print- Sachs, R. 1967. Liveweights and body measurements of ing Office. Reprint 1971. Serengeti game animals. E. Afr. Wildl. J., 5, 24-36. packer, C. 1986. The ecology of sociality in felids. In: Schaller, G. B. 1972. The Serenngcti Lion. Chicago: Ecological Aspects of Social Evolution (Ed. by D. 1. University of Chicago Press. Rubenstein & R. W. Wrangham), pp. 429451. Snedecor, G. W. & Cochran, W. G. 1967. Statisrical Princeton, New Jersey: Princeton University Press. Methoak. 2nd edn. Ames, Iowa: Iowa State University Packer, C. & Ruttan, L. 1988. The evolution of Press. cooperative hunting. Am. Not.. 132, 159-198. Stander, P. E. 1992a. Foraging dynamics of lions in a .Packer. -~ C.. Scheel. D. & Pusev. A. E. 1990. Whv semi-arid environment. Can. J. Zoo/., 70, 8-21. lions form groups: food is not enough. Am. Nut., 138, Stander, P. E. 1992b. Cooperative hunting in lions: the I-19. role of the individual. Behav. Ecol. Sociohiol., 29, de V. Pienaar, U. 1969. Predator prey relationships 445454. amongst the larger of the . Koedae, 12, 108-176. Stander, P. E. & Albon, S. D. 1993. Hunting success of pulliam, H. R. & Caraco, T. 1984. Living in groups: is lions in a semi-arid environment. Symp. zool. Sot. there an optimal group size? In: Behavioural Ecolagy Lund., 65, 127-143. (Ed. by J. R. Krebs & N. B. Davies), pp. 122 147. Taylor, C. R., Heglund, N. C. & Maloiy, G. M. 0. 1982. Sunderland, Massachusetts: Sinauer. Energetics and mechanics of terrestrial locomotion. I. Rannala, B. H. & Brown, C. R. 1994. Relatedness and Metabolic energy consumption as a function of speed conflict over optimal group size. 7iendy Ecol. Evol., 9, and body size in birds and mammals. J. esp. Bial.. 97, 117-119. I 21. Reich, A. 1981. The behavior and ecology of the African Ulmer, H. V. 1983. Nutrition. In: Human Physiology wild dog (Lycaon picrus) in the Kruger National Park. (Ed. by R. F. Schmidt & G. Thews). pp. 573-586. Ph.D. thesis, Yale Ilniversity. Berlin: Springer-Verlag.