AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 141:646650 (2010)

Brief Communication: Reaction to Fire by Savanna Chimpanzees (Pan troglodytes verus) at Fongoli, Senegal: Conceptualization of Fire Behavior and the Case for a Chimpanzee Model
Jill D. Pruetz1* and Thomas C. LaDuke2
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Department of Anthropology, Iowa State University, Ames, IA 50010 Department of Biological Sciences, East Stroudsburg University, East Stroudsburg, PA 18301 KEY WORDS hominid; control of re; apes environment at Fongoli, similar to the savanna mosaic thought to characterize much of hominid evolution, makes these apes ecologically important as a living primate model as well. Chimpanzees at Fongoli calmly monitor wildres and change their behavior in anticipation of the res movement. The ability to conceptualize the behavior of re may be a synapomorphic trait characterizing the human-chimpanzee clade. If the cognitive underpinnings of re conceptualization are a primitive hominid trait, hypotheses concerning the origins of the control and use of re may need revision. We argue that our ndings exemplify the importance of using living chimpanzees as models for better understanding human evolution despite recently published suggestions to the contrary. Am J Phys Anthropol 141:646650, 2010. V 2009 Wiley-Liss, Inc.
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ABSTRACT The use and control of re are uniquely human traits thought to have come about fairly late in the evolution of our lineage, and they are hypothesized to correlate with an increase in intellectual complexity. Given the relatively sophisticated cognitive abilities yet small brain size of living apes compared to humans and even early hominins, observations of wild chimpanzees reactions to naturally occurring re can help inform hypotheses about the likely responses of early hominins to re. We use data on the behavior of savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal during two encounters with wildres to illuminate the similarities between great apes and humans regarding their reaction to re. Chimpanzees close relatedness to our lineage makes them phylogenetically relevant to the study of hominid evolution, and the open, hot and dry

Of the many ways in which humans are considered unique, the use and control of re are behavioral characteristics that are thought to have come about fairly late in the evolution of our lineage. Although data from certain sites suggest the ability to control re extends beyond one million years ago (Gowlett et al., 1981; Brain and Sillen, 1988; Bellomo, 1994), the issue is far from resolved. It was long held that Homo erectus used and controlled re during the Middle Paleolithic, based primarily on data from Zhoukoudian (Black, 1931; James, 1989; Shen et al., 2004). Recently, however, the evidence has come under criticism (Weiner et al., 1998; Pickering et al., 2008). Data indicate that late Lower Paleolithic Homo may have used re (Karkanas et al., 2007), but in general the issue is still debated. The use of re is thought to correlate with an increase in intellectual complexity and sophisticated cognitive abilities. Given the relatively sophisticated cognitive abilities yet small brain size of living apes compared to humans and even early hominins, such as Australopithecus, observations of wild chimpanzees reactions to naturally occurring re can help in constructing hypotheses about the likely responses of early hominids to re. A recent article by Sayers and Lovejoy (2008) takes issue with chimpanzee (Pan troglodytes) models of human evolution, noting that scholars have overemphasized the importance of this species to the exclusion of others. Here, we argue that looking to particular aspects of chimpanzee behavior provides an important perspective informing our understanding of human evolution in
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distinguishing human autapomorphies from hominoid synapomorphies. We use data on the re-related behavior of savanna chimpanzees (P. t. verus) at Fongoli, Senegal to illuminate the similarities between great apes and humans regarding their reaction to re. In addition to their close phylogenetic ties to our lineage as a sister taxon, chimpanzees in Senegal inhabit an open, hot, and dry environment, making them ecologically important as a living primate model as well (Moore, 1996). As Fessler (2006) notes, humans may possess evolved psychological mechanisms dedicated to controlling re and others have noted that such mastery entails some

Grant sponsors: National Geographic Society, Leakey Foundation, National Science Foundation, Wenner-Gren Foundation for Anthropological Research, American Society of Primatologists, U.S. Fish & Wildlife Great Ape Conservation Fund, Primate Conservation Inc., Great Ape Trust of Iowa, Iowa State University College of Liberal Arts & Sciences, Department of Anthropology, Institute for Science and Society, Center for the Study of Violence. *Correspondence to: Jill D. Pruetz, Department of Anthropology, 324 Curtiss Hall, Iowa State University, Ames, IA 50010. E-mail: pruetz@iastate.edu Received 13 April 2009; accepted 6 November 2009 DOI 10.1002/ajpa.21245 Published online 21 December 2009 in Wiley InterScience (www.interscience.wiley.com).

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CHIMPANZEE RESPONSE TO FIRE degree of self-restraint in terms of the urge to ee (Goudsblom, 1986). This seems to be supported, in part, by the general avoidance behavior associated with re in most other animals (Sunquist, 1967; Geluso et al., 1986; De Ronde et al., 2004). However, little information on the proximate responses of animals to the presence of re is available. Indeed, numerous studies have addressed the responses of living people to re (Fessler, 2006), but we know little about how other primates react to re, save for their behavior in areas that have been burned (Berenstain, 1986; OBrien et al., 2003; Peres et al., 2003; Jaffe and Isbell, 2009). Given the history of difculty in assessing re use archaeologically (Pickering et al., 2008), estimates of which hominid species exhibited such behavior are necessarily conservative. Understanding how other hominoids react to re can assist anthropologists in developing hypotheses regarding which selective traits inuenced the development of re control in our own lineage, a characteristic that is unique in the animal kingdom (Goudsblom, 1986). Although savanna-dwelling animals can be routinely subjected to wildre, its presence can be detrimental to them in a number of ways, and avoidance behavior can be crucial to their survival. However, little has been published about the responses of large African mammals to wildre. Other animal species in such environments as well as savanna-dwelling mammals elsewhere provide some insight into the effects of such stresses. West African reed frogs (Hyperolius nitidulus) living in a savanna environment in Ivory Coast were observed to cue in to and ee the sound of re played back to them but did not react signicantly to white noise or a reverse playback of a re (Grafe et al., 2002). Although Christensen (1980) found that brush-tailed bettongs (Bettongia penicillata), a marsupial, were able to survive a high intensity re in a dry forest in Australia, they became more susceptible to predation because they were prone to becoming dazed or disoriented following the re. Woolley et al. (2008) found that, in addition to a general increase in mortality and injury, African elephants (Loxodonta africana) exhibited increased stress hormone levels after exposure to re. Here, we provide information on the proximate responses of wild, West African chimpanzees living in a savanna environment to seasonally occurring bush res. The chimpanzees conceptualization of re behavior (Trollope et al., 2004) is examined in light of the intellectual ability thought to characterize the earliest hominids to master the control of re.

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month, and a target of 100 hours of focal subject data were collected each month. Data on encounters with the seasonal res were collected opportunistically. Wildres are set yearly by humans for land clearing and hunting, and most areas within the Fongoli chimpanzee home range experience burning to some degree.

OBSERVATIONS Case 1. Late dry season re at Djendji Ravine, 14 March 2006

The observer (JP) rst heard sounds of re, such as grass stems and saplings popping, at 1100 hours, but could not yet see the re. By 1210 hours, the re had advanced to the woodland slope beyond a eld at the foot of the gallery forest ravine where the chimpanzees (a party of seven: ve adult males, one subadult male, one subadult female) were resting. Smoke was beginning to block out the sun. By 1320 hours, the sun was obscured by smoke. The adult male focal subject continued resting and grooming in the shade of this tiny patch (approximately 150 m in length and from 10 to 100 m in width) of gallery forest. At 1410 hours, he moved less than 100 meters from the southern to the northern slope of the ravine, but away from the re nearing the southern slope. At 1430 hours, the re reached the ravine area. Flames shot several meters high in grassy areas. Chimpanzees calmly moved away from the ravine, seemingly to avoid the re, and termite shed in woodland 200 m north. At 1645 hours the re was burning large trees on the southern slope, but at 1649 hours individuals were heard again in the ravine near where the re had been. At 1700 hours the ravine vegetation here burned also. Chimpanzees were in the vicinity of the re, but the observer was unable to see the distance between the re and the chimpanzees. The observer lost sight of chimpanzees at this point and, given that res were burning all around, left the area.

Case 2. 1 April 2006A late dry season re at Kerouani stream

During the late dry season a small party of chimpanzees encountered a wildre at a dry streambed where elephant grass (Pennisetum purporeum) was tall (over 2 m in height), dry and unburned (Table 1). Before the arrival of the re (i.e., while it was estimated to be more than 100 m away), unfamiliar vocalizations were heard twicea variation on a wraah-bark (Nishida et al., 1999), which may have been re-specic. Such a vocalization had never been recorded in over 2300 contact hours by JP with the Fongoli chimpanzees. The vocalization was most likely given by the alpha male, who moved in the direction of the re several minutes before the vocalizations were given, following his and the partys attention to the noise of the approaching re. However, the individual vocalizing was not visible to the observer, so this interpretation of the nature of the call remains somewhat speculative. The remainder of the chimpanzee party rst ascended a baobab (Adansonia digitata) tree upon seeing the approaching re. This organized move was not directed to feeding in the tree, although Saba senegalensis vines could be found there, and some individuals plucked fruit before they left. In ascending the tree, they moved closer, initially, to the re and then to heights of greater than 10 m. The remaining dominant male exhibited a slow American Journal of Physical Anthropology

Study site and subjects

The Fongoli chimpanzee community ranges within the Kedougou region in southeastern Senegal (12840N, 12813W). The savanna environment here is a mosaic of habitats, including woodland, grassland, bamboo, and small patches of gallery forest (Pruetz, 2006; Pruetz et al., 2008). Approximately 97% of the Fongoli communitys 65 km2 home range (excluding only the 3% that is gallery forest) can be considered re climax vegetation (Tappan et al., 2004). The dry season lasts over 7 months, and rainfall averages less than 1,000 mm annually (Pruetz, 2006). Study began at Fongoli in 2001 and, after four years of habituation effort, systematic behavioral data collection was initiated in the spring of 2005. The chimpanzee community consisted of 33 individuals: 9 adult females, 10 adult males, and 14 immatures. Chimpanzees were observed for at least 20 days per

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TABLE 1. Chimpanzees reaction to late, dry season re at Kerouani dry streambed, transcribed from audiotapea

Time of day 1149 hours 1152 hours 1202 hours 1220 hours 1225 hours Moments later 1229 hours 1300 hours

Observations Fire is several hundred meters away and can be heard (vegetation popping) approaching. Chimpanzees do not appear bothered, though dead grass is between them and the re. Flames are about 6 m high. Chimpanzees simply eat Saba fruit. Flames now 200 m away and [6 m high. One chimpanzee sleeps. BO eats fruit. Observer leaves to avoid the re. Chimpanzees vocalize again; seems like a re specic vocalization. There is also a re on the opposite side of the dry streambed. It is almost dark, due to smoke. Observer cannot see the chimpanzees. Chimpanzees approach, with YO displaying. BO carries fruit with him, eating. Fire is 20-30 m from the chimpanzees, as they sit in a tree, mainly to avoid YO, who displays below. The display appeared to be towards the re, given its similarity to the rain dance. Chimpanzees climb vines to reach a taller tree. Fire approaches from the opposite side. They appear to be escaping from the re. BI plucked a fruit before he ascended. Chimpanzees climb higher as the re nears. BI waited until he nished his fruit. Chimpanzees could have moved across the streambed but would have to keep moving or to skirt back around the re and move into a burned area. Instead, they climb a tree to about 20 m in height. Chimpanzees leave. YO descends, perhaps to avoid the re; ame shoots up high. Other chimpanzees leave. BI descends. Subjects hurry: LU with infant, YO, TI, and BI. They climb into vines. All have moved up the slope. All have moved. Fire was on three sides: east, west, and south. Chimpanzees moved to north slope and sit under vines. Fire came within 15 m of them. Most of the ames were low but at times would are higher; chimpanzees would ascend higher in the treesalthough the trees were only about 10 m talland would travel through them rather than coming to the ground. When the ames got higher in an area of tall grass about 20 m from them, they nally descended and moved up the slope. They sit, grooming. BI was one of the last to leave, nishing his fruit before he came to the ground rather than going through the trees. The movement through the trees was either to watch the re or to stay off of the ground. Fire returns from the south and moves around west as well as east. YO moved here rst, with others following from the east. No one appears bothered. FR sits with YO and his mother (FA), watching the re rather than following everyone else, as it approaches the hillside. Chimpanzees groom, periodically watching the re. Fire is about 30 m away and burning dry elephant grass. Chimpanzees nally move off, walking up the slope. A juvenile male looks back periodically as he unhurriedly moves off after the rest, about 15 m in front of the ames. They appear to have given up on avoiding it and are leaving as the re has come around on three sides of them.

1313 hours 1316 hours 1322 hours 1325 hours

a A total of 11 chimpanzees in this party: 2 adult males (BI, YO), 2 adult females (FA, LU), 1 subadult female (TI), 2 adolescent males (BO, JM), 2 juvenile males (FR, LT), 1 infant male (LX), 1 infant female (FN).

and exaggerated display toward the re in a manner analogous to the rain dance exhibited here and elsewhere where chimpanzees have been studied (Goodall, 1986). As the re approached, individuals moved into the very top of the crown. Immediately before the re reached the tree, however, the entire party led out and moved ahead of the re. The ames climbed more than halfway to the top of the tree. As the party moved away from the re, they often sat, rested, and/or ate Saba fruit they carried with them. The re varied in its speed but could cover distances of 10 m in a matter of seconds in areas of tall grass (i.e., high fuel levels). Rather than move long distances at once, they moved short distances of less than 2030 m, waited until the re neared almost to the point (about 1015 m depending on the intensity of the ames) where heat could be feltand moved a short distance again. Signicantly, the re swept through the area at midday, when chimpanzees normally rest in the few sources of shade that can be found, and activity is severely restricted. At 1300 hours, temperature in the shade was measured with a handheld thermometer as 378C.

DISCUSSION
Chimpanzees at Fongoli calmly monitor bush res at close range and change their behavior in anticipation of the res movement. We interpret the chimpanzees American Journal of Physical Anthropology

behavior as being predictive rather than responsive in that they showed no signals of stress or fear (e.g., fear grin, piloerection, Nishida et al., 1999) other than avoiding the re as it approached near them. Therefore, we maintain that they were predicting that the re would continue its pace of burning and its movement and were unconcerned, for example, that it would suddenly leap forward and burn them. The fact that they sat directly in front of the advancing re at a proximity that made the observer feel uncomfortable for her safety demonstrates that they were comfortable with their understanding of its behavior and their ability to predict its movements and adjust their responses to it. Although data are otherwise lacking regarding the responses of wild apes to re, anecdotal accounts of captive or formerly captive apes also reveal their ability to overcome a ight response. Captive chimpanzees exposed to an experimentally introduced campre were not distressed by its presence, and a male displayed toward the re, even disrupting it by pulling out pieces of wood with his hands as he passed (Charmove, 1996). Rehabilitant orangutans (Pongo pygmaeus) have been observed to imitate humans in making res (Russon and Galdikas, 1999), and chimpanzees in a zoo in South Africa were observed to light and extinguish cigarettes without being burned (Brink, 1957). These behaviors demonstrate the cognitive ability to adjust to a potentially harmful agent. In the case of high-intensity wildres, such as those experienced by chimpanzees at Fongoli, a number of

CHIMPANZEE RESPONSE TO FIRE variables must be taken into account in order to successfully negotiate such a re. The behavior of re is complex, as its movement depends on wind speed and fuel sources as well as other factors, such as slope and relative humidity (Trollope et al., 2004). We suggest that the control of re by humans is the endpoint of a complex evolutionary process that involves the acquisition of at least three cognitive stages: 1) conceptualization of re, i.e. an understanding of the behavior of re under varying conditions that would allow one to predict its movement, thus permitting activity in close proximity to the re. 2) The ability to control a re, involving containment, providing or depriving the re of fuel, and perhaps the ability to put it out. 3) The ability to start a re. The rst ability is a prerequisite to the other two and has been overlooked in models of the stages of re use by humans (Frazer, 1930; Goudsblom, 1986). It likely preceded two and three, and we propose that it characterizes chimpanzees living today. The chimpanzees at Fongoli specically must predict the behavior of a head re or the leading edge of a re, which, in a surface re fed by ne fuels such as grasses and twigs, can move rapidly (Trollope et al., 2004). This suggests that they have acquired the rst cognitive ability mentioned above: conceptualization or forming a mental prediction of the res movement. Variables that should be taken into account include ame height and width and re intensity, as inuenced by topographic and climatic factors. Considering such variables and predicting the behavior of re is a complex task (Trollope et al., 2004). As Sayers and Lovejoy (2008) note in a recent essay detailing reasons chimpanzees have been overemphasized as models in the study of human evolution, the differences between nonhuman and human primates (including hominids) are the traits on which paleoanthropologists should focus (i.e., autapomorphies). We concur on this point but argue that paleoanthropologists should continue to recognize the specic importance of studies of chimpanzees in informing a subdiscipline that is necessarily hindered by the fact that behavior does not fossilize. For example, forming hypotheses about the earliest stages of humans control and use of re can be little informed from ethnographic records of living humans, as evidence suggests that no society on record has lacked such ability (Goudsblom, 1986). Recognizing similarities among hominoids, for example, works to this end, although in a manner different from the one Sayers and Lovejoy (2008) suggest. Rather than focusing on differences between humans and other species, identifying similarities between our species and chimpanzees in fact furthers our attempts to identify uniquely human traits and, importantly, commonly aids in correcting our views of what is and is not a derived hominid trait or behavior (e.g., Hernandez-Aguilar et al., 2007). Just as studies of fossilized remains of extinct organisms may elucidate the range of possible morphological variation within a taxon, thereby clarifying the systematic relationships within that taxon (by serving as outgroups), behavioral studies of modern organisms that are evolutionary outgroups can elucidate the range of possible behaviors within taxa that include extinct species. Additionally, the issue often in question is the underlying cognitive ability relative to a particular type of behavior (e.g., hunting, communication). For example, the suggestion that australopithecines could have exhibited tool-assisted hunting refers to the cognitive abilities underlying this behavior rather than the selective pressures or motoric

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abilities associated with tool-assisted hunting (Pruetz and Bertolani, 2007). This issue is key to the argument for using closely related species as referential models and one that Sayers and Lovejoy (2008) as well as their list of respondents in Current Anthropology give relatively little overt attention. Given that brain size of fossil hominids is used as a proxy for cognitive sophistication, understanding the range of behaviors of living primates with similar brain size in referential models can continue to provide paleoanthropologists with information that allows them to construct conceptual models. The ability to conceptualize the behavior of re may be a synapomorphic trait characterizing the humanchimpanzee clade, and this possibility should be investigated more closely. Although living humans preoccupation with re in some cases is seen as aberrant, Fessler (2006) points out that we know little about learning processes associated with re as well as cultural differences in behavior toward it. Rolland (2004) argues that, like language, the use and control of re has no behavioral antecedents in the animal world, while Goudsblom (1986) further notes that, while rudimentary language abilities are seen in some nonhuman primates, only our own lineage has mastered the use of re. We counter that, while the control and use of re is uniquely human, the cognitive capabilities necessary to conceptualize the behavior of re appear to be present in our closest living relatives and likely other primates (as well as some other cognitively sophisticated mammals such as elephants) similar to what has been observed in great apes regarding language abilities (Segerdahl et al., 2005). Our observations at Fongoli suggest that, like humans, chimpanzees are able to control their fear impulse in response to re (Goudsblom, 1986). If we accept the idea that conceptualization of re is prerequisite to its control and use, then identifying such conceptualization at a more basal node in the hominid phylogeny than previously hypothesized implies that control and use might have been possible earlier than formerly thought based on the date of rst appearance in the fossil record. Thus, if the cognitive underpinnings of re conceptualization are a primitive hominid trait, hypotheses geared to explaining the control and use of re in certain taxa may need to be re-examined.

ACKNOWLEDGMENTS
The Republic of Senegal, Department of Eaux et Forets, and the Arrondissement du Bandafassi granted kind permission for work in Senegal. Assistance in the eld at Fongoli has been provided by D. Kante, M. Camara, M. Sahdjaro, P. Bertolani, S. Bogart, K. Boyer, M. Cook, M. Gaspersic, S. Johnson-Fulton, S. Lindshield, T. Martz, and M. Waller. Thanks to T. Pickering, K. Walkup, and K. Klag for helpful comments on an earlier version of this manuscript. Research reported here complies with the regulations enforced by the Iowa State University Institutional Animal Care and Use Committee (IACUC).

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