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Reconstructing hominin interactions with mammalian carnivores (6.0 - 1.8 Ma)

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Reconstructing hominin interactions with mammalian carnivores (6.0 - 1.8 Ma)
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  17 Reconstructing Hominin Interactionswith Mammalian Carnivores(6.0–1.8 Ma) Adrian Treves and Paul Palmqvist Introduction Several hominin genera evolved to use savanna and woodland habitats acrossPliocene Africa. This radiation into novel niches for apes occurred despite adaunting array of carnivores (Mammalia, Carnivora) between 6.0 and 1.8 Ma(Figure 17.1). Many of these carnivores would have preyed on hominins if giventhe opportunity. In this paper we ask what the behavioral adaptations were thatpermitted hominins to survive and spread, despite this potentially higher risk of predation in ancient Africa.When considering hominin anti-predator behavior, many scholars looked firstto material culture, such as fire or weaponry (Kortlandt, 1980; Brain, 1981). How-ever, the idea that deterrent fire or weaponry freed early hominins from threatsposed by predators is unsatisfying for several reasons. First, the modern carnivoresnow roaming Africa are survivors of humanity’s repeated and systematic cam-paigns to eradicate problem animals, trade in skins, and so on. (McDougal, 1987;Treves & Naughton-Treves, 1999), whereas Pliocene carnivores would not havehad a history of conflict with armed hominins. Second, thousands of modernhumans fell prey to leopards ( Panthera pardus ), lions ( P. leo ) and tigers ( P. tigris )in the twentieth century despite their sophisticated weapons and fire (Turnbull-Kemp, 1967; McDougal, 1987; Treves & Naughton-Treves, 1999; Peterhans &Gnoske, 2001). Although, thorn branches, stone tools, fire brands, pointed sticks,orbonescouldpotentiallyhelptorepelcarnivoresfromtheirkills(Kortlandt,1980;Bunn & Ezzo, 1993; Treves & Naughton-Treves, 1999), such weaponry seemswholly inadequate for personal defense when large carnivores achieve surprise,attack in a pack, or are accustomed to overcoming heavier prey defended by horns,hooves, or canines. Therefore, we assert that weaponry by itself does not nul-lify the risk posed by predators. Moreover, controlled use of fire and stone tooltechnology appear late in the archaeological record relative to the evolution of  355  356 A. Treves and P. Palmqvist 6  5 4 3 2 CARNIVORAHOMININSArdipithecusOrrorinAustralopithecus *  includes KenyanthropusParanthropusMaHomo * AgriotheriumDinofelisMachairodusChasmaporthetesHomotherium 1 PachycrocutaAcinonyxMegantereonLycaonPantheraCrocuta, Hyaena, Parahyaena F IGURE  17.1. Time spans of paleopredator and hominin genera in Africa. semi-terrestrial hominins in Pliocene Africa (Bellomo, 1994; Brain, 1994; Wolde-Gabriel et al., 1994; Brunet et al., 1997; Leakey et al., 1998; Haile-Selassie, 2001).Hominin anti-predator behavior remains a key puzzle of our human ancestry.InthenextsectionofthischapterwereviewAfricanlargecarnivoreecologyandhunting behavior in extant taxa and that reconstructed for Plio-Pleistocene forms(“paleopredators” hereafter). Following this, we review the anti-predator behaviorof hominins by analogy with monkeys and apes; this analogy is parsimoniousbecauseoftheobservedcross-taxonomicconsistencyoftheirbehavioralresponsesto predators. Vigilance behavior in relation to social organization is particularlyinformative. Finally, we integrate the two reviews to reconstruct the range of anti-predator behaviors open to hominins.  African Large Carnivores, Past and Present  Africa has long contained diverse carnivore communities (Figure 17.1). Carni-vores have repeatedly radiated into various niches, including specializations for  17. Hominin Interactions with Mammalian Carnivores 357 predation, active or passive scavenging, open-country or forested habitats, andsmall or large ungulate prey (Table 17.1).Following Sunquist & Sunquist (1989) we define a “large carnivore” as anyspecies with average individual or group body mass  > 34 kg (e.g.,  Hyaena hyaena or  Lycaon pictus , respectively). Subsequent reference to large/small prey relate tothe carnivore under discussion.Large carnivore diversity was greater in Africa’s past than it is today (Figure17.1). Between 6 and 3.6 Ma there were five genera of large carnivores withoutextant analogues (the long-legged ursid  Agriotherium , the large coursing hyaenid Chasmaporthetes , and the saber-toothed felids Homotherium,  Machairodus  and  Dinofelis ). Then, from the mid-Pliocene (3.6 Ma), the archaic genera were joinedby one large canid (  Lycaon lycaonoides ) (Mart´ınez-Navarro & Rook, 2003), threenew large felid genera (  Acinonyx, Megantereon  and  Panthera ), and four new gen-era of hyaenids ( Crocuta, Pachycrocuta, Hyaena , and  Parahyaena ). At some sites,8–10 species appear to have been coeval and broadly sympatric (Barry, 1987;Turner & Anton, 1997)(Figure 17.1). Niche separation under such conditions isnot yet clear.As the Pleistocene wore on (1.8 Ma onward) the archaic carnivores went extinctinAfrica,partlyasaresultofaglobalcarnivoreguildturnoverandspeciesreplace-ment (Figure 17.1). The African faunal turnover coincided with a decrease inwoodland relative to grassland, more herd-living grazing ungulates, and fewersolitaryor small-group-living large herbivores like giraffids, rather than from com-petition between the modern carnivore guild and archaic forms (Hendey, 1980;Turner, 1990; Werdelin & Turner, 1996; Turner & Anton, 1998).Coexistence of hominins and carnivores is insufficient by itself to concludethat hominins evolved effective anti-predator defenses against such paleopreda-tors. Coexistence would have had little selective impact if (a) carnivores didnot kill Pliocene hominins regularly, or (b) if such predation were random withrespect to hominin traits. Thus, in the following sections we assess whetherpaleopredators killed hominins regularly, and if so, were there consistent patternsof hominin-carnivore interactions that might have produced directional selectionamong hominins.  Habitat Selection Carnivoresgenerallygowherepreyaremostabundant,butmanywillestablishanddefend territories year-round. Except for the leopard, all the extant African largecarnivores are most abundant in open savannas and savanna-woodlands (variablemixtures of trees, grassland, and bushland where visibility is less than 100 m onaverage), coincident with highest ungulate densities (Table 17.1). Nevertheless,several carnivores can breed successfully within very arid regions or dense forest(Leakey et al., 1999; Bailey, 1993). The leopard is the greatest habitat generalisttoday, breeding from rainforest to desert, albeit preferring habitat with vegetationcover.      T     A    B    L    E     1    7 .    1 .    A    f   r    i   c   a   n    l   a   r   g   e   c   a   r   n    i   v   o   r   e   s ,   p   r   e   s   e   n    t   a   n    d   r   e   c   o   n   s    t   r   u   c    t   e    d .     P   r   e    d   o   m    i   n   a   n    t    H   u   n    t    i   n   g    B   e    h   a   v    i   o   r    G   e   n   u   s    M   a   s   s ,    k   g    B   r    i   e    f    d   e   s   c   r    i   p    t    i   o   n   o   r   c   o   m   m   o   n   n   a   m   e    A   c    t    i   v    i    t   y    G   r   o   u   p    i   n   g    A    t    t   a   c    k    H   a    b    i    t   a    t    *     A   c    i   n   o   n   y   x 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   w   o   o    d    l   a   n    d    ”    i   n    d    i   c   a    t   e   s    l   e   s   s    t   r   e   e   c   o   v   e   r    t    h   a   n   g   r   a   s   s   c   o   v   e   r .    S   o   u   r   c   e   s   :    A   n   y   o   n   g   e ,    1    9    9    3 ,    1    9    9    6   ;    A   r   r    i    b   a   s    &    P   a    l   m   q   v    i   s    t ,    1    9    9    9   ;    B   a    i    l   e   y ,    1    9    9    3   ;    B   a   r   r   y ,    1    9    8    7   ;    B   e   r    t   a ,    1    9    8    1   ;    B   o   a   z   e    t   a    l . ,    1    9    7    9   ;    B   r   a    i   n ,    1    9    8    1 ,    1    9    9    4   ;    B   u   s   s   e ,    1    9    8    0   ;    C   a   r   o ,    1    9    8    7 ,    1    9    8    9   a ,    b   ;    C   o   o    k   e ,    1    9    9    1   ;    F   a   n   s    h   a   w   e    &    F    i    t   z    G    i    b    b   o   n ,    1    9    9    3   ;    F   e   r   r   e    t    t    i ,    1    9    9    9   ;    F    i    t   z    G    i    b    b   o   n ,    1    9    9    0   a ,    b   ;    G   e   r   a   a    d   s ,    1    9    9    7   ;    H   e   n    d   e   y ,    1    9    7    4 ,    1    9    8    0   ;    H   o    l   e    k   a   m   p   e    t   a    l . ,    1    9    9    7   ;    H   u   n    t ,    1    9    9    6   ;    K   e   y   s   e   r ,    1    9    9    1   ;    L   e   a    k   e   y ,    1    9    9    9   ;    L   e   w    i   s ,    1    9    9    7   ;    M   a   r   e   a   n ,    1    9    8    9   ;    M   a   r    t    i   n ,    1    9    8    9   ;    M   a   r    t    ´   ı   n   e   z  -    N   a   v   a   r   r   o   a   n    d    P   a    l   m   q   v    i   s    t ,    1    9    9    5 ,    1    9    9    6   ;    M    i    l    l   e   r    &    C   a   r   r   a   n   z   a ,    1    9    9    6   ;    M    i    l    l   s ,    1    9    8    9   ;    P   a    l   m   q   v    i   s    t ,    2    0    0    2   ;    P   a    l   m   q   v    i   s    t    &    A   r   r    i    b   a   s ,    2    0    0    1   ;    P   a    l   m   q   v    i   s    t   e    t   a    l . ,    1    9    9    6 ,    1    9    9    9   ;    P   e    t    t   e   r   e    t   a    l . ,    1    9    9    4   ;    R   o   o    k ,    1    9    9    4   ;    S   c    h   a    l    l   e   r ,    1    9    7    2   ;    T   a   y    l   o   r ,    1    9    8    9   ;    T   u   r   n    b   u    l    l  -    K   e   m   p ,    1    9    6    7   ;    T   u   r   n   e   r    &    A   n    t   o   n ,    1    9    9    6 ,    1    9    9    7 ,    1    9    9    8   ;    T   u   r   n   e   r ,    1    9    9    0 ,    1    9    9    7   ;    W   e   r    d   e    l    i   n ,    1    9    9    4 . 358  17. Hominin Interactions with Mammalian Carnivores 359 As far as micro-site selection for hunting, only the leopard is known to huntarboreal prey within 10–15 m of the ground. Leopards also kill in caves, cliff sides and houses (Simons, 1966; Turnbull-Kemp, 1967).Among extinct carnivores, habitat use varied (Table 17.1).  Agriotherium  andmachairodonts  Dinofelis  and  Megantereon  are believed to have selected moreforested habitats based on their postcranial morphology, typical of stalking,ambush hunters. The latter two genera show relatively more robust forelimbsthan hindlimbs. A comparative study of the postcrania in modern and Plio-Pleistocene carnivores shows  Dinofelis  resembles pantherine felids craniodentally,and its postcrania resembling modern prey-grappling lions, tigers, and leopards(Marean, 1989; Anyonge, 1996; Lewis, 1997). The postcrania of   Megantereon reveal tree-caching and long-distance dragging capabilities, as in modern leopardsand jaguars ( Panthera onca ) (Lewis, 1997; de Ruiter & Berger, 2000).  Homoth-erium  and  Machairodus  postcrania suggest cursorial tendencies in more openhabitats, given their comparatively higher values for both brachial and cruralindexes (Table 17.1).  Chasmaporthetes  and  Lycaon  have been associated withopen-country habitats as well—although it should be noted that  Lycaon  today canhunt quite successfully in dense shrub land (Creel & Creel, 1995). The giant hyena Pachycrocuta  was associated with more open habitats, particularly where mediumto large ungulate carcasses were left by machairodont felids (Arribas & Palmqvist,1998) (Table 17.1).Associations of fossil hominins with remains of   Chasmaporthetes ,  Dinofelis ,  Homotherium ,  Machairodus , Megantereon, and  Pachycrocuta  indicate sympatryin the period 6.0–1.8 Ma in habitats reconstructed as a mixture of woodlandsand open country (Cooke, 1991; Keyser, 1991; Brain, 1994; Brantingham, 1998;Dominguez-Rodrigo & Pickering, 2003; Palmqvist et al., 2005). At a finer level,felid and hyaenid activity was considerable in and around the same caves withhominin remains (Brain, 1981, 1994; Turner, 1990). Deep caves would there-fore have been dangerous resting sites (for vivid examples, see Simons, 1966;Brain, 1981). However, there seems to be evidence that hominins went voluntar-ily to caves used by paleopredators. For example, the presence of Plio-Pleistocenestone tools in South African caves without evidence of their manufacture(Brain, 1981) suggests that hominins came to some of these sites voluntarily (car-rying tools) most likely, or, less likely, that predators transported their carcasseswithout losing the tools (e.g., in a portable container that neither fossilized nordropped off the carcass when dragged).The extant carnivores hunt by day and night, but seem to do so most oftenor most successfully between 19.00 and 07.00 with the exception of the diurnalcheetah (Table 17.1). There is some indication that carnivores hunt less by daywhenhumans poseathreattothem (Turnbull-Kemp, 1967; van Schaik&Griffiths,1996), a benefit hominins would not have enjoyed in the Pliocene for the reasonsmentioned before. Moreover, observations of predation reveal that carnivores killprimates in the day as well as at night (reviews in Treves, 1999a; Boinski et al.,2000). Thus, hominins could not have escaped predation simply by diurnality.
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