Hunting strategies of and with canids in the Southern Cone and their effects on prey adaptations
It is difficult, based on morphology alone, to deduce behavioural strategies, given the amount of inter- and intraspecific variation in canids ( Sillero-Zubiri & Macdonald 2004). Paleontologists, however, have noted that morphological specializations can favour particular hunting strategies in contemporary canids and felids, and thus by analogy there may have been similar tradeoffs in extinct mammals (Andersson 2005). For example, limbs can be cursorially specialized, elongating in the course of evolution to allow a wider stride, or specialized to “supinate the forearm, and thus grapple prey” when pouncing on it (Andersson 2005 p. 57). The latter is a common strategy used by ambush predators like felids. When the prey cannot be effectively grappled with the forearms, bringing down large prey with the mouth is the alternative; this latter kill strategy favours pack hunting and is usually seen in cursorial hunting tactics. Nevertheless, Andersson (2005) highlights exceptions. For example, lions, which can grapple excellently, are group hunters (using an ambush strategy), while S. venaticus , the bush dog, has very poor cursorial adaptations but is a group hunting species that brings prey down by biting. Ecological conditions such as shrub cover, by creating physical constraints, can also affect hunting strategies (Thibault & Ouellet 2005, Karanth & Sunquist 2000; Fanshawe & Fitzgibbon 1993). Group hunting may contribute flexibility to hunting. Fanshawe and Fitzgibbon (1993), for example, show that African wild dogs, Lycaon pictus, unlike solitary ambush predators, were equally successful in killing prey in different amounts of cover, and alone or in groups of different sizes. While pack-hunting cursorial species such as C. lupus typically “troll” for and “test” prey in a cursorial mode, they also have group hunting tactics that include ambushes (Fox 1987). Many contemporary canid species hunt small prey alone and large prey in packs (e.g. Canis lupusC. latransC. adustusS. venaticus ; Andersson 2005). All of this indicates that it is not possible to clearly delimit two hunting strategies, since predator behaviour will depend on multiple factors.
The variation in predator behaviour makes a difference to prey. When faced with ambush vs. cursorial predators, large prey animals alter their anti-predator behaviours, notably by being more vigilant in habitat types associated with stalking and ambushing predators (Makin et al. 2017). Preissner et al. (2007) conclude that predator identity, e.g. hunting strategy, matters to prey life-history tradeoffs.
D. avus was the only large Pleistocene canid found in Chile (Castillo, 2007). Prevosti et al. (2009) report that D. avus likely predated Lama spp. as well as other large and medium animals, although they also indicate that Lama spp. may often have been scavenged rather than killed (Prevosti & Vizcaíno 2006; Prevosti & Martin 2013). The extinct Dusicyon spp. are described as “large foxes” (e.g. Prevosti et al. 2009), a description that may suggest that they did not live or hunt in packs, although like large-fox-like coyotes (Canis latrans) they might have lived in packs but hunted alone or in pairs. Coyotes are cursorial predators adapted to hunting in the open plains and prairies of North America (Thibault & Ouellet 2005). At least part of the distribution of Dusicyon was open plains, so it could have shared a similar hunting strategy. Yet, as for L. pictus (Fanshawe & Fitzgibbon 1993), they might be equally successful with a lone-hunting cursorial strategy in areas with tree cover, at least for certain prey. The maned wolf, Chrysocyon brachyurus is also large (up to 23 kg) (Sheldon 1992) and described as fox-like (Dietz 1985), and in some accounts a sister genus of Dusicyon spp (Austin, 2013). While it has never occurred in Chile to date (Torres et al. 2013), it would have coexisted with Lama spp., rheas (Rhea spp. ), and other prey species whose ranges included Chile during the Holocene. It can thus both suggest potential D. avus hunting strategies by analogy, and suggest the kinds of strategies species found in Chile would also have been adapted to. Maned wolves are opportunistic and flexible omnivores that eat a variety of plant parts, insects, small animals, and medium sized animals of a similar mass to its own, including armadillos, Pampas deer (Ozotoceros bezoarticus , 22-34 kg) and Greater rheas (Rhea americana , 20-27 kg) (de Almeida Jácomo et al. 2004; Aragona & Setz 2001). Maned wolves hunt alone, and usually at night (Sheldon 1992; de Melo et al. 2007). They are typically found in open woodlands (cerrado) and tropical grasslands. Neither Dietz (1985) nor Sheldon (1992) clarify their hunting strategy or behaviour, but the long legs might indicate a cursorial adaptation. Alternately, it might simply (or also) be an adaptation to survey prey over the high grasses of the tropical grasslands. However, it was considerably smaller than an adult guanaco (L. guanicoe), and thus is unlikely to have actively hunted adults if it hunted alone. Thus, as for the culpeo fox, or coyotes in the context of llamas, we might expect that adult guanacos may have been able to counter-attack lone-hunting maned wolves as well as D. avus .
The surviving Pleistocene Lycalopex spp. found in Chile, as reported above, anecdotally attack juvenile camelids and prey on entrapped adults. However presumably it is difficult for the lone-hunting Lycalopex spp. to bring down adult guanacos or native deer, due to their small size, and their lack of a group hunting strategy (cf. the equally small bush dogs, Speothos venaticus , which are only able to bring down deer by biting them in large packs (Biben 1982; Sheldon 1992)). Otherwise, Lycalopex spp. can prey on fawns of medium and large ungulates (Corti et al. 2010).
We cannot necessarily conclude that widely distributed species such as guanacos, which would have been exposed prehistorically to cursorial pack-hunting, had different anti-predator adaptations in the populations to the west and the east of the Andes. However, it is possible that with the only remaining pack-hunting species in South America being the bush dog in the Amazonian basin, prey in the Southern Cone like guanaco may have lost, since the Pleistocene extinctions of the hypercarnivorous species, adaptations specific to surviving group predation tactics, whether cursorial or ambush.
By contrast, domesticated dogs may have been incorporated into particular roles in hunting with humans. Lupo (2017) presents ethnographic evidence that of the many forms of hunting with dogs, most of them, under most circumstances, have no appreciable benefit for hunting success. She argues that dogs are least useful for large prey that need to be hunted by stealth, and often interfere in ambushes and traps, but can be useful for finding and flushing prey, and handling certain prey. Although a large increase in effectiveness of hunting (compared to hunting without dogs) is observed when dogs are introduced as novel predators to islands and used in packs in combination with guns, other factors such as colonialism and land-use change also simultaneously alter technologies, economic drivers, and environmental conditions, such that the role of dogs alone is unclear in driving hunting outcomes (Lupo 2017). We cannot assume that even if hunting with larger dog packs might be more effective, hunters would necessarily prefer this tactic, since hunter-gatherers and many agriculturalists do not purposefully maximize productivity (Sahlins 1974[2017]). Rather, the evidence from Lupo (2017) is more consistent with viewing dogs as part of the social group (see the discussion of taming, above), than as hunting tools. How, in fact, were dogs incorporated into hunting tactics in Chile and the Southern Cone?
The dogs of the northern Andean civilizations and the north of Chile are all rather small, but some of them may have been used to corner game. The munutru was reportedly also small. The dogs of the Selk’nam, which appear to be medium-sized and shaped like dingos (Alvarado et al. 2007), had roles including chasing down guanacos that ran off after being stalked and struck by an arrow, taking down guanacos that the hunters were ambushing, or helping chase guanacos towards an ambush site (Legoupil 2011). Belardi et al. (2017) describe early Holocene communal hunting strategies to the East of the Andes, in Patagonia, that involved one group of hunters driving guanacos towards another group lying in wait; later on, constructed blinds were used. Santiago & Salemme (2016) report that in addition to using group ambush tactics such as those described above, Selk’nam men often hunted alone, and women sometimes hunted with dogs: they do not clarify whether this was by pursuit or stalking. Across the Americas and across the Holocene, communal hunting driving large prey towards ambushes, opportunistic hunting of large animals mired in natural “traps” such as bogs and tar pits, and building traps including nets, are among the many tactics thought to have been used, with or without dogs (Davis & Reeves 2014). The use of trapping technologies and tactics can be seen as a kind of displaced ambush, which like other forms of ambush require the prey to be highly alert and discerning about danger cues that try to blend into the environment (Gell 1996). We have not found clear evidence, before horses and cars, of pursuit followed by killing as a human hunting tactic for guanaco or other prey in Chile and the Southern Cone. Consequently, if indeed human hunting with dogs to the west of the Andes and the extreme south of the continent predated Europeans (as seems plausible), in itself this does not mean that guanacos or other prey were exposed to the same selective pressures as pack-hunting cursorial feral dogs pose today.
Conclusions
Why are the key large and medium prey species of Chile—camelids and deer—apparently unadapted to cursorial pack hunting by contemporary feral dogs? In summary, west of the Andes (within Chile) there is no clear evidence of group-hunting species or cursorial specialists large enough to bring down adults of these species (with the possible exception of the very small pudu Pudu puda ). However, the guanaco in particular has had a range throughout the Southern Cone, and should have been exposed to Pleistocene pack-hunting species that would have used cursorial strategies. It is unclear evolutionarily speaking whether appropriate anti-predation strategies might have been found only in the guanaco populations exposed to these predation pressures East of the Andes. The biology and evolution of predation defense suggests that suites of different kinds of defense behaviours can be maintained as long as any kind of predators are present (Blumstein 2006). Since there have always been puma and foxes in Chile, we would thus expect the prey species to maintain the capacity for any anti-predator behaviours that they had had to evolve.
For guanacos with their wide distribution in particular we can thus outline three possible scenarios or hypotheses: (1) only certain guanaco lineages east of the Andes were adapted to group-hunting cursorial strategies, and these populations have gone extinct (or are isolated from the Chilean populations); (2) the entire species had these adaptations but have lost them with the extinction of the predators in question at the end of the Pleistocene, contrary to the implication of Blumstein (2006); (3) these adaptations are latent in all guanacos’ repertoire of possible adaptative responses to predation but for some reason which remains unclear (see below) seem to be expressed inadequately in Chilean populations. As for deer species and other camelids of Chile, these species have smaller ranges largely restricted to the Andes and the west of the Andes, so their prehistoric exposure to group-hunting cursorial strategies of predators east of the Andes is less clear, but possibly null.
However, these hypotheses are complicated by also considering the factor of indigenous people’s Pre-Columbian hunting strategies involving domesticated canids, whether dogs or other species. It is probable that human hunters throughout the Southern Cone including Chile used dogs to assist in hunting well before the arrival of Europeans. These strategies were probably, given historical and comparative ethnographic evidence, primarily ambush strategies, although we cannot totally rule out cursorial-type strategies since group hunting in canids (and in humans) is usually quite flexible in form. So, we suggest two further hypotheses: (4) Pre-Columbian indigenous hunters also had an advantage in killing prey with canids similar to what is currently observed in canid hunting in Chile today, and for some reason prey species largely did not adapt to this group-hunting attack strategy perhaps due to lack of historical depth or continuity of the practices, or a continually changing habitat structure under human influences (which would affect predation strategies and success rates); (5) prey species did adapt to the group-hunting human-dog strategy, but only to the dominant ambush strategies, which is why they remain vulnerable to cursorial attacks. Again, it is not totally clear from the biology of adaptation to predation whether these two anti-predation strategies can be neatly separated in this way, since most examples of flexible prey anti-predation adaptations come from prey exposed to both ambush and cursorial strategies. This is an implicit sub-hypothesis that requires further research.
Although the logic of Geist (1998) suggesting a lack of cursorial predators is persuasive to explain deer adaptations, when also considering camelids, and when carefully looking at the evidence, the situation appears to be more complex than his argument accounts for.
We conclude that our state of knowledge about the biology and evolution of prey adaptations to different predation strategies, and the state of the base evidence for the existence of different canids in Chile and the rest of the Southern Cone of South America, are inadequate to address how the problem of feral dog predation on native species in these regions might be solved. For example, if native prey species really have no evolutionary background of exposure to group-hunting cursorial species, then only a natural process of selection could lead to its emergence. Whether it would be better, or more feasible, to expose native prey species to feral dogs in the hope of their eventual adaptation, or to eradicate feral dogs, is an open question. On the other hand, if guanacos in particular, and perhaps other species, have latent adaptations to group-hunting cursorial species, it is somewhat mysterious as to why they are so often killed by feral dogs. Here, there may be two more hypotheses to consider, which are quite different. (6) It could be the case that the rate of feral dog hunting success is not exceptionally high compared to “natural” predation, but is problematized due to its seeming unnaturalness. With current information this is difficult to assess. (7) It could be that camelids and deer need to learn socially (from parents or others) how to use certain anti-predator strategies of which they are behaviorally capable (Wiedenmayer 2009), but this possibility has been lost at some point in history due to loss of the behavioral expression among Chilean populations, either due to a Pleistocene predator extinction, or the local discontinuation of human hunting with dogs among relictual deer and camelid populations. In the latter case, training individuals to respond correctly to cursorial group attacks (Griffin et al. 2000) could be an approach to mitigate the problem by allowing social learning to spread.
Although we cannot offer any answers, we have clearly outlined the existing relevant knowledge and developed an array of hypotheses that should be tested, both to advance general knowledge of adaptations to predation, and to assist conservationists in designing appropriate interventions to conserve camelids and deer in South America and Chile in particular.
Acknowledgements
MR-B thanks WCS Chile for including her in the Grupo Núcleo Guanacos, which has been a useful forum to share information on threats to guanaco conservation.
Data accessibility statement
This is a qualitative review and analysis and there is no original or re-analysed data to share beyond the text found in this document.
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