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 lupus , C.
latrans , C. adustus , S. 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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