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\begin{document}

\title{History of canids in Chile and impacts on prey adaptations}



\author[1]{Benjamín Silva}%
\author[2]{Meredith Root-Bernstein}%
\affil[1]{Pontifical Catholic University of Chile}%
\affil[2]{CNRS}%


\vspace{-1em}



  
  \date{May 25, 2024}


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\selectlanguage{english}
\begin{abstract}
Artiodactyl prey species of Chile, especially guanacos (\emph{Lama
guanicoe}) are reported to be very susceptible to predation by pack
hunting feral dogs. It has been previously suggested that guanacos and
endemic South American deer may have evolved in the absence of
pack-hunting cursorial predators. However, the paleoecology of canid
presence in southern South America and Chile is unclear. Here, we review
the literature on South American and Chilean canids, their
distributions, ecologies and hunting behaviour. We consider both wild
and domestic canids, including Canis familiaris breeds. We establish two
known antipredator defense behaviours of guanacos: predator inspection
of ambush predators, e.g. \emph{Puma concolor}, and rushing at and
kicking smaller cursorial predators, e.g. \emph{Lycalopex culpaeus}. We
propose that since the late Pleistocene extinction of hypercarnivorous
group-hunting canids east of the Andes, there were no native species
creating group-hunting predation pressures on guanacos. Endemic deer of
Chile may have never experienced group hunting selection pressure from
native predators. Even hunting dogs (or other canids) used by indigenous
groups in the far north and extreme south of Chile (and presumably the
center as well) appear to have been used primarily within ambush hunting
strategies. This may account for the susceptibility of guanacos and
other prey species to feral dog attacks. We detail seven separate
hypotheses that require further investigation in order to assess how
best to respond to the threat posed by feral dogs to the conservation of
native deer and camelids in Chile and other parts of South America.%
\end{abstract}%



\sloppy


History of canids in Chile and impacts on prey adaptations

Benjam\selectlanguage{ngerman}ín Silva Rochefort\textsuperscript{1} \& Meredith
Root-Bernstein\textsuperscript{2,3,4}

\begin{enumerate}
\tightlist
\item
  Facultad de Ciencias Biológicas, Universidad Católica de Chile,
  Santiago, Chile
\item
  Center of Applied Ecology and Sustainability, Santiago, Chile
\item
  Institute of Ecology and Biodiversity, Santiago, Chile
\item
  UMR CESCO, CNRS, Musée National d'Histoire Naturelle, Paris, France
\end{enumerate}

\textbf{Abstract}

Artiodactyl prey species of Chile, especially guanacos (\emph{Lama
guanicoe} ) are reported to be very susceptible to predation by pack
hunting feral dogs. It has been previously suggested that guanacos and
endemic South American deer may have evolved in the absence of
pack-hunting cursorial predators. However, the paleoecology of canid
presence in southern South America and Chile is unclear. Here, we review
the literature on South American and Chilean canids, their
distributions, ecologies and hunting behaviour. We consider both wild
and domestic canids, including \emph{Canis familiaris} breeds. We
establish two known antipredator defense behaviours of guanacos:
predator inspection of ambush predators, e.g. \emph{Puma concolor} , and
rushing at and kicking smaller cursorial predators, e.g. \emph{Lycalopex
culpaeus} . We propose that since the late Pleistocene extinction of
hypercarnivorous group-hunting canids east of the Andes, there were no
native species creating group-hunting predation pressures on guanacos.
Endemic deer of Chile may have never experienced group hunting selection
pressure from native predators. Even hunting dogs (or other canids) used
by indigenous groups in the far north and extreme south of Chile (and
presumably the center as well) appear to have been used primarily within
ambush hunting strategies. This may account for the susceptibility of
guanacos and other prey species to feral dog attacks. We detail seven
separate hypotheses that require further investigation in order to
assess how best to respond to the threat posed by feral dogs to the
conservation of native deer and camelids in Chile and other parts of
South America.

Keywords: anti-predator defense; prehispanic dogs; cursorial; ambush;
hunting; ethnography; evolution of prey behavior; predation strategy

\par\null

\textbf{Introduction}

The damage to biodiversity caused by feral dogs (\emph{Canis
familiaris)~}is well-established globally. Young et al. (2011), in a
global review of feral dog impacts, show that the negative effects of
\emph{C. familiaris} on native species is widespread. Feral dogs are a
serious problem for conservation, mainly because of their predation on
native species, transmission of diseases, and competition with other
predators, which may destabilize ecosystems (Ritchie et al. 2014; Hughes
\& Macdonald 2013). For example, on the Galapagos Islands a significant
decline in marine iguanas (\emph{Amblyrhinchus cristatus}) resulted from
the establishment of dogs on the island (Kruuk \& Snell 1981). Feral and
freely ranging dogs in Brazilian protected areas displace native
predators, spread diseases, and predate small and medium sized wildlife
(Lessa et al. 2016). South America is a hotspot of \emph{C.
familiaris~}negative impacts on threatened species (Doherty et al.
2017). Why exactly are dogs such a destabilizing conservation threat in
South America in particular? Geist (1998) partly addressed one angle of
this question when he concluded that the South American deer are not
adapted to cursorial predators like dogs, and that deer traits like body
size, locomotion, and reproductive investment, imply that there must
have been only ambush predators present during the radiation of the
South American deer species. In this paper, we re-examine and extend
this hypothesis, to ask whether it can explain why deer and also
camelids of South America are so vulnerable to feral dogs. We look more
completely at all the evidence for the hunting strategies of prehistoric
predators as well as domesticated canids associated with Precolumbian
human presence. We focus on the case of Chile, and we end our review by
evaluating the state of knowledge and suggesting hypotheses that need to
be researched in order to better understand how the evolution and
ecology of predator-prey interactions affect possible responses to the
conservation challenge posed by feral dogs.

\par\null

In Chile, feral dogs represent economic as well as environmental
damage.~\emph{C. familiaris} is estimated to cause 57,000 sheep deaths
annually (Bonacic \& Muñoz 2014), although it should be noted that this
includes deaths caused by both feral and domestic dogs allowed to freely
roam agricultural areas. Both feral and freely ranging human-dependent
dogs cause significant reductions in the populations of threatened
species in Chile. Freely ranging dogs that are poorly fed are more
likely to prey on and harass wildlife (Silva-Rodríguez \& Sieving 2011).
Rural dogs are interference competitors with the native
fox~\emph{Lycalopex griseus~}(Silva-Rodríguez et al. 2010), eat the eggs
of migratory birds that nest in Chile (Shuttler et al. 2009), and
predate on native deer species, including pudu~\emph{Pudu puda}
(Silva-Rodríguez et al. 2009; Silva-Rodríguez \& Sieving 2012), and
huemul~\emph{Hippocamelus bisulcus} (Flueck \& Smith-Flueck 2006; Corti
et al. 2010). Reports from Peru (Barrio 2007) state that fawns of
taruka,~\emph{Hippocamelus antisensis,} a deer also found in northern
Chile, may be killed by shepherd and feral dogs. The native camelids,
principally guanaco (\emph{Lama guanicoe}), are considered by experts
(pers. comm. WCS Chile Grupo Núcleo Guanacos 2020) to be particularly
susceptible to being killed by feral dogs, and dog presence is
considered to be a factor preventing the conservation and reintroduction
of~\emph{Lama guanicoe} in large parts of its historical range (Farías
et al. 2010; González 2010). According to McLaren et al. (2018), vicuña
(\emph{Vicugna vicugna} ), a high-altitude relative of the guanaco, are
hunted and harassed by humans with dogs, but feral dogs primarily
predate on young offspring. It should be noted that the extent of the
threat to species posed by dogs in Chile is partly due to the lack of a
legal or cultural basis for their control, lethal or otherwise.

However, there are also behavioural and evolutionary features of
predators to consider when trying to understand why feral dogs present a
large and difficult-to-control threat to Chilean wildlife. When
understanding predation, there are two key factors to consider: the
predator's niche, and its hunting strategy. In terms of their niches, in
general most canids (Carnivora: Canidae) are omnivores that do not kill
(but may scavenge) prey larger than themselves (Sheldon 1992). Small
canids \textless{} 6 kg are solitary foragers, canids 6-13 kg are
facultative cooperative foragers, and larger ones are obligate
cooperative foragers (Sheldon 1992) likely to be specialized as
carnivores, with some exceptions (Wang et a. 2004). Canids, like other
predators, may use either cursorial (pursuit) or ambush strategies. In
cursorial strategies, the predator relies on its running adaptations and
stamina to exhaust the prey, and may lack specific killing tactics,
rather relying on bringing the prey down by biting (Fox 1987; Sheldon
1992; González 2010). In ambush strategies, the predator stalks the
prey, remaining undetected until it can pounce on or rush at the prey
from a short distance; it may rely on specialized killing tactics (Fox
1987; Sheldon 1992). For example, puma (\emph{Puma concolor} ) are
ambush predators that kill with a specialized crushing bite to the head
(Sarno et al. 1999).

In turn, prey species have adaptations such as scanning for predators,
alarm calls, escape tactics, group defence tactics, and so on, which may
be suited for different predator body sizes, group sizes, and either
pursuit or ambush predation strategies (see Caro et al. 2004).
Essentially, to escape cursorial predators, prey need to outrun or lose
them. By contrast, the advantage of ambush predators is their ability to
take a targeted prey individual by surprise from a short distance. An
effective defence against ambush predation is ``predator inspection''
which consists of staring at, approaching, surrounding and/or following
the predator so that it cannot gain this advantage (FitzGibbon 1994
provides an excellent review). In a study of gazelle, FitzGibbon (1994)
found that they were much more likely to inspect ambush than cursorial
predators. Inspecting cursorial predators is likely to be risky and
maladaptive, as by shortening the distance to the predator it may
increase the chances of being overtaken.

In its feral state,~\emph{C. familiaris} hunts in packs using a
cursorial strategy (Farías et al. 2010; Ritchie et al. 2014). Geist
(1998) notes that all of the South American deer lack appropriate
defenses against what he calls ``culling'' predators, by which he
appears to mean a cursorial strategy where one or more predators picks
out and runs down its prey. Geist notes that South American deer are
``curious'' which suggests they approach possible threats. They also
respond too late to attacks, are saltatorial bounders rather than
runners, and lack stamina (Geist 1998). It has also been suggested that
guanaco lack adaptations to escape cursorial pack hunters
(Root-Bernstein \& Svenning 2016). The guanaco demonstrates group
vigilance and predator inspection, suitable for countering its main
native predator, the puma (\emph{P. concolor}), which hunts by solitary
ambush (Darwin 2015 {[}1839{]}; Marino \& Baldi 2008; pers. obs. MR-B).
It is not entirely clear whether guanacos inspect dog packs, but
González (2010) attributes high dog-caused mortality to guanacos' short
bursts of fast flight, leaving them unable to outrun dogs with greater
stamina. Oyama (2006) describes vicuña (\emph{V. vicugna} ) in the
Peruvian Andes as approaching possible threats such as dogs, and facing
predators while giving an alarm call. Similarly to the case of guanaco,
vicuñas' main predators, including puma and foxes, hide in the long
grass and ambush them.

If South American deer and camelids lack appropriate defense behaviours,
does this imply that this is an evolutionary problem---that they lack an
adaptive trait? Although prey must respond differently and appropriately
to different predators, this does not necessarily imply the evolution of
completely independent responses to each predator. Lind and Cresswell
(2005) make the valuable point that ``antipredation behaviours are a
composite of many behaviours that an animal can adjust\ldots{}'' (p.
945). Components of anti-predator behaviour can be lost under loss of
predation pressure, but the presence of any kind of predator may be
enough to retain a whole suite of antipredation behaviours for tens of
thousands of years (Blumstein 2006). So, to assess the hypothesis that
key native Chilean prey species---camelids and deer---lack evolutionary
adaptations to cursorial group predation, we would need to establish
that they never in the past experienced selection pressures in the form
of cursorial group hunting predators. Thus we need to understand when
cursorial group-hunting canids first appeared in the southern cone of
South America, which is not entirely clear. We focus on canids simply
because there are not, and have never been, any group-hunting or
cursorial felids in South America.

It is clear that European conquistadors brought domestic dogs to the
Americas starting around 500 years ago. However, the history of canids
in South America, and ecological and behavioural descriptions of any
kinds of prehispanic South American canids, have not been synthesized in
a way accessible to conservationists. Here, we review existing evidence
for the presence of prehispanic canids and their behaviours and
ecological roles, focusing on Chile and the southern cone of South
America.

\par\null

\textbf{Canids and their hunting ecology}

\emph{South American canids}

Up to 16 extinct canids (Wang et al. 2004), depending on the criteria
used for phylogenetic studies, inhabited South America since around 3
Mya. Canids originated in North America and spread first to Eurasia upon
the formation of the Bering Strait in the late Miocene (7-8 Mya), where
they radiated into multiple lineages. Later, at 3,5 Mya they also spread
to South America, in the Great American Biotic Interchange (GABI), which
however may have started as early as 20 Mya (see Bacon et al. 2015,
O'Dea et al. 2016, and Jaramillo et al. 2017). The GABI led to the
evolution of new canid species, of which at least 7 were hypercarnivores
(Prevosti \& Forasiepi 2018). Hypercarnivores are those species with a
diet composed of at least 70\% meat (Holliday \& Steppan 2004), and so
have unique ecological characteristics and requirements, distinct from
other species of the group. Particularly in Canidae, hypercarnivores are
large in size due to energetic constraints (Wang et al. 2004) and thus
also very likely to be obligatory cooperative group hunters (Sheldon
1992). The South American ancient hypercarnivores came from 4
genera,\emph{Theriodictis} ,~\emph{Protocyon} ,~\emph{Speothos,}
and~\emph{Canis,} the latter having three species\emph{, C. gezi, C.
nehringi, and C. dirus} (the dire wolf, also found in North America,
although it has been recently included by Perri et al. (2021) in the
genus~\emph{Aenocyon}, according to molecular data) (Wang et al. 2004;
Prevosti 2010). Members of these genera all went extinct at the end of
the Pleistocene, except the bush dog (\emph{Speothos venaticus}) which
is a small, short-legged group-hunting carnivorous canid living near
water in the tropics. The hypercarnivorous Pleistocene species,
distributed as far south as Argentina (Prevosti 2018), would most likely
have hunted megafauna.~\emph{Protocyon troglodytes,} a group-living
cursorial predator, as well as the non-hypercarnivorous
canids~\emph{Dusicyon} spp in Uruguay hunted camelids, giant sloths, and
other large and medium sized prey (Prevosti et al.
2009).~\emph{Dusicyon} spp. went extinct much later than the
hypercarnivores (see below).

The 13 contemporary South American canids are divided into two lineages
(Perini et al. 2010), one of species similar to Old World foxes
(genera~\emph{Lycalopex}, \emph{Atelocynus}, and \emph{Cerdocyon} ), and
the other of species similar to Old World wolves (\emph{Chrysocyon and
Speothos} ). Although it is unclear, some authors (see Austin et al.
2013; Slater et al. 2009) consider this last lineage to be related to
the extinct genus \emph{Dusicyon} mentioned previously.

\emph{South American canids in Chile}

The only one of the now-extinct South American Pleistocene canids known
to have had a distribution in Chile was~\emph{Dusicyon avus} , found in
Chilean Patagonia and Tierra del Fuego, on the southern tip of Chile
(e.g. Méndez et al. 2014; Prevosti et al. 2009). The
genus~\emph{Dusicyon} , to which~\emph{D. avus} and~\emph{D.
australis~}(also known as the Falklands wolf) belonged, went extinct
fairly recently, although the extinction process may have been initiated
by reductions in its distribution begining in the Pleistocene (Prevosti
et al. 2011; Austin et al. 2013; Prevosti et al. 2015). In spite of
this, there have been reported sightings of~\emph{D. avus} by English
explorers as late as 1870. There are also theories
that~\emph{Dusicyon~}spp\emph{.} underwent selection or hybridized with
European~\emph{C. familiaris} and thus became unrecognisable but not
necessarily extinct (Borrero 2009). The English natural historian
Charles Hamilton Smith wrote in 1839 that
mainland~\emph{Dusicyon}~\emph{spp.} were tamed and used for hunting by
South American tribes (quoted in Clutton-Brock 1977).~\emph{D.
australis,} restricted to the Falklands Islands, and extinct by the
1880s, shows morphological traits typical of domesticated canids, such
as a bulbous head and white patches on its pelage (Clutton-Brock 1977),
all of which may be a consequence of a long interaction with humans. As
humans arrived in the Americas at the end of the Pleistocene, and in
southern Chile around 14.5 kya (Gómez-Carballa et al. 2018; Braje et al.
2020), this gives any~\emph{Dusicyon} -human relationship considerable
potential historical depth.

The other South American Pleistocene canids, especially the
hypercarnivorous species, are not known to have occurred in Chile
(Carrasco O. 2009). Since the uplift of the Andes occurred around 14 Mya
(Le Roux 2012), well before the GABI, it may have formed a barrier to
their dispersal. Other Pleistocene carnivores, including \emph{Smilodon
populator} , \emph{Panthera onca} and \emph{Arctotherium tarijense} are,
like \emph{D. avus} , found only in the extreme south of Chilean
Patagonia (Carrasco O., 2009) where large parts of the Andean cordillera
are below 1000 m and thus less of a barrier to dispersal. Indirect
evidence suggests that they also travelled further north into northern
Patagonia at least (Labarca et al. 2014). Other large predators
(\emph{P. concolor}) were (and are still) found in central Chile
(Nielsen et al. 2015). New evidence could change known distributions:
part of a skeleton of a large canid has been found in the Pampa de
Tamarugal (Region of Tarapacá), undated but conjectured to be from
before the arrival of the conquistadors to Chile (C. Latorre pers. comm.
2018, N. Villavicencio pers. comm. 2020).

Currently, three native canids exist in Chile, all of the
genus~\emph{Lycalopex} (\emph{L. culpaeus} , \emph{L. griseus} , and
\emph{L. fulvipes} ). These species are omnivores, consuming small prey
(Iriarte 2007). They have also been reported to opportunistically hunt
juvenile guanacos (Novaro et al. 2009) in the Karukinka National
Reserve, in Tierra del Fuego, where there are no pumas and the
population of guanacos is high. According to Novaro et al. (2009),
culpeo foxes~\emph{L. culpaeus} are cursorial predators. Juvenile
guanaco, though much larger than culpeo foxes, are recorded to have been
chased by them in two cases, and in at least one case, the attacking fox
was kicked by several adult guanacos (Guzmán 2009). \emph{Lycalopex
culpaeus} also prey on huemul (\emph{Hippocamelus} \emph{bisulcus} )
fawns (Corti et al. 2010). It is thus likely that \emph{Lycalopex} can
exert a cursorial hunting selection pressure on juvenile large prey
species, but since juveniles are often protected by adults who may
attack the fox, this may mitigate the selection pressure. Juvenile
anti-predator responses might also be adaptations distinct from adult
behaviors. Franklin \& Powel (1994) report that guanacos as well as
llamas respond to coyotes by alerting to them, alarm calling, walking or
running towards them, chasing them, kicking and stomping on them.

\emph{South American dogs}

Though~\emph{C. familiaris,} which co-evolved along with humans in
Eurasia, is found early in the archeological record of North America,
probably as a result of Bering Strait crossings, they arrive much later
in South America. Dogs are reported from 4,5 kya in Ecuador and the
coast of Peru (Stahl 1984; Salomon and Stahl 2008; Stahl 2012), and up
to 10 kya in other parts of the Americas (Allison et al. 1982; van Asch
et al. 2013, Perri et al. 2018).

It seems most probable that any dog present in Chile arrived along
migratory or trade routes from the Andes, with the earliest evidence of
dogs in the form of artistic representations from cultures such as the
Moche (1900 ya -- 1300 ya) or the Chimú (1100 kya -- 500 ya) (Vásquez
Sánchez et al. 2009). Prates et al. (2010) hypothesize that the presence
of dogs in South America was mainly related to complex societies, such
as the Andean societies of Peru and Ecuador previously mentioned, and
that their arrival in the Southern Cone is related to an increase in
long-distance communication and trade by egalitarian hunter-gatherer
societies of the Pampas and Patagonia. The authors come to these
conclusions based on archaeological deposits in northern Argentina
dating from around 1000 ya (around 500 years before the arrival of the
Spanish), which accords with other archaeological evidence of early dog
presence in southern Brazil (1600 ya, Guedes Milheira et al. 2017) and
in Uruguay (undated remains, but found in multiple-use structures
starting around 1000 ya, López Mazz et al. 2018).

Many morphotypes of dogs may have been present in South America at
different times. Gilmore (1950) lists nine possible breeds of South
American dogs, remarking that parallel selection pressures or
morphological constraints resulted in a terrier-like dog (the Fuegian
dog), a setter-like dog (the Ona dog), a foxhound/greyhound type dog
(the Tehuelche dog), and another terrier-like dog (the Techichi dog), as
well as hairless dogs. The Pre-Columbian origins of these possible
breeds is unknown. Van Asch et al. (2013) confirmed the pre-Columbian
origins of several formally recognised American breeds, including the
Peruvian \emph{perro sin pelo} (``hairless dog''). Vásquez Sánchez et
al. (2009) summarize many possible morphotypes of South American dogs,
including a short-nosed dog from the Chicama valley, the medium-sized
``helping dog'', the ``pet'' dog with a long body and short limbs, the
miniature dog similar to a chihuahua, the hairless dog, a short-haired
dog, a Peruvian bulldog, a Peruvian sausage dog, a long-haired Incan
dog, and the Chiribaya shepherd used to herd llamas. Vásquez Sánchez et
al. (2009) also discuss dogs depicted on Moche pots, including a
morphotype consisting of a small or medium-sized spotted dog with a
bulbous head, which appears in a deer-hunting scene among other
contexts. They speculate that this kind of dog might have been used to
corner deer during hunting. Like the Peruvian pitbull (Cossios E. 2018),
many morphotypes may have been bred locally for certain periods of time,
but later allowed to outcross with other morphotypes.

It is unclear whether prehispanic dogs of South America would have
formed significant feral populations, although large packs of feral dogs
are reported by the 18\textsuperscript{th} century (Prevosti et al.
2015), and it is hypothesized that feral populations existed soon after
the beginning of dog domestication in Eurasia (Boitiani \& Ciucci 1995).
Dogs, when feral or free ranging, are omnivorous opportunists (Campos et
al. 2007). Dogs that are fed by humans tend to predate much less on wild
prey (Vanak \& Gompper 2009). This indicates that, if there were either
feral or free-ranging dogs in South America before the arrival of the
Europeans, the pressure against succeptible~wildlife may have a
reasonable historical depth, although maybe not enough to allow prey to
evolve adequate anti-predator behaviours.

\emph{South American dogs in Chile}

Chile can be grouped into three large cultural areas: the North, with
Incan influence, the Central-South zone, dominated by the Mapuche
peoples, and the Southern region, associated with coastal
hunter-gatherer cultures, which includes Patagonia. We discuss South
American dogs in Chile in the sequence of these three cultural areas
from north to south. However, the evidence for prehispanic dogs in each
zone is relatively weak.

In Arica (northern Chile), eight mummified dogs, dated to circa 1450-
500 ya, were found by Allison et al. (1982), belonging to the San
Miguel, Inca, Cabuza, Alto Ramirez and Maitas Chiribaya phases and
periods. These are terrier-like dogs 46-52 cm high, apparently similar
in form to the Moche hunting dogs discussed by Vásquez Sánchez et al.
(2009).

It is commonly held in Chile, as well as recorded by historians (Vial
2010) and the ethnographer Ricardo Latcham (1822) that the contemporary
Mapuche indigenous people of the center-south of Chile previously had
two words for one or more kinds of dogs that they had:~\emph{thegua}
or~\emph{tregua} and~\emph{munutru} . Mapuche dogs may have been
obtained from Argentinian Patagonia pre-Columbian populations (Prates et
al. 2010), or from the Inca (Uribe \& Sánchez 2016). According to
Latcham (1822, p. 62),~\emph{munutru} refers to a dog with long curly
hair on its face, or any ``small ugly dog'', similar to what is today
called a ``\emph{quiltro} '', usually translated as ``mutt'', (ibid p.
62), and is reported by contemporary Mapuche people to mean anything
with an ``ugly face'' (Loncon, E. pers. comm. 2019), while~\emph{tregua}
simply means ``dog.'' Some contemporary dogs in areas of central Chile,
where indigenous populations persisted for several centuries after
colonization, are ``small ugly dogs'', with bulbous foreheads and eyes,
and short floppy ears, but no curly hair (pers. obs. MR-B). Perhaps
these resemble the munutru or another indigenous dog morphotype.
However, no specimens of Mapuche dogs are known to us either in the
present day nor the archeological record.

The case of Patagonia in the extreme south of Chile also does not have
good biological evidence about dogs. Ethnohistorical records of the
19\textsuperscript{th} and 20\textsuperscript{th} Century (Coiazzi 1914)
mention the presence of dogs in societies that had little or no contact
with the Spanish or other colonists. Coiazzi documents the coexistence
of~\emph{C.~familiaris} with hunter-gatherer societies of southern
Patagonia, in particular the Selk'nam (also known as Ona), who used dogs
to hunt guanacos among other animals. Martin Gusinde, famous Selk'nam
etnographer, also recognized the presence of, at least in some form,
domesticated dogs in Selk'nam encampments. These dogs, described as
loud, aggressive, and with a pointed snout (Gusinde 1951), were very
appreciated by the natives for their loyalty and protection, and as
Gusinde described ``In each house there are at least four of these
acrimonious and irreconcilable dogs''. Gusinde also described Selk'nam
using dogs to hunt guanacos, even apparently picking up and following a
scent in packs, just like hunting scent hounds in the UK and other areas
with a tradition in hunting (Gusinde, 1951). The initial Spanish
colonization of Chile only extended to Chiloé, which does not rule out a
European origin of Selk'nam dogs via long-distance trade, but also makes
it plausible that these dogs had a previous origin in pre-Columbian
trade networks or migrations. Coiazzi (1914) suggests that the Selk'nam
dogs had displaced the populations of a native canid similar to a fox,
though whether he means by competition (cf. Vanak \& Gompper 2009) or as
a favoured domesticate is unclear. Carlos R. Gallardo (1910) also wrote
about the relationship between Selk'nam people and domesticated,
supposedly native, dogs, described as ``\emph{Canis}
(\emph{Pseudalopex}) \emph{lycoides'',} also described as an important
part of guanaco hunting. These domesticated dogs are described
thouroughly in his book ``The Onas'':

\begin{quote}
``The fuegian dog is a not very big, wild looking animal. Some of them
retain such a striking similarity to their ancestors, one can easily
mistake them for a big fox, but (\ldots{}) There are some of yellowish
grey colour, of clear and almost white background, and with dark tints
from black to ashen yellow. They have a broad forehead, straight,
pointy, and fairly large ears, the eyes are somewhat oblique, the snout
is long and even pointy, the neck is short, and the legs are notable for
having very developed membranes between the fingers. The tail is long,
covered in also long hair that coats the rest of the body as well.''
\end{quote}

Some of this description can be seen in Figure 1, from the same book
(Gallardo 1910). In this context and with such characteristics, the
possibility of a native canid being domesticated in Tierra del Fuego by
Selk'nam is certainly more plausible. In addition to this, the study by
Petrigh and Fugassa (2013), in which they genetically identified a
taxidermized canid belonging to native people of the area showed that
this specimen was closely related or identical to~\emph{L. culpaeus.}

\emph{Other, native, domesticated canids}

Mitchell (2017) proposes a theory to explain the slow dynamics of spread
of \emph{C. familiaris} in South America, including the Southern Cone
and Chile, after its original arrival across the Bering Strait. He
argues that the canids of South America were better adapted to local
diseases such as l\emph{eishmaniasis canina} and distemper than \emph{C.
familiaris,} and acted as reservoirs and vectors that led to almost
epidemic levels of these diseases in dog populations, limiting their
adoption by native peoples. The Andes may also have acted as a barrier
to dog migration into Chile. An alternative perspective is developed by
Prates (2014), who argues that many societies may have failed to
incorporate C\emph{. familiaris} into their social practices and daily
lives if other canids already played roles such as keeping watch,
hunting, or companionship. Prates bases this hypothesis on the discovery
of a \emph{D. avus} skeleton at a funerary site of the ``Loma de los
Muertos'' archeological deposit in Río Negro Province, on the southern
half of Argentina, dated around 2000-3000 ya (Stahl 2012, 2013; Prates,
2010). This is most probably a deliberate burial. In the context of
cosmological systems in many areas of South America in which taming of
individual animals is a common practice and a fundamental part of their
understanding of social relations between species (Stahl, 1984; Erikson,
2000), this suggests the hypothesis of possible taming of canids in many
societies of the continent. It is thus not impossible to imagine that
some sort of domestication or taming of native canids by Patagonian
hunter-gatherers happened in Chile (Gusinde, 1951; Gallardo, 1910).

\emph{C. familiaris} was thus not necessarily the only canid either
tamed or domesticated by humans. However, further research would be
necessary to determine what kinds of relationships may have existed
between native canids and humans in Chile, and to what extent this
domestication took place, as at present we do not have archeological nor
paleontological data to support these hypotheses.

\emph{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~\emph{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,~\emph{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~\emph{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.~\emph{Canis lupus} ,~\emph{C. latrans} ,~\emph{C. adustus}
,~\emph{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.

\emph{D. avus} was the only large Pleistocene canid found in Chile
(Castillo, 2007). Prevosti et al. (2009) report that~\emph{D.
avus~}likely predated~\emph{Lama} spp. as well as other large and medium
animals, although they also indicate that~\emph{Lama} spp. may often
have been scavenged rather than killed (Prevosti \& Vizcaíno 2006;
Prevosti \& Martin 2013). The extinct~\emph{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 (\emph{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~\emph{Dusicyon} was open plains, so it could have shared a similar
hunting strategy. Yet, as for~\emph{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,~\emph{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~\emph{Dusicyon} spp (Austin, 2013). While it has never
occurred in Chile to date (Torres et al. 2013), it would have coexisted
with~\emph{Lama} spp., rheas (\emph{Rhea} spp\emph{.} ), and other prey
species whose ranges included Chile during the Holocene. It can thus
both suggest potential~\emph{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 (\emph{Ozotoceros bezoarticus} , 22-34 kg) and Greater rheas
(\emph{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 (\emph{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~\emph{D. avus} .

The surviving Pleistocene \emph{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 \emph{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, \emph{Speothos venaticus} ,
which are only able to bring down deer by biting them in large packs
(Biben 1982; Sheldon 1992)). Otherwise, \emph{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.

\par\null

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 \emph{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.

\par\null

\textbf{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 \emph{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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\begin{figure}[H]
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\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

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Figure 1. Reproduction of a drawing found in ``The Onas'' by Carlos
Gallardo (1910) titled ``Educating the dog'' (\emph{Educando al perro}
). Some characteristics mentioned in the text can be seen in the drawing
of this Fueguian dog. The image is free of copyright
(http://www.memoriachilena.gob.cl/602/w3-article-8403.html)

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