Most ungulates are surprisingly inept at using their horns against predators

Most species of ungulates have pointed 'weapons' of some kind on their heads, be these horns, antlers or long canine teeth.

These are used in masculine rivalry. In species in which they occur also in females, they are used for other social threats in certain situations.

One might expect, then, that the horns, antlers and teeth of ungulates could be applied to stabbing predators.

Surprisingly, most species of ungulates turn out to be inept in fighting off predators. The pointed structures on their heads function far more as adornments than as weapons; and their ritual rather than violent deployment is analogous with the gentlemanly sport of fencing.

The problem seems to be that the 'hardware' lacks suitable 'software', i.e. the brains of ungulates do not seem to be programmed for aiming and striking at predators, regardless of how precise their use is in masculine sparring.

Ungulates generally rely on rapid fleeing and reproduction, not self-defence, in order to survive predation. When cornered, they lack the behavioural versatility to use their 'weapons' in more than a tokenistic way. Once wounded, they go into shock and seem to lose will and coordination.

The few exceptions appear to include

All of the above are ecologically and/or socially extreme in ways suggesting that fleeing and reproduction alone would not allow them to survive predation. And all have horn-designs unusually suited to stabbing and hooking.

Other ungulates with defensive reputations in the semi-popular literature, such as oryxes (https://www.inaturalist.org/observations?taxon_id=42307), are more inept than the fearsome shapes of their horns might suggest.

The African savanna buffalo (Syncerus caffer, https://www.inaturalist.org/taxa/42405-Syncerus-caffer) is certainly dangerous to humans when wounded. However, its horns are too blunt to stab an antagonist.

Various video-clips show how timorously the African buffalo behaves versus the lion (Panthera leo), even when there are opportunities to strike blows (see https://www.sabisabi.com/blog/14593/lions-vs-buffalo/ and https://www.youtube.com/watch?v=zzI8ya3ARWM and https://www.youtube.com/watch?v=SObYT06hnm4 and https://www.youtube.com/watch?v=MY6conyVLlc and https://www.youtube.com/watch?v=aCwwecrasWI). The lion is intimated more by the bulk and gregariousness of buffaloes than by the horn-tips.

The following video-clip (https://www.youtube.com/watch?v=Bu2JspPdMy0 ), taken near Kruger National Park, shows:

  • how obtuse the African buffalo can be, with the mother wandering out from the group with her infant, 'just asking for trouble',
  • that the mother does in fact put up a spirited defence against one adult male of the lion, but loses her infant anyway because several individuals of the lion are hunting together, and
  • how poorly co-ordinated the collective defence is: there are several mature males near the infant, but they do not cooperate to defend it.

And even when the predators, in the act of eating, are vulnerable to retribution, the mature male African savanna buffalo holds back, as if knowing that if several individuals of the lion grab him, he will not be able to rely on solidarity from the rest of his own group.
 
Furthermore, here is a point that I suspect will be overlooked by most naturalists: the mother, when attacked by the lion, does not flee into the nearby water.
 
If this were the river buffalo (Bubalus bubalis) or the water buffalo (Bubalus carabanensis), I expect that the mother would take refuge in the river immediately.

I realise that in plain view was a group of the hippopotamus (Hippopotamus amphibius), and that species would not necessarily show any solidarity with a buffalo. However, this footage suggests that the African savanna buffalo is quite different from its tropical Asian counterparts in its relationship to water. It obviously sees no refuge in the river.

Not only waterbucks (Kobus ellipsiprymnus and Kobus defassa), but even antelopes such as Strepsiceros strepsiceros and Hippotragus niger that have no particular aquatic affinity, would, I expect, have gone straight into the water to save themselves (http://www.dailymail.co.uk/news/article-2609382/Kudu-tell-African-antelope-escaped-pack-hyenas-Clever-prey-conceals-pack-zebra.html).

It is in the light of this general pattern, in which most ungulates are almost as vulnerable to predators as they would be were their horns absent, that we can understand the basic nature of Spanish bullfighting: choreography rather than a real contest of impalation. The matador pierces the bull, which is incapable of reciprocating - except by accident.

Publicado por milewski milewski, 17 de mayo de 2021

Comentarios

The hook-lipped rhino (Diceros bicornis) is certainly defensive, to the point of seeming aggressive. However it too tends to be inept in actually striking blows, partly owing to incongruously poor eyesight.

Publicado por milewski hace más de 1 año (Marca)

Interesting. The elk sometimes will use antlers to try to fend off wolves, but they aren't well equipped to fight off a whole pack very well.

Publicado por beartracker hace 5 meses (Marca)

The difference between Carnivora and their prey (e.g. ungulates) is not so much weaponry but skill, i.e. a behavioural programming to use weaponry to lethal effect./It’s true that Carnivora have sharp teeth and sharp claws. But these organs are, in themselves, no more potentially dangerous than the organs possessed by most ungulates and many other herbivores: horns, antlers, hooves, and even teeth (e.g. in pigs and rodents). If anything, it is the herbivores that have the more impressive weapons in the sense of sheer mass and force of weaponry. All the teeth and claws of a lion add up only to a small fraction of each horn of an African buffalo, even if we look at the female of the bovine. And even a cane rat has such formidable teeth that, if used well, these could really hurt any attacker.

The real difference is not weaponry as such; it is the ability, willingness, eagerness, skill, and versatility to use this weaponry.

Any intact adult human has fists, elbows, feet, knees, a forehead, teeth, and sheer weight. These count for little in a pusillanimous person, unpractised in fighting, who has not learned how to use these weapons and who has not adopted a habitually defensive attitude. But with the right training and attitude, even a small woman can inflict lethal harm on another human in a fight.

Consider the difference between a channel-swimmer and one who drowns in a pond: same human physique, but incomparably different skills in using the arms, legs, breathing, co-ordination, etc. The human species can swim well, not because we have ‘swimming organs’ but because we use what organs we have to excellent effect if trained and ready.

Many herbivores do have remarkable ‘fighting’ skills, or at least sparring skills, and the males practise hard to master their ‘martial arts’, which tend to be species-specific because the purpose of the antagonism is to gain mating rights. So, for example, many antelopes and deer spar with their horns/antlers, and it may take half of the life of the male to perfect his skills to the point that he can use his weaponry well.

So it is not just the case that herbivores have opted to abandon any thought of fighting and just get on with what they are good at, i.e. eating and breeding and growing rapidly. They do ‘fight’ and they do practise their fighting. It is just that all their martial arts seem to count for little when they are attacked by a predator.

Publicado por milewski hace 5 meses (Marca)

 As part of the same explanation, I can point out a consistent difference between carnivores and herbivores. Whereas the herbivores gain independence as soon as they are weaned, it is normal for carnivores to continue to depend on their mother for years. This is because a leopard or cheetah, once weaned, can still not hunt for itself until it has received enough lessons from its parent and practised enough. Herbivores have no such lag because there is so little to learn about eating plants. My point is that the superior deployment of weaponry by carnivores is consistent with their ‘schooling’ period between weaning and independence; this is their time to learn ‘martial arts’.

Of course, it is true that much sparring and even fighting among male herbivores is ‘ceremonial’ rather than intended to be lethal. Even the mutual ramming by males of the bighorn sheep, which muster impressive force, do no more than stun, for which the skull is adapted with well-developed shock absorbers. But on the other hand many sharp-horned bovids do have ‘skin-shields’ on their necks, shoulders and flanks, to reduce the risk of being impaled.

The only ungulates that deter predation by means of their weaponry seem to be certain suids, e.g. Potamochoerus larvatus. It seems significant that the canine teeth of bushpigs, so dangerous to predators, are not even visible - being hidden inside the mouth. Even in suids, the weaponry is partly in compensation for relatively short legs and small eyes.

Publicado por milewski hace 5 meses (Marca)

Great point about how the diet affects this. Eating plants doesn't take much time to learn, but hunting does take time to learn well.

Publicado por beartracker hace 5 meses (Marca)

@beartracker @davidbygott @maxallen

Example of reluctance of wildebeest to use horns even when determinedly defending infant against African hunting dog:
  
The following video clip (https://www.youtube.com/watch?v=cZUBC1SSWN8) is interesting w.r.t. the parental defence by a mother of the western white-bearded wildebeest (Connochaetes mearnsi) of its infant against a group of the African hunting dog (Lycaon pictus).

This individual ostensibly succeeds on this occasion. However, I found two aspects of this footage particularly interesting.
 
Firstly, the mother seems remarkably unwilling, or unable, actually to deliver blows with her horns. It is merely the threat of this that eventually wins the day against the predators. However, it might have been more effective to actually inflict wounds.
 
Secondly, I was surprised to see that, as the mother and infant escape the scene, the mother style-trots for for most of the way, as if to say to the dogs ‘I’m still fit, so if you try again you’ll fail again’.

This post-crisis ‘stotting’ is new to me. It is particularly interesting because wildebeests (at least of other spp.) also use a different fitness-display, that of ‘prancing/cavorting’. This is typically associated with the black wildebeest (Connochaetes gnou), but also occurs in other spp. (https://www.flickr.com/photos/130290763@N08/50155063702/ and https://www.shutterstock.com/nb/video/clip-5778965-cavorting-animal-mad and https://www.youtube.com/watch?v=K6fQrirdP2w and https://www.warrenphotographic.co.uk/23373-wildebeest-capering).

I would like to understand better why, in antagonism with predators, wildebeests prance/cavort in some situations, and style-trot in others.

Also see https://blog.londolozi.com/2021/01/26/wildebeest-are-anything-but-stupid/.

Publicado por milewski hace 5 meses (Marca)

Although Oryxes are inept at self defense, it’s apparent that Sable (Hippotragus niger) have highly efficient defenses/bluffs

Publicado por paradoxornithidae hace 4 meses (Marca)

https://youtu.be/WAPh8Bch5xU
In this video, the H. niger doesn’t use it’s horns directly so to say, but keeps distance by charging and displaying horns to the L. pictus pack, whilst being in the water. Combing multiple means of escape, and deterrent.

Publicado por paradoxornithidae hace 4 meses (Marca)

According to Valerius Geist, even Oreamnos americanus hastily quits the area when a group of the wolf arrives (https://www.youtube.com/watch?v=TBFGlCipnqo). This is remarkable, because this species a) can evade predators by means of its sure-footedness on cliffs, b) is renowned for its intimidating horns, and c) usually seems more phlegmatic than other ungulates in the proximity of carnivores.

Publicado por milewski hace 4 meses (Marca)

Many thanks to @paradoxornithidae for this exceptionally informative video.

@beartracker @maxallen @davidbygott @tonyrebelo @jeremygilmore @tandala @dejong @capracornelius @oviscanadensis_connerties

Please note the following.

Firstly, this footage shows a behaviour that may not have previously been recorded. This is the rapid and continual horizontal wagging of the tail by Hippotragus niger in a stand-off with Lycaon pictus.

The use of the tail as a caudal flag in Hippotragus is poorly documented in the literature. What is significant in this case is that the horizontal wagging of the tail differs from the erection of the tail seen in various other ungulates when alarmed. Furthermore, the horizontal movement means that the caudal flagging fails to create a 'strobing' effect against the white patch on the buttocks, even if viewed from behind.

Also note another behaviour possibly never recorded before: as the individual of H. niger lowers its horns in an almost choreographed way, it simultaneously utters a loud, protracted snort - different from the normal snort of alarm, and perhaps having a vocal component.

So, the defensive display consists of a) the dynamic component of the lowering of the horns, b) the dark-pale contrast presented by the face, which is a kind of 'frontal flag' in terms if adaptive colouration, and c) the audial accompaniment of the specialised snort.

(Another bovid with a combination of snort and vocalisation, 'warning off' intruders, is Connochaetes gnou, https://www.facebook.com/watch/?v=870370330095038.)

Please note that, although H. niger does lower its horns defensively, the main form of counter-intimidation is the flailing of the fore feet while feinting forward (similar to giraffes in similar situations).

It is significant that even a hippotragin bovid, belonging to a clade somewhat renowned for the anti-predator use of the horns, seems to advertise its capacity to kick forwards, in this emphatic way. The implication is that the risk posed by the horns has been overstated in the literature.

It seems significant that this adult male individual of H. niger, when harassed by a group of L. pictus, ran down to a pool of shallow water. Was this because it sought the option of retreating even further, into deeper water, were the carnivore to persist with the attack? Was the idea to use Hippopotamus - which tends to be irascible in an unintelligent way - as a form of distraction (collateral defensiveness)? Was the idea merely to find a situation as much out in the open as possible (as opposed to backing into a patch of thorny cover to protect the hindquarters, and then fight with the horns)? Was this individual somewhat 'torn' between an attraction to water as a refuge, and fear of Crocodylus niloticus?

On the topic of caudal flags, please note the various gradations of caudal flagging in L. pictus, ranging from a) inert tail while standing attentively but stationary, or while stalking with head lowered, through b) erratic flicking of the lowered tail to one side when walking/trotting indecisively, to c) raising the tail above the horizontal when either breaking into a run (however brief the acceleration), or approaching the prospective prey dangerously closely.

Lycaon pictus is a prime example of a carnivore possessing a caudal flag (https://www.dreamstime.com/african-wild-dog-lycaon-pictus-namibia-image198639762 and https://www.dreamstime.com/african-wild-dog-lycaon-pictus-pack-kill-greater-kudu-female-carcass-namibia-image196305261). The above video reveals low-intensity aspects of this, and suggests that the various positions of the tail are complex enough to form a rudimentary 'sign language' in L. pictus. The wolf does not show similar caudal flagging, and the dhole needs further investigation.

Also note that the individual of Hippopotamus (which is adult male, based on the proportional size of the head) has an inert tail as it stands still, watching the drama unfold.

Publicado por milewski hace 4 meses (Marca)

While we are on the topic of Hippotragus niger, https://www.youtube.com/watch?v=Le7MPaVDDhU contains some relevant points, particularly a defensive formation of gregariousness surrounding infants.

However, what this commentator does not mention, is that infants/juveniles tend to lag behind the group during locomotion, exposing themselves to predation (Estes 1991).

Note that, in this population in Texas, many adults seem to have somewhat deformed horns.

Publicado por milewski hace 4 meses (Marca)

Estes (1991), on page 119, states: "Though horse antelopes are considered by hunters to be among the most dangerous antelopes, and even lions are sometimes fatally stabbed, their dangerous horns and stabbing technique do not deter predators from hunting them and are probably rarely effectively employed (Eloff 1964)...The habitats frequented by these antelopes usually have relatively few predators, perhaps permitting a certain slackness in their antipredator behaviour".

By 'horse antelopes', Estes means hippotragin bovids.

Publicado por milewski hace 4 meses (Marca)

Interesting post!

Publicado por beartracker hace 4 meses (Marca)

@tonyrebelo @davidbygott @maxallen @beartracker @ludwig_muller @paradoxornithidae @botswanabugs @jeremygilmore @simontonge @tandala @oviscanadensis_connerties

Bos grunniens vs Panthera uncia:

The following
(https://www.youtube.com/watch?v=APuua4Gci5U) seems fairly typical of the ineptness of anti-predator defence in ungulates.

The mother of the attacked infant does eventually succeed in saving its offspring from predation. And it does score one impressive hit, in which it uses the horns to lift the predator off the ground, possibly inflicting internal injury.

However, a) it uses its horns clumsily despite these horns being more suitable-looking for such defence than are the horns of most other bovines, and most other ungulates, b) it fails even to try to use its superior weight to trample the predator, and c) the other adults in the group, although attending excitedly, fail to take obvious opportunities to horn, kick, or trample the predator.

Based on several similar video-clips I have watched in the case of Syncerus caffer vs Panthera leo, I find the two spp. of bovines similar in their 'dumb-looking' behaviour in these situations, despite a) the reputation of the African savanna buffalo for being a formidable adversary, and b) the fact that the domestic yak is not a wild species (https://www.inaturalist.org/taxa/889089-Bos-grunniens).

Publicado por milewski hace 4 meses (Marca)

@botswanabugs Many thanks for posting that informative video.

Publicado por milewski hace 4 meses (Marca)

@beartracker @maxallen @davidbygott @tonyrebelo @jeremygilmore @tandala @dejong @capracornelius @oviscanadensis_connerties @paradoxornithidae

The following shows how timid and tentative even Oryx gazella is vs predators, even in the case of a mother, and even vs a foe as small and weak as Acinonyx jubatus: https://www.youtube.com/watch?v=sMKkkW6jbug.

Publicado por milewski hace 2 meses (Marca)

@milewski have you seen this article ?
Dying for Dinner: A Cheetah Killed by a Common Duiker
Illustrates the Risk of Small Prey to Predators
Author(s): https://orcid.org/0000-0003-2702-5200Graham I.H. Kerley
Source: African Journal of Wildlife Research, 48(2)
Published By: Southern African Wildlife Management Association
https://doi.org/10.3957/056.048.024001
URL: http://www.bioone.org/doi/full/10.3957/056.048.024001

Publicado por botswanabugs hace 2 meses (Marca)

@botswanabugs Many thanks for this article, which is new to me.

Publicado por milewski hace 2 meses (Marca)

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