Why is there a caudal flag in the leopard (Panthera pardus) but not the puma (Puma concolor) or the jaguar (Panthera onca)? part 1

The leopard (Panthera pardus), the puma (Puma concolor) and the jaguar (Panthera onca) are like-size ecological counterparts on different continents. All have long tails, which they tend to twitch when excited.

However, the leopard alone has a caudal flag, which is sometimes activated while walking in daylight. And it is an extreme example of a caudal flag, even compared to deer and other ruminants (see https://www.youtube.com/watch?v=sWo7BiyDT04 and https://www.youtube.com/watch?v=LoVG75u72SQ).

The conspicuousness of the leopard's tail is owing to its white distal underside, which is exposed by curling the tail up to various heights.

For illustrations, please see:
https://www.alamy.com/leopard-panthera-pardus-tail-botswana-image184121172.html and https://www.alamy.com/leopard-panthera-pardus-side-body-face-looking-back-eyes-long-whiskers-tail-curled-up-partially-hidden-by-grass-samburu-national-reserve-kenya-africa-image242458468.html and https://www.alamy.com/leopard-panthera-pardus-walking-on-savanna-masai-mara-national-reserve-kenya-africa-image401804791.html and https://www.alamy.com/stock-photo-leopard-panthera-pardus-in-the-savannah-masai-mara-kenya-123182747.html and https://www.alamy.com/african-leopard-panthera-pardus-pardus-adult-male-at-dusk-going-to-a-waterhole-kruger-national-park-south-africa-africa-image243578242.html).

The puma has a tail of similar dimensions. However, its inconspicuous tip is not curled up while walking (https://www.youtube.com/watch?v=2bqgh40e6hU and https://www.mindenpictures.com/stock-photo-patagonian-puma-puma-concolor-patagonica-female-walking-over-open-naturephotography-image90744095.html and https://www.catster.com/the-scoop/see-mountain-lions-or-pumas-in-chile-patagonia and https://www.alamy.com/cougar-in-the-snow-5-image188075869.html).

The jaguar, although belonging to the same genus as the leopard, has a relatively short tail with an inconspicuously dark tip (https://twitter.com/Joe_LoCicero/status/957691134238576640/photo/1 and https://www.youtube.com/watch?v=flVdCHC1v-A and https://www.inaturalist.org/observations/39862424 and https://www.alamy.com/stock-photo-jaguar-walking-in-a-shady-area-of-the-riverbank-of-the-cuiaba-river-136707647.html).

Why is the leopard the species with a showy tail?

On their American supercontinent, the puma and the jaguar are the top predators in their main habitats.

By contrast, the leopard coexists with the tiger (Panthera tigris, in Asia), the lion (Panthera leo, in Africa and part of Eurasia) and/or the spotted hyena (Crocuta crocuta, in Africa and formerly in extratropical Eurasia) - all of which outrank it in a hierarchy of violence.

Superior carnivores are routinely inimical towards inferior carnivores, stealing their prey and also killing them whether for food or for spite.

Thus the leopard is at continual risk, both while patrolling its territory and in the interval between catching prey and hoisting it into a tree. If the superior carnivores hear a ruckus, they are likely to run towards the noise to capitalise on an opportunity for both food and gratuitous murder.

The main prey species of the leopard play on this vulnerability: when they spot the predator they not only denounce it, they embarrass it in what can seem like a superfluously noisy way.

The snorts, hoots and barks of e.g. impala (Aepyceros melampus, https://www.youtube.com/watch?v=U_GbVH8AOEY), chital (Axis axis, https://www.youtube.com/watch?v=hODA3OEzvaU and https://www.youtube.com/watch?v=WHrINU-E9Sk), sambar deer (Rusa unicolor, https://wildambience.com/wildlife-sounds/sambar-deer/), baboons (Papio spp., https://www.youtube.com/watch?v=za839cpwUh0) and langurs (Semnopithecus spp., https://www.youtube.com/watch?v=khxxVOg2Gtw) can be so frenetic and sustained that they seem beyond any purpose of informing each other or letting the leopard know that it has lost the element of surprise.

Could it be that the prey animals are calling in the superior carnivores to punish the leopard?

It is hardly surprising that, once the stalking leopard is spotted, it tends to reveal itself. What is surprising is that it tends to walk off in the open with its caudal flag activated. Since it lacks credibility in protesting its innocence, could this be a case of exaggerated appeasement?

Further observations are needed, comparing daytime and nighttime, to test whether the caudal flag reassures ruminants or monkeys enough to shoosh them and to curtail the risk from superior carnivores (https://www.youtube.com/watch?v=EuZo4PkIti4). However, if so this would mean that the caudal flag is an anti-predator adaptation, albeit one mediated by the leopard's prey animals.

The leopard tends to be both killer and victim, whereas the puma and the jaguar tend to be unambivalent killers. For the leopard, better to raise a flag of surrender to the whistleblowing prey than risk the arrival of those to which surrender means death. So, does the leopard emulate various species of ruminants in possessing a caudal flag because it - more than any comparable big cat - remains vulnerable even as it predates?

to be continued in https://www.inaturalist.org/journal/milewski/54706-why-is-there-a-caudal-flag-in-the-leopard-panthera-pardus-but-not-the-puma-puma-concolor-or-the-jaguar-panthera-onca-part-2#...

Posted on 23 de julho de 2021, 10:33 AM by

milewski

Comentários

Interesting observations... though it leaves me with a question regarding the position the cougar and jaguar occupy as relatively unrivalled top predators in the Americas, namely that this was a position they did not occupy until the end of the Pleistocene and that they would have coexisted with other large carnivores that would surely have outranked them in a "hierarchy of violence" as you quite wonderfully put it - American lions Panthera atrox spring to mind as the top contenders, but the various sabretooth cats and giant short-faced bears may also have posed a threat to mid-sized felids. Even keeping in mind that - at least in North America; my knowledge of South America is much more limited - Pleistocene jaguars grew to larger sizes than their modern counterparts, they still would have been considerably outclassed in size by the other carnivores mentioned above.

Should we then suspect that the jaguars of the Pleistocene (whether they are merely subspecies of the extant P. onca or, as some of my colleagues suspect, a separate species in North America, P. augusta) might've had leopard-like caudal flags or other forms of obvious supplicatory display in environments thick with other serious competitors? Or might this be a feature which owes existence to a very specific set of habitat conditions to evolve? That you note it occurs in leopards across their range, I suspect the latter option would not be the best explanation...

Anyway, it is interesting to think about, and as a paleontologist (studying proboscideans, which are of course not animals noted for their striking coloration or patterning!) I find that I am ever-increasingly curious about the evolutionary histories of features like unusual coloration or strange quirks of behavior in modern animals. I am, as I'm sure you can tell, no expert on the topic but I find it fertile ground for questions and like to imagine how such scenarios in the modern world and their evolutionary emergence might be applied to lives and histories of animals in the past.

Cheers!
Matt

Publicado por matthewinabinett quase 3 anos antes

@matthewinabinett Many thanks for your thoughts. The differences between the Pleistocene and the present in the large mammal fauna are indeed profound and inescapable in any topic like this one. However, if we keep the questions as direct as possible, I guess we should begin with why the leopard signals to its prey species in a way that other large felids do not. How might it benefit both the leopard and deer/monkeys for the leopard to offer communication of this sort? I suspect that when we figure this out there might be the same eureka moment as we had when we realised the value of stotting behaviour: the prey species cooperates with the scanning predator, at the expense of some other member of its own species, to ensure that the 'right' (least-fit to survive) individual is killed, with minimum risk to all involved except the victim. Please do let's think more on this. As a minor version of the same puzzle, why do felids of many species twitch their tail-tips while stalking? This movement may be invisible to the stalked, but it seems to make the stalker unnecessarily visible to others, such as birds or squirrels nearby, which might give the game away. Surely it would be more efficient for the nervous system of felids to be adapted to avoid this nervous twitching altogether in this context? The stalking felid seems to be signalling to some 'third party', but which and why?

Publicado por milewski quase 3 anos antes

The presence of white on the tail of the Sri Lankan leopard but a paucity of photographs of it engaging in the caudal flagging posture certainly seems like a good indicator that this behavior might be something related to the presence of competitive predators. Given that, to the best of my knowledge, the leopards of Sri Lanka have been in an environment free of larger cats since at least the end of the Pleistocene, that certainly seems like more than enough time for a behavior like this to be lost. I think perhaps the leopards of mainland India would be the place to turn next, as they coexist with similar prey animals to the Sri Lankan leopards, but crucially with the presence of tigers (and historically lions at least regionally) - if there seems to be a higher incidence of caudal flagging in these leopards it can probably be hypothesized that it really does represent some form of "appeasement" behavior. I began to look over the Indian leopard photos on here on iNat but so far no dice... This is certainly a curious path!

Publicado por matthewinabinett mais de 2 anos antes

@matthewinabinett Many thanks for these thoughts, with which I agree. At least one of the links I gave in the text of my Post, above, does indeed show a particularly striking case of the caudal flag in the Indian leopard. So I do predict that a photo-comparison of the Indian and Sri Lankan leopards will support this hypothesis.

By the way, I'd like to mention another puzzle about the leopard. When I was in Wilpattu, Sri Lanka, several years ago, I clearly observed several females of the Indian peafowl (Pavo cristatus) walk right in front of a leopard semi-hidden in a thicket, without being attacked. In fact, this individual did not even fix its gaze on the birds; I got the impression that the leopard regards the Indian peafowl as 'non-prey'. Then, while looking for videos illustrating the caudal flag in the Indian leopard, I found this case of an individual leopard activating its caudal flag to an extreme degree right close to a group of females of the Indian peafowl (https://www.youtube.com/watch?v=sWo7BiyDT04). Based on these instances, I'm starting to suspect that the leopard ignores the Indian peafowl as prey but fears it as an an embarrassor (if there is such a word). And if it really is true that the Indian peafowl has some sort of chemical defence making it 'non-prey' to both leopard and tiger, what does this add to that classic example of Zahavi's handicap principle w.r.t. sexual selection? Everyone familiar with peafowl must wonder how this bird survives the encumbrance of the masculine tail in a world of predators, but is it possible that something as obvious has been hiding in plain sight as that P. cristatus is simply unpalatable?

Publicado por milewski mais de 2 anos antes

Here is one good view of the activation of the caudal flag in India: https://www.youtube.com/watch?v=76gQoSwtQW0.

Publicado por milewski mais de 2 anos antes

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@matthewinabinett Hi again Matt, I have just tested the Sri Lankan ssp. (kotiya) of the leopard by seeking photos of the caudal flag on that island.
Out of many photos, so far only this one shows the caudal flag: https://www.dreamstime.com/walking-sri-lankan-leopard-panthera-pardus-kotiya-big-spotted-wild-cat-nature-habitat-yala-national-park-lanka-widlife-image192857408.

Most of the photos do not show the flag. It seems possible that, although at least some white occurs on the underside of the tail in all populations of the leopard, there may be enough regional variation such that, partly by behavioural modification, the flag is reduced in a ssp. finding itself to be the top carnivore throughout its range. Your further thoughts?

Here is an alarm call of the Indian peafowl: https://www.youtube.com/watch?v=2zOpK31b7nQ. However, apparently there are six different alarm calls (https://www.researchgate.net/publication/238351528_Seasonal_and_diurnal_use_of_eight_different_call_types_by_Indian_peafowl_Pavo_cristatus).

@matthewinabinett

The following series illustrates the caudal flag in females of Panthera leo, particularly the sheen on the pale pelage of the ventral surface of the distal tail-stalk.

However, this individual female has an unusually small tail-tassel (normally, the black tail-tassel is larger in females than in males of P. leo):

https://www.dreamstime.com/stock-photo-big-male-lion-gorgeous-mane-goes-savanna-national-park-kenya-tanzania-maasai-mara-serengeti-excellent-illustration-image79069317

https://www.dreamstime.com/stock-photo-meeting-lion-lioness-savannah-national-park-kenya-tanzania-masai-mara-serengeti-excellent-illustration-image79069747

https://www.dreamstime.com/stock-photo-lioness-savannah-national-park-kenya-tanzania-masai-mara-serengeti-excellent-illustration-image79105582

https://www.dreamstime.com/stock-photo-meeting-lion-lioness-savannah-national-park-kenya-tanzania-masai-mara-serengeti-excellent-illustration-image79069703

For comparison, caudal flag in males of Panthera pardus: https://www.dreamstime.com/leopard-patiently-waiting-wildebeest-to-cross-nile-river-migration-leopard-patiently-waiting-image124421306

Publicado por milewski cerca de 1 ano antes

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Publicado por milewski 9 meses antes