Comparative Canid Behavior
This summary doesn’t begin to address the subject in depth, to do so would require a book, but I hope you’ll find what I’ve written interesting nonetheless. My biggest problem with this was not spending all week on it… the topic is so interesting, and almost every fact led to a question. (Territorial? Well, that’s a highly variable concept. How does it differ between species, say dogs and wolves for example.) You can see the problem here, but one of the great things about science is that it every answer generates new questions. That’s heaven for discovery junkies like me.
AFRICAN WILD DOGS Lycaon pictus:
Also known as Cape Hunting Dogs or Painted Dogs, these canids are not “dogs” at all (note they are in a different genus than wolves and dogs). Called the wolves of Africa, they are highly social, territorial carnivores who are more dependent on group hunting than any other canid. They specialize in medium sized prey (Impala are their preferred prey) and like Hyenas, tend to rely on running their prey to exhaustion rather than the classic stalk and pounce of some canids and most big cats. (They also, like Hyenas, hunt mostly at night, thus stalking is of less importance under the cover of darkness.) They do NOT run in “relays” as often described, but tend to run in loose groups, well spread out, so that a zig-zagging quarry has a higher chance of running into another pack member when they change direction. AWDs have incredible stamina: they can run 5 km at 30 mph.
I’m not sorry I missed seeing a kill. AWDs have an incredibly high success rate as hunters (about 85%) and usually disembowl and begin consuming their prey before the animal is dead. Granted, it is probably in shock, and well might not be feeling anything at all, but I’ve seen a crocodile consume a Gazelle, and 3 African Bullalo kill another full grown buffalo, and a lion with a freshly killed zebra foal–and that’s enough for me.
Their social system is similar in many ways to that of wolves. There is usually only one breeding pair (but note there are exceptions in wolves, not sure if true in AWDs), along with non-breeding adults who also care for the young and provision the mother and pups. Unlike most social mammals in which the males leave the group and the females stay, related males stay in the pack in this species and older females emigrate if the pack becomes too large. They are completely reliant on being in a group. One study I read said that less than a group of less than 6 or 8 AWDs were incapable of successful reproduction (due no doubt to less predatory success). Their packs can become quite large, sometimes as many as 20 to 40 dogs will live in one pack.
What’s different about AWDs and Wolves is how the breeding pair maintain their rank. Apparently there are very few overt displays of aggression; you see almost none of the facial displays, growls or dominance related actions that are commonly seen in wolves. AWDs were believed at one point not to have a social hierarchy, but careful study found that wild dogs assert their social status by assuming the same stalking posture that they use when hunting or by simply supplanting (taking the space of) another. You DO see a lot of submissive behavior, from ears-back, toothy-grinned submission displays, food begging & whining. Adults will often regurgitate food to adult pack members who missed out on a kill (which happens often, since the hunting pack is so dispersed and the food is usually consumed within just a few minutes.)
BLACK-BACKED Canis mesomelas & GOLDEN JACKALS Canis aureus :
This is a Black-backed Jackal in Botswana, found feeding on what was left of a kill with its mate and a group of White-backed Vultures.
Also highly territorial, Jackals do not live in large groups like AWDs and Wolves. The usual core social unit is a monogamous pair, although both species sometimes have yearling young who stay with their parents and acts as “helpers at the nest.” (Who often trespass on neighboring territories, unlike the breeding pair who rarely do.)
Jackals could be called the “coyotes of Africa,” being of similar size, not as reliant on group hunting behavior as AWDs and Wolves, and being one of the least specialized of the canidae. Jackals eat just about anything, including insects (a large part of their diet), rodents (who they pounce upon exactly like foxes and coyotes, not to mention lots of domestic dogs), lizards, eggs, and the young of ungulates like Thompson’s Gazelles. They are most successful at preying upon fawns when in pairs: one individual will harass the mother while the other grabs the young and runs off with it.
They also, like coyotes, have developed a fondness for domestic livestock. In a study by Gusset (2009) in South Africa (or Botswana?), it was found that jackals were responsible for 724 of the 938 acts of wild animal predation on cattle and goats. And like coyotes, jackals are doing relatively well, in spite of the conflicts between them and ranchers. Lesson learned: Be a generalist- it pays off!
Jackals howl in chorus, probably as a way of keeping track of their mates, staying in contact with their young and warning off adjacent territory holders. Black-backed Jackals (BBs) tend to show more aggression within the group: siblings become “increasingly quarrelsome” and they have a “more rigid rank hierarchy” if they stay with their parents. Golden Jackals appear to be far more laid-back, with little aggression between individuals, and no instances in which the parents drive the young away (as seen in BB’s). This is ironic, because BB’s reproductive success is higher with helper’s than it is if the breeding pair goes it alone.
Breeding is always done by one pair only. Sexual behavior is suppressed in non-breeders: yearling males have small, undeveloped testes and don’t scent mark at all until they have their own territory. The guide books will tell you that you usually see jackals in pairs, and in previous trips to Africa that’s what we’ve always seen. They tend to travel in single file, the male often in front, and I’ve often seen them scent mark the exact same spot, one at a time, as they trot along a line. However, on this trip, most of the jackals we saw (mostly Black Backed) were alone. That MIGHT be because of the drought. Food was scarce I’d suspect, and there’s no value in group hunting for grasshoppers….
For more information about African Canids, and the behavior of all large African mammals, go to my African behavioral bible: Richard Estes’ Behavior Guide to African Mammals. It’s amazing.
GREY WOLF Canis lupus lupus:
Photo taken at Wolf Park, Battleground, Indiana. I took the photo, but can’t take the credit: Monty Sloan, wolf photo journalist extraordinaire, lent me his incredible camera. This was taken at the beginning of the breeding season (February), when “who’s who in the social hierarchy” was pretty much the topic of the week.
Highly territorial and highly social, “the soul of the wolf is the pack, and the soul of the pack is the wolf”. Although single wolves can and do live alone successfully, there is no debate that the wolf is essentially a pack animal who is most comfortable and most reproductively successful when living in groups. Like AWDs, wolves specialize on large prey: Elk, Bison and White-Tailed Deer. Thus, their social system is driven by the ecological niche: their specialization on large prey requires group living and cooperative hunting. Wolves are the widest ranging canid in the world, with the possible exception of the Red Fox. They are one of the few circum-polar species, being found in northern Canada, northern Europe and northern Asia.
You don’t need me to tell you that wolves usually have one breeding pair, that reproduction by other adults is usually suppressed by the alpha female who hip slams and harasses the other females at high rates before she goes into estrous. On the other hand, you are also probably aware there are a lot of exceptions: the pack in the Lamar Valley of Yellowstone that Jim and I watched was believed to have three litters.
The most interesting difference to me between Wolf and AWD behavior is the difference in their social organization and signaling. Wolves are famous for their clear facial and postural signals that continually reinforce social status. High ranking individuals use a lot of displays (rank is within sex both in Wolves and AWDs, not between them), from offensive puckers to tooth displays to growls to standing over.. etc etc to maintain their social ranking. AWDs are rarely seen to give ‘offensive’ displays, so much so that researchers initially thought they had no social hierarchy at all. And yet they do, but it is primarily maintained by submissive displays rather than displays of dominance. In the end, they seem to have a very similar social system: highly pack oriented, adults caring for the young of other individuals, a high degree of care and concern for other pack members, a high degree of play (not sure about AWDs here) even as adults. But these similar social systems are maintained in very different ways, with far more tooth displays, growls and postural displays in wolves than in AWDs.
I took this photo of “Village Dogs” on a previous trip to Africa. These were the classic ‘unowned’ foraging dogs that live off of scraps and garbage in human settlements.
I use this photo in seminars when I talk about the D word in dog behavior. (Wayne Hunthausen and I once came up with calling dominance “the concept formerly described as dominance,” complete with a Prince-like icon to avoid using that loaded term.) I could, of course, write forever about the behavior of the domestic dog, but here’s the part that seems most relevant to the species above:
1) Social hierarchy does seem to be important to at least SOME dogs. Look at the clear displays in the photo above. I don’t think we can toss out the concept just because it has been so mis-used by so many (and that is an understatement to say the least). “Dominance” is a relationship (thus, there is no such thing as a “dominant dog”) between individuals, and it simply means who has the most social freedom in a context in which they both want the same thing. It can be maintained or achieved in many ways. Fighting is the LEAST effective way to get it, and given the dangers of fighting if you have carpet knives in your mouth, hierarchies are a way to AVOID fighting, not encourage it. Usually it is maintained by visual and acoustic displays that signal to others where a dog/wolf believes itself to be in the social group.
2) Here are my speculations about dogs, wolves and their social system: Domestic dogs may be the same species as wolves, but their behavior is profoundly different from wolves. But that doesn’t mean it is not related. My impression is that dogs vary greatly, much more so than wolves, in how important social hierarchy is to them (as well as to how territorial they are). This fits with my observations of thousands of dogs over 22 years, but more importantly it also fits with the biology and genetics of the domestic dog. The physical and behavioral variability of dogs is remarkable, and is potentially caused by something called a Regulator Gene that creates changes in the developmental rates of individuals. It also creates a high degree of variability, which is exactly what we see in dogs. Dogs can weigh 2 pounds or 250, be obsessed with ball play or oblivious to it, uncomfortable with strangers or party animals who love everyone. And they are SO much less serious than wolves about hierarchy and territory–but some less than others, yes?
Exactly how dogs end up being so genetically plastic is still under investigation, but the dog’s variation in size, structure and behavior correlates with what I see as a highly variable sense of social hierarchy. Some dogs just couldn’t seem to care less, other dogs display very stereotyped and “serious-looking” social displays. Some dogs are highly territorial, others would invite the entire dog park home if they could.
I’m curious what you think. And please, before you react to the “D” word, keep this in mind. Dominance, as it should be used and understood, has NOTHING to do with good dog training. By suggesting that social status is still relevance to some dogs (between dogs that is, between dogs and people is an entirely different conversation!) I am NOT saying that it has any relevance to teaching our dogs how we would like them to behave, any more than I would teach a child that she could get anything she wanted just by throwing her social weight around… That said, what do you think? Have you found that the domestic dogs you know and have worked with vary in their interest in social status with other dogs? Or not?
Meanwhile, back at the farm: Lassie and Will are great. Lassie’s chem panel just came back and her kidney values are not just h olding steady, they are improving. Wheeeee! (She was diagnosed at Stage 2 Kidney Failure almost 2 years ago.) I still have 3 sheep who can’t seem to get enough oxygen in their lungs. The vet was out for the 3rd time, we have 2 more diseases to run tests on. I split out the 2 thin ones (Martha 2.0 and Barbie) and kept them with the lambs. That way I can grain them all together, while the other 7 ewes are stuck in the orchard pasture. The lambs and ewes can still greet through the fence, which has been found to be the least stressful way to wean off the lambs. Please explain this to the lambs, who were bawling non-stop this morning. Poor babies. They’ll stop by tomorrow, but still…..