That's a well-written article but it does contain assumptions that also beg investigation. Namely, it's well and good to say that altruism doesn't necessarily exists in dogs but you are assuming it exists in humans, who are also animals. That is, I see an assumption in the humans studying the other animals and that is the assumption that the human is not an animal but somehow, above above animals. That has yet to be proven. Zoologically, man is a great ape as far as size goes, though our closest relative is the chimpanzee. We are pretty much completely bi-pedal. Relatively hairless for an ape. Hence, CS Lewis' quaint description of man by Lucifer in "The Screwtape Letters" as a relatively hairless, bipedal ape (that God has taken a fancy to.)
There is an assumption that humans do things that merit them no favor or reward and this is called altruism. But I propose that humans do things for others because it pleases them to do so. Let me call it the "It's a Wonderful Life" principle. It is a good feeling, a sense of importance and where one fits in the world to know that you made a difference. And that is totally personal.
I've known a few people that have an endless list of people they have taken care of, doled out money for to help with this or that cost, etc, and they sound sometimes as if they have been put upon. Wrongo, they enjoy being the "savior" of others and they often include their type of control along with the help. Others in the same situation would say, you are on your own, you worry about. Which one is moral? The one that leaves you to your own devices or the one that bails you out and in return, you end up having to listen to them direct your life?
Morality is slippery as it gets in semantics. For many people, morality is something outlined in a religious text, such as the Ten Commandments and the endless corollaries found right after them. But, it seems in religious texts, it is dependent on the context. For example, it is considered immoral, and illegal nowadays, to have sex with your daughters. But Lot did that and was not judged or condemned for it. Yet, in the story of Elisha, some children laugh at an old man's bald head and he gets his widdle feeling hurt and curses them and God sends to she-bears who tear the 40 children to bits. Is that just and moral?
I once read that moral behavior is that which preserves the life of self, family, community, tribe, or species, not always in that order. So, if some of this survival includes avoiding psychological damage by not engaging in incest, so be it. And, such as in war, killing someone else who would do damage to your family, country, township, whatever, that is also moral.
Is it also moral to kill another animal for food? Yes it is but I don't think it was immoral to begin with. However, we have defined some morals around how to kill an animal. If we kill another animal, it should be as quick and painless as possible. In that regard, my friend, John is a humane and moral deer hunter. Why? He hunts with a .50 cal black powder muzzleloader with his own homemade .50 cal ammo. Lighter bullets can ricochet off a shoulder blade or another bone and the animal may not die or die right away. But a .50 caliber slug will break through any bone and stop the heart. Every shot is a kill shot.
Why care at all? Well, from a mechanical standpoint, allowing a deer to die slowly allows adrenalin to get into the muscle tissue, giving the meat an off taste. Ted Nugent carries a sidearm when he is bow hunting, just in case the arrow didn't quite hit the target, which it usually does. If the shot didn't kill quickly, then a round from his Glock 9 will do it. In Texas, he has a conceal and carry permit and he is also a registered deputy sheriff of Genesee County, Michigan. Yeah, the Nuge is a cop.
We often describe a dog as having an "a-ha" moment, especially in clicker training. It's that moment, sometimes perceptible, when the dog has learned that doing this certain behavior leads to reward. It may not be accurate to state it that way but, even as a misnomer, it helps us relate a concept, especially in layman's terms.
So, we're back to the challenge of describing non-human things in human terms or models that help us to understand what we see. But even without the a-ha experience in human terms, all mammals do show the behavioristic effects of operant conditioning. That is a survival mechanism which, by that one definition, makes it moral, and I would think, regardless of species. The dog that is hunting for food is moral. So is he also moral when he attacks an intruder that threatens the survival of the group. So is he also moral when he steals the steak from your plate. After all, who's going to teach you to not leave your steak where the dog can get to it? 
But the dog may not think of it as moral and simply acting on instinct that is personal, akin to the non-linear dog theory. Attacking and warding off the intruder returns the surroundings to the status quo the dog is trying to maintain. What is that status quo? A sense of safety for oneself and one's resources, which leads to survival.
Just because a dog didn't write the chapter on the hyperbolic trig analysis of Einstein's relativistic mechanics by way of topology doesn't mean he isn't smart or is immoral.
First, what is moral in man? It's fine to say that dogs do not have morals or a-ha moments but that is only in relation to Man and first, I think, we must define those in Man. That is, what are we comparing to or against?