"Can animals be moral?" Well, can they be immoral? Is the question: can other animals emulate some of our standards of goodness? (Can animals be utilitarians? Kantians?) Or is it whether they have their own standards on which they can succeed or fail? Do animals have standards? (Certainly, many animals seem to have expectations.) Remember: human morality is not one thing. (At least at the level of anthropological description.) And much within our moral thinking is essentially contestable. (Do other animals have moral disagreements?)
On the study of animals in order to understand "the origins of morality": Which morality? Whose values?
(Thinking about this stuff and reading josh blog has put me in an aphoristic mood...)