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Thank you for that. Over the past few days he has started to bite (sometimes softly, sometimes too hard) much more often - and even when there are no toys in sight.
I've noticed that he is biting hard when he doesn't want to do something - like step up when he's on my shoulder, or when I go to give him a scratch and he doesn't want it. I have started to put him back in his cage for about 10 min without any fuss when he bites hard. Often I'll say no bite sternly but that hasn't really deterred him much. Hopefully he'll learn that it isn't acceptable to bite hard with a bit of training and maturity.
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