Hi all
I generally consult the OED for questions of word usage. Suffice it to say that they acknowledge that "who" is sometimes used for animals, "usually with implication of personality." As in the following:

76. The bee-keeper who understands the language of bees, can turn it to his advantage. Here are some examples:

"When something seems to irritate the bees, who are in front of a hive, on the alighting-board, they emit a short sound, z-z-z-, jumping at the same time towards the hive. This is a warning. Then they fly and examine the object of their fears, remaining sustained by their wings, near the suspected object, and emitting at the same time, a distinct and prolonged sound. This is a sign of great suspicion. If the object moves quickly, or otherwise shows hostile intent, the song is changed into a piercing cry for help, in a voice whistling with anger. They dash ‘forward violently and blindly, and try to sting.

"When they are quiet and satisfied, their voice is the humming of a grave tune; or, if they do not move their wings, an allegro murmur. If they are suddenly caught or compressed, the sound is one of distress. If a hive is jarred at a time when all the bees are quiet, the mass speedily raise a hum, which ceases as suddenly. In a queenless hive, the sound is doleful, lasts longer, and at times increases in force. When bees swarm, the tune is clear and gay, showing manifest happiness."

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