What also needs to be considered is that some colony deaths are the result of unique short-term events and  other colony deaths or dwindling are due to longstanding conditions.

Some locations and types of enclosures may be more vulnerable to certain events than others, so while we necessarily consider each factor in isolation, we have to understand that everything is a trade off and part of a system.  

Moreover, while environmental conditions, may be predictable in general, they are subject to unexpected or unusual events.  Every year is different and the optimum, if it exists, is a moving target.

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