Nymphal growth efficiencies are high, and P/R ratios are in excess of 1.00, but the population's P/R ratio declines during the non-growing adult period to 0.57. Growth: ingestion efficiency for an individual declines from 35 % at the end of instar one, to 10 % at the beginning of September. Using the individual energy budget as a model, it can be shown that, although actual yields may vary two fold, given that almost half the population must survive to September to ensure population survival, population growth: ingestion efficiencies lie between 10% and 13 %, regardless of the pattern of mortality.
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John Hinton (1971) studied this question.
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