All possible crosses have been made between the 4 species A. punctulatus, A. koliensis, A. farauti No. 1 and A. farauti No. 2. In 3 of the crosses the F1 larvae failed to survive. In the other 9 crosses both the male and female hybrid adults were sterile. A study of the development of the F1 individuals has shown that the 2 most closely related species are A. farauti No. 1 and A. farauti No. 2. A. koliensis is more closely related to these species than is A. punctulatus.
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Joan H. Bryan (1973) studied this question.
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