Physiological age-grading in females of the dung beetle Euoniticellus intermedius (Reiche) (Coleoptera: Scarabaeidae)

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Abstract
Newly emerged females of Euoniticellus intermedius (Reiche) have no differentiated oocytes in their single ovariole. The eggs mature sequentially during the maturation period, and at 27°C the first egg is laid 4–8 days after emergence. The rate of oviposition increases with temperature between 18 and 32°C. The threshold for oviposition is 19·6°C, and one egg is produced for every four day-degrees above this threshold. Most females can be sorted into the nulliparous or one of three parous categories, covering the period from emergence to the time when between 75 and 127 eggs have been deposited, on the basis of the quantity of yellow body in the ovariole. The hardness of the cuticle, the degree of oocyte development, and the presence or absence of fat-body in the abdomen can be used to subdivide the nulliparous category into three parts and to recognise old or ‘spent’ beetles. The degree of wear on the tibial teeth gives an age estimate that is independent of nutritionally induced delays and with which the ovarian age can be compared. Actual age cannot be estimated in field-caught-beetles, however, since many factors besides temperature alter the rate of brood production.