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Reversal to a Two-Ovaries State in Scarabaeinae (Coleoptera: Scarabaeidae)

The Scarabaeinae, typically dung beetles, forms a monophyletic group according to the criteria agreed upon by all authors. Perhaps the character that most strongly supports this unique origin is the reduction of the female gonads to a single ovary (the left one) with a single ovariole; a true synapomorphy. The first to note this reduction as a distinctive characteristic of Scarabaeinae was Heymons (1929, 1930). E. Willimzik (1930), a student of Richard Heymons, confirmed it in several other species. As a basic and distinguishing character of Scarabaeinae, ovarian reduction has been amply discussed and illustrated by Halffter and Edmonds (1982), and more recently by Scholtz (2009). All other authors who have examined a scarabaeine specimen in general studies of the ovaries in Scarabaeoidea mention an ovary with a single ovariole. The only equivalent case (one ovary, one ovariole) that we know of in Insecta occurs in the aphid family Pemphigidae (Woodward et al. 1970). Furthermore, although Scarabaeinae and Aphodiinae (the closest subfamily) share other synapomorphies, such as membranous incisive areas in their maxillae and mandibles, the Aphodiinae do not have the ovarian reduction found in the Scarabaeinae. The first case of a scarabaeine female with two ovaries was published by Heymons (1930), who found the character state in a specimen of Scarabaeus sacer L., which, in addition to a normal left ovary, also had a right ovary with a nonfunctioning ovariole. Pluot (1979) published on a second species, Onthophagus lecontei Harold, with this same condition. This author, in addition to describing the relictual gonad, did a detailed study of organogenesis in the female genital apparatus in two species of Scarabaeinae. To summarize Pluot (1979), in the larva there were two ovarian precursors, although the right one was considerably atrophied compared to the left one. The left ovary had six ovarioles, of which only one developed normally, the others were strongly atrophied. In the larva, the right ovarian precursor,