N. Jb. Geol. Paläont. Abh.

297/2 (2020), 193–203 Article

Stuttgart, August 2020

Discovery of complemental males in a Pliocene accumulation of

Chelonibia testudinaria (LINNAEUS, 1758), with some notes on the
evolution of androdioecy in turtle barnacles
Alberto Collareta
With 3 figures

Abstract: While mostly hermaphrodite, thoracican cirripedes (stalked and acorn barnacles) are re-
markable in presenting also androdioecy and dioecy. With few androdioecious exceptions, the major-
ity of the acorn barnacles of the order Balanomorpha are hermaphroditic. One of the balanomorphs in
which androdioecy has been verified is the turtle barnacle Chelonibia testudinaria (Linnaeus, 1758)
(Coronuloidea: Chelonibiidae), an epibiont represented by various morphs found on several hosts,
including sea turtles, crabs, horseshoe crabs, sirenians, sea snails, sea snakes and crocodylians. An
accumulation of shells of C. testudinaria from the late Pliocene of Italy is here reappraised, and com-
plemental males are discovered among them. These complemental males consist of miniature shells
occurring largely on the radii of much larger hermaphrodites, a condition known for living C. testu­
dinaria. To the author’s knowledge, this record represents the first report of fossilized complemental
males for a member of Balanomorpha worldwide. The evolution and persistence of miniature males in
C. testudinaria is likely due to their insuring cross-fertilization when distances between neighbouring
hermaphrodites are great and speeding the rate of reproduction in this relatively short-lived species.
Chelonibia testudinaria is known from fossil specimens as old as the Pliocene and the record herein
demonstrates that androdioecy evolved in the lineage of C. testudinaria not later than the Piacenzian.
It is thus tempting to propose that a combination of high phenotypic plasticity, diverse host habits, and
androdioecy is responsible for the persistence of C. testudinaria, a refugial species.

Key words: Sexual systems, sexual determination, hermaphroditism, protandry, species longevity,
Chelonibiidae, Piacenzian, Valdelsa Basin, Italy.

1. Introduction rodite, the stalked and acorn barnacles (Cirripedia:

Among modern metazoans, hermaphroditism and di-
oecy are the most common sexual systems (Leonard
2010). In hermaphroditism, both male and female sex-
ual organs occur in the same individual during adult
life, either simultaneously or at different times, where-
as in dioecy individuals are either male or female. Oth-
er sexual systems include the rare gynodioecy (i.e., the
coexistence of females and hermaphrodites) and the
even rarer androdioecy (i.e., the coexistence of males
and hermaphrodites). While being mostly hermaph-
Thoracica) are remarkable in presenting various sex-
ual systems (namely, hermaphroditism, androdioecy
and dioecy) (e.g., Høeg et al. 2009; Yamaguchi et al.
2012; Lin et al. 2015). Although the male individuals
of Thoracica display a high degree of morphological
diversity, they are invariantly minute (i.e., dwarf) rela-
tive to their female or hermaphrodite counterparts. Fol-
lowing a practice first established by Darwin (1852,
1854), thoracican males are referred to as “dwarf
males” when coexisting with females, whereas they
are named “complemental males” when occurring be-

© 2020 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/njgpa/2020/0920 0077-7749/2020/0920 $ 3.00