Analysis of the juvenile shell of Lingula anatina (Brachiopoda: Linguliformea) provides insight into the evolution of life cycles of fossil brachiopods

Inferences on the development and morphology of extinct brachiopods must be informed by the ontogeny and shell ornamentation of extant brachiopods. Although the adult shells of extant brachiopods are well studied, detailed descriptions of the embryonic and juvenile shells of extant lingulides are lacking. Here, we describe in detail the shells of juveniles of Lingula anatina Lamarck, 1801 from Vietnam and the Republic of the Philippines. The following previously unknown properties of the lingulide shell are described: (1) a distinct border between the protegulum and the brephic shell; (2) drapes that develop on both the protegulum and brephic shell; and (3) the notched anterior margin of the brephic shell. The drapes and cogs on the brephic shell may be caused by the formation of setal follicles during the planktonic stage. Specimens of L. anatina from the Philippines have larger brephic shells than those from Vietnam, probably because the former have a longer planktonic stage. Based on comparisons of the first-formed shells of extant brachiopods with published data on fossil brachiopods, we suggest that the life cycle of extant lingulides, in which planktotrophic juveniles with a shell hatch from the egg envelope, is the most evolutionarily advanced brachiopod life cycle and appeared in the early Silurian. We suggest criteria for determining the type of life cycle based on the structure of the first-formed shell of brachiopods. Finally, we consider hypothetical scenarios of life cycles of fossil brachiopods, including true planktotrophic larvae in the Cambrian linguliforms.