Helcionellids and the Early Molluscan Morphology
Helcionellids, which appear in the fossil record over 540 million years ago, are among the earliest identifiable molluscs, characterized by their snail-like shells. These organisms predate the earliest trilobites and indicate that shelled molluscs were among the first complex life forms to evolve. The helcionellids display a variety of sizes, which may represent different growth stages, and some paleontologists propose that they could be ancestral to modern gastropods. However, the definitive evidence for gastropod torsion, a unique twisting of the body that is characteristic of this class, is not present in these early fossils.The Origin and Evolution of Molluscan Shells and Cephalopods
The molluscan shell is thought to have evolved from a primitive mucus layer that hardened into a protective cuticle and later became mineralized. This evolutionary innovation led to the development of more complex respiratory structures, such as gills, to support the enclosed body. The earliest molluscs likely had shells composed of aragonite. The fossil record of early cephalopods, including the Middle Cambrian Nectocaris and the Late Cambrian Plectronoceras, provides evidence of shelled existence. Plectronoceras, in particular, shows complex shell features such as septa and a siphuncle, although these early cephalopods were not active swimmers but rather benthic, crawling along the ocean floor.The Diversification of Bivalves and Emergence of Modern Forms
The earliest bivalve fossils, Fordilla and Pojetaia, date back to the Early Cambrian period. By the Ordovician, bivalves resembling modern forms had emerged. During the Cretaceous period, a group of bivalves known as rudists became prominent reef builders but were wiped out in the Cretaceous–Paleogene extinction event. Despite such losses, bivalves have thrived and today represent a highly diverse and widespread class of molluscs, with numerous ecological roles in aquatic environments.Ongoing Controversies in Molluscan Phylogenetic Relationships
The phylogenetic tree of molluscs is a subject of active debate among scientists. This debate includes the relationships among extant mollusc classes and the reevaluation of the classification of certain fossil groups. As members of the Lophotrochozoa superphylum, molluscs share a common ancestry with annelid worms and other marine organisms. The characteristics of the last common ancestor of all molluscs, such as whether it exhibited segmentation, remain elusive. Morphological and molecular data have led to competing hypotheses, such as the "Aculifera" and "Testaria" models, which propose different evolutionary relationships among molluscan classes based on shared anatomical features.