The Fossil Record of the Clam Shrimp (Crustacea; Branchiopoda).

Clam shrimp (the paraphyletic assemblage of spinicaudatans, laevicaudatans, cyclestherids and the extinct leaiins) are small, bivalved branchiopod crustaceans that specialize in ephemeral freshwater habitats. They have a long fossil record (Devonian onward) that has often been overlooked. Here we briefly review the fossil record of the major groups of clam shrimp and clear up some misconceptions in the literature as to their origin. The dominant group of clam shrimp in the fossil record is the Spinicaudata, which have a diverse fossil record beginning in the Devonian. The clam shrimp suborder Laevicaudata are known from the Permian, with possible soft-part preservation from the Jurassic. However, owing the character-poor nature of these fossils, it is impossible to tell if they represent crown group or stem group laevicaudatans. In contrast, the total group Spinicaudata have a rich record of mostly carapace fossils- the earliest from the Early Devonian. The leaiins are an enigmatic extinct diplostracan lineage thought to be closely related to the spinicaudatans. They have a record that extends from the Middle Devonian to the Permian. The Cyclestherida have a somewhat problematic fossil record: there are no examples of cyclestherids preserved with soft-parts, so the only character used to assign fossils to this lineage is the carapace shape. According to that metric, cyclestherids have a record that begins in the Middle Devonian. Exceptionally preserved clam shrimp are found in the Paleozoic and Mesozoic. Assessing holistically what is known about the clam shrimp fossil record along with carapace morphology, carapace ornamentation and examples of exceptional preservation will ultimately contribute to a synthetic paleontological and neontological understanding of the group, its systematics and evolution.