Postembryonic development ofDalmanitina, and the evolution of facial suture fusion in Phacopina

Abstract

AbstractA large sample of postembryonic specimens ofDalmanitina proaeva elfridaandD. socialisfrom the Upper Ordovician (Sandbian to Katian) Prague Basin allows for the first reasonably complete ontogenetic sequence of Dalmanitoidea (Phacopina). The material provides an abundance of morphological information, including well-preserved marginal spines in protaspides and meraspides, and hypostome external surfaces throughout. The development ofD. proaeva elfridais unusual due to variability in timing of the first trunk articulation. This broadens our developmental understanding of Phacopina, a diverse group of phacopid trilobites, and also allows us to study the evolution of their specializations in exoskeletal molting behavior. Adult phacopines, unlike most other trilobites, had fused facial sutures. This means that rather than molting through the sutural gape mode, characterized by opening of the facial sutures and separation of the librigenae, they disarticulated the entire cephalon in Salter’s mode of molting. For other phacopine clades (Phacopoidea) the transition to Salter’s mode occurs during the meraspid period or at the onset of holaspis, and its developmental timing is intraspecifically fixed. However, owing to the large sample size, we can see that facial suture fusion likely occurred later inDalmanitina, usually during the holaspid period, and was intraspecifically variable with holaspides of varying sizes showing unfused sutures. Further,D. proaeva elfridaspecimens showed an initial librigenal–rostral plate fusion event, where the librigenae began as separate entities but appear fused with the rostral plate as one structure (the “lower cephalic unit”) from M1, and are discarded as such during molting. Dalmanitoidea is considered to represent the first phacopine divergence, occurring earliest in the fossil record. This material therefore provides insight into how linked morphologies and behaviors evolved, potentially suggesting the timing of facial suture fusion in Phacopina moved earlier during development and became more intraspecifically fixed over geological time.