A quantitative assessment of ontogeny and moulting in a Cambrian radiodont and the evolution of arthropod development

Radiodonta is a clade of stem euarthropods of central importance to our understanding of the evolution of this phylum. Radiodonts include some of the largest early Paleozoic animals, however little is known about their ontogeny. We present an analysis of moulting patterns and ontogeny in the radiodont Stanleycaris based on 263 exceptionally preserved specimens from the mid-Cambrian (Wuliuan) Burgess Shale. Ranging in size from 10 to 83mm, this constitutes the most extensive radiodont ontogenetic series known. Using a novel morphospace approach, we show that putative carcasses and exuviae can be quantitatively distinguished by the particular suites of structures preserved and their modes of preservation. We propose that Stanleycaris, and probably other radiodonts, moulted via a suture near the anterior of the trunk. Similar anterior moulting strategies, with a suture located at the head-trunk boundary, are shared with some Cambrian euarthropods and are potentially ancestral. Allometric analyses suggest that as Stanleycaris body size increases, the head sclerite and neck become relatively broader, while the trunk and flaps become slightly longer. The eyes developed precociously indicating an important role of visual processing in juveniles. Finally, we find evidence for an initial anamorphic developmental phase, where segment number increases at least from 11 or 12 up to 17, followed by an epimorphic phase, in which growth continued without segment addition. This is consistent with the hypothesis that finite post-embryonic segment addition (hemianamorphosis) is ancestral for arthropods and refines the timing of the origin of this important developmental mode.Data files can be opened in R (R Core Team) with the code provided. See README file for further details of file contents.Funding provided by: Natural Sciences and Engineering Research Council of CanadaCrossref Funder Registry ID: http://dx.doi.org/10.13039/501100000038Award Number: 341944Funding provided by: Natural Sciences and Engineering Research Council of CanadaCrossref Funder Registry ID: http://dx.doi.org/10.13039/501100000038Award Number: Vanier CGSFunding provided by: Government of OntarioCrossref Funder Registry ID: http://dx.doi.org/10.13039/100013873Award Number: Ontario Graduate Scholarshi