†Henningsmoenicaris scutula, †Sandtorpia vestrogothiensis gen. et sp. nov. and heterochronic events in early crustacean evolution

Abstract

A detailed account of the morphology and ontogeny of the late Middle Cambrian crustacean †Henningsmoenicaris scutula is presented. Ten successive ontogenetic stages could be recognised in the material collected from various localities in Sweden. Morphogenetic changes include the development of a pair of stalked lateral eyes and the increase in the number and size of appendages and their setal armature. Notably, early stages lack ‘proximal endites’ on all post-antennular appendages; such a spine-bearing endite has previously been thought to appear simultaneously on these limbs. In †H. scutula a single functional endite appears on the third limb in an advanced stage; an additional endite appears on the second limb and, subsequently, further endites appear on more posterior limbs. Furthermore, a single specimen of †Sandtorpia vestrogothiensis gen. et sp. nov. is described. Based on this new information and data of other ‘Orsten’ taxa, particularly those assigned already to the early evolutionary lineage of Crustacea, a small-scale computer-based phylogenetic analysis was performed. This resolved the basal branchings of Crustacea s. l. as follows: †Oelandocarididae (=†Oelandocaris oelandica+†H. scutula+†S. vestrogothiensis)+(†Cambropachycopidae (=†Goticaris longispinosa+†Cambropachycope clarksoni)+ (†Martinssonia elongata+Labrophora (=†Phosphatocopina+Eucrustacea))). Plotting ontogenetic data on the phylogram and comparing the ground pattern at every node led to the detection of three peramorphic heterochronic events in the evolutionary lineage towards Eucrustacea.