Comment on "The lower Cambrian lobopodian Cardiodictyon resolves the origin of euarthropod brains".

Strausfeld et al. (Report, 24 Nov 2022, p. 905) claim that Cambrian fossilized nervous tissue supports the interpretation that the ancestral panarthropod brain was tripartite and unsegmented. We argue that this conclusion is unsupported, and developmental data from living onychophorans contradict it.

Expression of engrailed in the developing brain and appendages of the onychophoran euperipatoides kanangrensis (Reid).

We have cloned an engrailed-class gene in the onychophoran Euperipatoides kanangrensis and investigated its expression using in situ hybridisation. The expression pattern was found to differ drastically from that previously described for another onychophoran species. In the present investigation, engrailed transcripts were detected in a subset of developing neurons in the brain anlage, and in the mesoderm as well as ectoderm of the developing limb buds. The engrailed positive cells of the brain are of differing developmental maturity, ranging from subepidermal neuronal precursors to neurons located basally in the embryo with developing axons. The lack of the traditional expression in the posterior compartment of segments reported earlier in onychophorans is discussed, and we suggest that onychophorans may have acquired two copies of engrailed with different functions.

Head development in the onychophoran Euperipatoides kanangrensis with particular reference to the central nervous system.

The neuroectoderm of the Euperipatoides kanangrensis embryo becomes distinguishable during germ band formation when the antennal segment is evident externally. During later stages of development, the neuroectoderm proliferates extensively and, at the anterior part of the head, newly-formed neuron precursor cells occupy most of the volume. The antenna forms from the dorsolateral side of the anterior somite. The antenna has no neuroectoderm of its own at the onset of its formation, but instead, neurons migrate out to the appendage from the nearby region of the developing brain. When the antennal tract is formed it is positioned horizontally in the brain, in line with the antennal commissure. Only later, and coincidentally with the anterior repositioning of the antenna, is the tract's distal part bent anteriorly and positioned laterally. The eye starts to develop posteriorly to the antenna and the antennal commissure. This suggests that the segment(s) associated with the onychophoran eye and antenna are not serially homologous with segments carrying equivalent structures within the Euarthropoda. Evidence is presented to further support the presence of a terminal mouth in the ground plan of the Onychophora and, hence, an acron may not exist in the arthropod clade.