Plectronoceratids (Cephalopoda) from the latest Cambrian at Black Mountain, Queensland, reveal complex three-dimensional siphuncle morphology, with major taxonomic implications

Abstract

The Plectronoceratida includes the earliest known cephalopod fossils and is thus fundamental to a better understanding of the origin and early evolution of this group of molluscs. The bulk of described material comes from the late Cambrian Fengshan Formation in North China with isolated occurrences in South China, Laurentia, Kazakhstan and Siberia. Knowledge of their morphology and taxonomy is limited in that most specimens were only studied as longitudinal sections, which are prone to misinterpretations due to variations in the plane of section. We describe more than 200 new specimens, which exceeds the entire hitherto published record of plectronoceratids. The material was collected by Mary Wade and colleagues during the 1970s and 1980s, from the lower Ninmaroo Formation at Black Mountain (Mount Unbunmaroo), Queensland, Australia. Despite the collecting effort, diverse notes and early incomplete drafts, Mary Wade never published this material before her death in 2005. The specimens provide novel insights into the three-dimensional morphology of the siphuncle based on abundant material, prompting a general revision of the order Plectronoceratida. We describe Sinoeremoceras marywadeae sp. nov. from numerous, well-preserved specimens, allowing investigation of ontogenetic trajectories and intraspecific variability, which in turn enables improved interpretations of the three-dimensional siphuncle morphology. The siphuncle of S. marywadeae sp. nov. and other plectronoceratids is characterised by highly oblique segments, an elongated middorsal portion of the septal neck (= septal flap) and laterally expanded segments that extend dorsally relative to the septal flap (= siphuncular bulbs). We show that this complex siphuncular structure has caused problems of interpretation because it was studied mainly from longitudinal sections, leading to the impression that there were large differences between specimens and supposed species. We revise the order Protactinoceratida and the families Protactinoceratidae and Balkoceratidae as junior synonyms of the Plectronoceratida and Plectronoceratidae, respectively. We reduce the number of valid genera from eighteen (including one genus formerly classified as an ellesmeroceratid) to three: Palaeoceras Flower, 1954, Plectronoceras Kobayashi, 1935 and Sinoeremoceras Kobayashi, 1933. We accept 10 valid species to which the 68 previously established species may be assigned. Sinoeremoceras contains 8 of the 10 plus the new species. Two species, previously referred to ellesmeroceratid genera, are transferred to Sinoeremoceras. This revised scheme groups plectronoceratids into distinct geographically and stratigraphically separated species, which better reflects biological realities and removes bias caused by preparation techniques. North China remains important containing the highest known diversity and was likely a centre of cephalopod diversification.