Homologous shell microstructures in Cambrian hyoliths and molluscs

Hyoliths were among the earliest biomineralizing metazoans in Palaeozoic marine environments. They have been known for two centuries and widely assigned to lophotrochozoans. However, their origin and relationships with modern lophotrochozoan clades have been a longstanding palaeontological controversy. Here, we provide broad microstructural data from hyolith conchs and opercula from the lower Cambrian Xinji Formation of North China, including two hyolithid genera and four orthothecid genera as well as unidentified opercula. Results show that most hyolith conchs contain a distinct aragonitic lamellar layer that is composed of foliated aragonite, except in the orthothecid New taxon 1 that has a crossed foliated lamellar microstructure. Opercula are mostly composed of foliated aragonite and occasionally foliated calcite. These blade or lath‐like microstructural fabrics coincide well with biomineralization of Cambrian molluscs rather than lophophorates, as exemplified by the Cambrian members of the tommotiid‐brachiopod linage. Accordingly, we propose that hyoliths and molluscs might have inherited their biomineralized skeletons from a non‐mineralized or weakly mineralized common ancestor rather than as a result of convergence. Consequently, from the view of biomineralization, the homologous shell microstructures in Cambrian hyoliths and molluscs strongly strengthen the phylogenetic links between the two groups.     

Interpreting ‘shelly’ fossils preserved as organic films: the case of hyolithids

Most studies of Burgess Shale‐type preservation have focussed on soft‐bodied organisms, but ‘shelly’ fossils are also preserved as carbonaceous films. These films are usually interpreted as coherent organic layers – often external sheaths or periostracal layers – that were present in the original mineralized elements. The example of hyolithids shows that the organic films of skeletal parts do not represent original ‘layers’, but a composite resulting from the coalescence, into a single carbonaceous film, of all the preservable organic matter present in the skeletal element. The diagenetic processes that led to Burgess Shale‐type preservation, which involve the polymerization of organic matter and the loss of original internal structure and chemical integrity of the original tissues, are entirely compatible with – and could account for – the characteristics observed in the fossil films of hyolithid skeletal elements. These observations have general implications for the interpretation of other organisms preserved as carbonaceous films, such as the diverse and often problematic Cambrian sponges.     

新疆下寒武统玉尔吐斯组软舌螺口盖和口盖状化石

文中系统描述了新疆乌什－阿克苏地区下寒武统玉尔吐斯组软舌螺口盖和口盖状化石５属７种,其中包括２新属３新种。Ｘｙｓｔｒｉｔｈｅｃａ的口盖具有复杂的锁骨构造,Ｐａｃｈｙｔｈｅｃａ的口盖有明显的背腹扇面之分,它们与具口唇的软舌螺Ｍｉｃｒｏｃｏｒｎｕｓ共生。这些演化上较高级的软舌螺与原牙形类Ｊｉａｎｇｓｈａｎｏｄｕｓ,Ｇａｐｐａｒｏｄｕｓ,Ａｍｐｈｉｇｅｉｓｉｎａ;假牙形类Ｒｈｏｍｂｏｃｏｒｎｉｃｕｌｕｍ     

扬子地台早古生代软舌螺化石记述

该文的标本采自云南昆明、晋宁下寒武统筇竹寺组,龙陵上寒武统核桃坪组、柳水组和江苏江宁上奥陶统顶部“新开岭层”。文中描述了软舌螺化石4属5种,1未定种。其中3新属3新种,再次丰富了早古生代碎屑岩地层中的软舌螺化石,对探讨软舌螺类演化进程,确定时代,进行地层对比均有重要价值。     

Plywood‐like shell microstructures in hyoliths from the middle Cambrian (Drumian) Gowers Formation,Georgina Basin,Australia

Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators.