Scale morphology and squamation play an important role in the study of fish phylogeny and classification. However, as the scales of the earliest osteichthyans or bony fishes are usually found in a disarticulated state, research into squamation patterns and phylogeny has been limited. Here we quantitatively describe the scale morphology of the oldest articulated osteichthyan, the 425-million-year-old Guiyu oneiros, based on geometric morphometrics and high-resolution computed tomography. Based on the cluster analysis of the scales in the articulated specimens, we present a squamation pattern of Guiyu oneiros, which divides the body scales into 4 main belts, comprising 16 areas. The new pattern reveals that the squamation of early osteichthyans is more complicated than previously known, and demonstrates that the taxa near the crown osteichthyan node in late Silurian had a greater degree of squamation zonation compared to more advanced forms. This study offers an important reference for the classification of detached scales of early osteichthyans, provides new insights into the early evolution of osteichthyan scales, and adds to our understanding of the early osteichthyan body plan.
The Sinacanthida ordo nov. and Mongolepidida are spine-and scale-based taxa whose remains encompass some of the earliest reported fossils of chondrichthyan fish. Investigation of fragmentary material from the Early Silurian Tataertag and Ymogantau Formations of the Tarim Basin (Xinjiang Uygur Autonomous Region, China) has revealed a diverse mongolepidid and sinacanthid fauna dominated by mongolepids and sinacanthids in association with abundant dermoskeletal elements of the endemic 'armoured' agnathans known as galeaspids. Micro-computed tomography, scanning electron microscopy and histological sections were used to identify seven mongolepid genera (including Tielikewatielepis sinensis gen. et sp. nov., Xiaohaizilepis liui gen. et sp. nov. and Taklamakanolepis asiaticus gen. et sp. nov.) together with a new chondrichthyan (Yuanolepis bachunensis gen. et sp. nov.) with scale crowns consisting of a mongolepid-type atubular dentine (lamellin). Unlike the more elaborate crown architecture of mongolepids, Yuanolepis gen. nov. exhibits a single row of crown elements consistent with the condition reported in stem chondrichthyans from the Lower Devonian (e.g. in Seretolepis, Parexus). The results corroborate previous work by recognising lamellin as the main component of sinacanthid spines and point to corresponding developmental patterns shared across the dermal skeleton of taxa with lamellin and more derived chondrichthyans (e.g. Doliodus, Kathemacanthus, Seretolepis and Parexus). The Tarim mongolepid fauna is inclusive of coeval taxa from the South China Block and accounts for over two-thirds of the species currently attributed to Mongolepidida. This demonstrates considerable overlap between the Tarim and South China components of the Lower Silurian Zhangjiajie Vertebrate Fauna.
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