“…Although the supposed living fossil Lingula has long been considered to virtually lack morphological evolutionary changes ( Schopf, 1984 ), more recent studies have shown that lingulide brachiopods have experienced dramatic modifications in many aspects ( Liang et al, 2023 ), including arrangement of internal organs ( Zhang et al, 2008 ), life mode ( Topper et al, 2015 ), shell structure ( Cusack et al, 1999 ), and even genome ( Goto et al, 2022 ; Luo et al, 2015 ). The complexity and diversity of linguliform shell architecture was increasingly recognised in the pioneering study of Cusack, Williams and Holmer ( Cusack et al, 1999 ; Holmer, 1989 ; Williams and Cusack, 1999 ; Williams and Holmer, 1992 ).…”