Fossil evidence for the origin of spider spinnerets, and a proposed arachnid order

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Abstract
Silk production from opisthosomal glands is a defining characteristic of spiders (Araneae). Silk emerges from spigots (modified setae) borne on spinnerets (modified appendages). Spigots from Attercopus fimbriunguis, from Middle Devonian (386 Ma) strata of Gilboa, New York, were described in 1989 as evidence for the oldest spider and the first use of silk by animals. Slightly younger (374 Ma) material from South Mountain, New York, conspecific with A. fimbriunguis, includes spigots and other evidence that elucidate the evolution of early Araneae and the origin of spider silk. No known Attercopus spigots, including the original specimen, occur on true spinnerets but are arranged along the edges of plates. Spinnerets originated from biramous appendages of opisthosomal somites 4 and 5; although present in Limulus, no other arachnids have opisthosomal appendage homologues on these segments. The spigot arrangement in Attercopus shows a primitive state before the reexpression of the dormant genetic mechanism that gave rise to spinnerets in later spiders. Enigmatic flagellar structures originally described as Arachnida incertae sedis, are shown to be Attercopus anal flagella, as found in Permarachne, also originally described as a spider. An arachnid order, Uraraneida, is erected for a plesion, including these two genera, based on this combination of characters. The inability of Uraraneida precisely to control silk weaving suggests its original use as a wrapping, lining, or homing material.
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