False scorpions (Arachnida, Pseudoscorpiones) from Lower Cretaceous Spanish amber

Abstract Pseudoscorpions, one of the earliest terrestrial lineages dating back to the Middle Devonian period ( c . 385 Ma), have a limited fossil record mainly as inclusions in amber. The available fossils are mostly from the Cenozoic era, making it challenging to comprehensively understand their evolutionary history and diversification over time. Here, we report the first fossils of the order from Spanish Lower Cretaceous (upper Albian) ambers. Two new genera and species in the family Garypinidae are described from Álava amber (Peñacerrada II outcrop, Álava Province): Cretogarypinus zaragozai gen. et sp. nov. and Ithioreolpium alavensis gen. et sp. nov. Remarkably, the holotype of Cretogarypinus zaragozai shows preservation of internal soft tissues. The variability observed in the arolium of both fossil garypinids serves as compelling evidence for the hypothesis of diphyletism within the clade, which was already well established before the Early Cretaceous. Additionally, an incomplete specimen consisting of an isolated palp ascribed to the family Pseudogarypidae (extant genus Pseudogarypus Ellingsen), is described from San Just amber (Teruel Province). These findings represent the earliest occurrences of the families Garypinidae (superfamily Garypinoidea) and Pseudogarypidae (superfamily Feaelloidea) in the fossil record. They not only contribute to our understanding of a diverse pseudoscorpion fauna in Spanish amber but also offer valuable insights into divergence time estimations for pseudoscorpion families and potentially for biogeographic studies.