First Perigondwanan record of actinolepids (Vertebrata: Placodermi: Arthrodira) from the Lochkovian (Early Devonian) of Spain and its palaeobiogeographic significance

Abstract

Different palaeogeographic models have been proposed for the position of Laurussia (including Baltica) and Gondwana-derived microcontinents (including Ibero-Armorica) during Ordovician to Late Carboniferous times. Principal differences concern the presence and duration of a large ocean, the Rheic Ocean, acting as a faunal barrier between these areas. The timing of the collision of Laurussia with Gondwana and/or Gondwana-derived terranes continues to be debated. Here we present new faunal data revealing close biogeographical relations between Ibero-Armorica (“Perigondwanan” or Gondwanan derivate terranes) and Podolia (SE margin of Baltica, in Laurussia). The placoderm assemblage found in the mid-late Lochkovian (Lower Devonian) of Celtiberia (north-central Spain), consisting of the ‘actinolepid’ species Kujdanowiaspis podolica, Erikaspis zychi and the acanthothoracid Palaeacanthaspis aff. P. vasta, is similar, both in terms of taxonomy and stratigraphic record, to that encountered in the Lochkovian of Podolia (Ukraine; Laurussia) and until now considered as endemic to this region. Moreover, vertebrate faunal links between Podolia and Celtiberia are also extended to the chondrichthyan scale-based species Seretolepis elegans and Altholepis composita previously documented exclusively from Laurussian localities (Podolia and Mackenzie Mountains in Canada), which occur together with the placoderm remains described herein. These evidences support the hypothesis that intermittent shallow neritic migration paths between Podolia (as well as “Avalonia”) and Iberia existed in the late Lochkovian, agreeing with a palaeogeographic reconstruction showing close proximity between peri-Gondwanan or Gondwana-derived terranes and Laurussia. It supports the palaeogeographic model of the non-oceanic Variscan Mobile Crustal Field and it corroborates the arguments against wide oceans, acting as biogeographical relevant barriers, between Baltica and Gondwana in early Devonian times. The distribution patterns of heavy-shelled ostracods, turbidicolous brachiopods, and Rhenish trilobites also support these conclusions.