The strophomenide brachiopod Ahtiella Opik in the Ordovician of Gondwana and the early history of the plectambonitoids

The Precordilleran species Ahtiella argentina Benedetto and Herrera, 1986 is redescribed and illustrated and Monorthis coloradoensis Benedetto, 1998b from northwestern Argentina is reassigned to the genus Ahtiella Opik, 1932. Ahtiella famatiniana new species from volcaniclastic rocks of the Famatina range (western Argentina) and Ahtiella tunaensis new species from the Precordillera basin (Cuyania terrane) are proposed. Paleogeographic and stratigraphic evidence strongly suggests that Ahtiella originated in the Andean region of Gondwana to further migrate to Avalonia, Baltica, and Cuyania. Contrary to previous assumptions, the fossil record from the Famatina volcaniclastic succession suggests that the plectambonitoid Ahtiella famatiniana n. sp. evolved from the hesperonomiid orthoid Monorthis transversa Benedetto, 2003 that always occurs in the underlying strata. Phylogenetic analysis of Ahtiella species shows that A. famatiniana n. sp. and the Peruvian A. zarelae Villas in Gutierrez-Marco and Villas, 2007 are not only the earliest species of the genus but also are morphologically intermediate between Monorthis Bates, 1968 and the later and more derived species of Ahtiella from Baltica and Cuyania. If, as empirical evidence presented here shows, Ahtiella originated from Monorthis through a series of minor transformations, then the impressive morphological gap between orthides and strophomenides was bridged through short-time cladogenesis events, suggesting that it might not have a definite discontinuity between the species level evolution and the origin of higher taxa (macroevolution).