Environmentally controlled phyletic evolution, blindness and extinction in Late Devonian tropidocoryphine trilobites

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Abstract
The Middle and Upper Devonian carbonate succession of the Montagne Noire, Southern France has been precisely zoned by an unbroken sequence of conodont zones. Stratigraphic control is excellent, and has allowed evolutionary changes in tropidocoryphine trilobites, which occur throughout the succession, to be directly established. The tropidocoryphines had been a stable group for some 40 million years prior to the Middle Givetian. In their last few million years, however, they underwent rapid evolution and exhibit some striking transformations of the cephalon and the regression and virtual disappearance of the eye within a relatively short space of time. They also show a marked diminution in size, and lost their original relief so that the glabella becomes virtually flush with the surface. There are two separate lineages, both of which show eye-reduction and subsequent blindness. In the earlier lineage Tropidocoryphe (Longicoryphe)-Erbenicoryphe, the main features of the ancestral rootstock are conserved and the stable, strongly divergent anterior sutural pattern of the cephalon remains the same. The eye, however, became reduced to a slightly convex surface lacking lenses, only indistinctly defined. Erbenicorphe is confined to well-oxygenated facies, and probably lived as a shallow burrower within the sediment; it became extinct in the early Frasnian. The second lineage T. (Longicoryphe)-Pterocoryphe-Pteroparia shows a remarkable backward migration of the suture, which progressively swings posteriorly in successive species spanning four Frasnian conodont zones (about three million years duration). At the same time the eye progressively degenerates so that the last forms are blind. Sutural migration and eye reduction are not genetically linked, however; the unusual form of the cephalon and suture probably resulted from an adaptation to the euxinic environment in which Pterocoryphe originated. The loss of the eye resulted from the adoption of an endobenthic habit in Pteroparia which descended from the ancestral Pterocoryphe but which had migrated to an oxygenated facies. Eye-reduction is therefore parallel in the two lineages, but superimposed upon a different original cephalic configuration. The last Pteroparia became extinct when the late Frasnian fauna became overwhelmed by the first pulse of the ''Kellwasser Event'' (probably an anoxic overturn). The evolving characters, through virtually all observed steps, show progressive unidirectional change without sudden breaks or saltations. Such unidirectional evolution is an adaptive response to constant long-lasting environmental influences.