Do principles of language processing in the brain affect the way grammar evolves over time or is language change just a matter of socio-historical contingency? While the balance of evidence has been ambiguous and controversial, we identify here a neurophysiological constraint on the processing of language that has a systematic effect on the evolution of how noun phrases are marked by case (i.e. by such contrasts as between the English base form she and the object form her). In neurophysiological experiments across diverse languages we found that during processing, participants initially interpret the first base-form noun phrase they hear (e.g. she. . .) as an agent (which would fit a continuation like . . . greeted him), even when the sentence later requires the interpretation of a patient role (as in . . . was greeted). We show that this processing principle is also operative in Hindi, a language where initial base-form noun phrases most commonly denote patients because many agents receive a special case marker ("ergative") and are often left out in discourse. This finding suggests that the principle is species-wide and independent of the structural affordances of specific languages. As such, the principle favors the development and maintenance of case-marking systems that equate base-form cases with agents rather than with patients. We confirm this evolutionary bias by statistical analyses of phylogenetic signals in over 600 languages worldwide, controlling for confounding effects from language contact. Our findings suggest that at least one core property of grammar systematically adapts in its evolution to the neurophysiological conditions of the brain, independently of socio-historical factors. This opens up new avenues for understanding how specific properties of grammar have developed in tight interaction with the biological evolution of our species.

Apart from common cases of differential argument marking, referential hierarchies affect argument marking in two ways: (a) through hierarchical marking, where markers compete for a slot and the competition is resolved by a hierarchy, and (b) through co-argument sensitivity , where the marking of one argument depends on the properties of its co-argument. Here we show that while co-argument sensitivity cannot be analyzed in terms of hierarchical marking, hierarchical marking can be analyzed in terms of co-argument sensitivity. Once hierarchical effects on marking are analyzed in terms of co-argument sensitivity, it becomes possible to examine alignment patterns relative to referential categories in exactly the same way as one can examine alignment patterns relative to referential categories in cases of differential argument marking and indeed any other condition on alignment (such as tense or clause type). As a result, instances of hierarchical marking of any kind turn out not to present a special case in the typology of alignment, and there is no need for positing an additional non-basic alignment type such as 'hierarchical alignment'. While hierarchies are not needed for descriptive and comparative purposes, we also cast doubt on their relevance in diachrony: examining two families for which hierarchical agreement has been postulated, Algonquian and Kiranti, we find only weak and very limited statistical evidence for agreement paradigms to have been shaped by a principled ranking of person categories.. We thank both audiences for helpful comments and questions. We also thank Sebastian Bank for an implementation of the algorithm for ranking calculation in Python, Katharina Haude for very helpful comments on an earlier draft, and two anonymous reviewers for very helpful reviews. Author contributions: AWM and BB wrote the paper; FZ and BB developed the basic idea of separating hierarchy effects from alignment typologies, while AWM developed the key proposal of 'exhaustive alignments' and TZ its computational implementation; LB collected most of the data and worked out the relevant morpheme analyses; AWM and TZ did most of the data aggregation work; BB performed the statistical analyses.