Do key innovations unlock diversification?A case-study on the morphological and ecological impact of

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Key innovations may allow lineages access to new resources and facilitate the invasion of new adap-tive zones,potentially influencing diversification patterns.Many studies have focused on the impact of key innovations on speciation rates,but far less is known about how they influence phenotypic rates and patterns of ecomorphological diversification.We use the repeated evolution of pharyngog-nathy within acanthomorph fishes,a commonly cited key innovation,as a case study to explore the predictions of key innovation theory.Specifically,we investigate whether transitions to pharyngogna-thy led to shifts in the rate of phenotypic evolution,as well as shifts and/or expansion in the occupa-tion of morphological and dietary space,using a dataset of 8 morphological traits measured across 3,853 species of Acanthomorpha.Analyzing the 6 evolutionarily independent pharyngognathous clades together,we found no evidence to support pharyngognathy as a key innovation;however,comparisons between individual pharyngognathous lineages and their sister clades did reveal some consistent patterns.In morphospace,most pharyngognathous clades cluster in areas that correspond to deeper-bodied morphologies relative to their sister clades,whereas occupying greater areas in diet-ary space that reflects a more diversified diet.Additionally,both Cichlidae and Labridae exhibited higher univariate rates of phenotypic evolution compared with their closest relatives.However,few of these results were exceptional relative to our null models.Our results suggest that transitions to phar-yngognathy may only be advantageous when combined with additional ecological or intrinsic factors,illustrating the importance of accounting for lineage-specific effects when testing key innovation hypotheses.Moreover,the challenges we experienced formulating informative comparisons,despite the ideal evolutionary scenario of multiple independent evolutionary origins of pharyngognathous clades,illustrates the complexities involved in quantifying the impact of key innovations.Given the issues of lineage specific effects and rate heterogeneity at macroevolutionary scales we observed,we suggest a reassessment of the expected impacts of key innovations may be warranted.
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