Application of integrative taxonomy combining phylogenetic and geometric morphometric techniques in a snapping shrimp (Alpheus) species complex (Decapoda: Caridea: Alpheidae)

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Abstract
Snapping shrimps of the genus Alpheus Fabricius, 1798 exhibit remarkable diversity, with over 300 described species. The genus is well-known for its prevalence of species complexes, necessitating the need of new tools to facilitate species discrimination. Traditional taxonomic methods in Alpheus have primarily relied on the examination of morphological traits or comparative morphometric measurements, with an emphasis on variation in the major chela and rostro-orbital region. We applied an integrated approach that combines molecular genetics and geometric morphometrics to investigate the A. gracilipes Stimpson, 1860 species complex. We additionally applied geometric morphometric techniques to study the major chela and the rostrum across different species, and used three mitochondrial genes (12S, COI, and 16S) to reconstruct phylogenetic relationships of this complex. Our results demonstrate the first application of geometric morphometric techniques to Alpheus snapping shrimps, and highlight the significance of the major chela and rostrum as taxonomically informative traits. Furthermore, we use DNA barcodes and geometric morphometric techniques to the A. gracilipes species complex to reveal two previously unidentified cryptic species. We present the first phylogenetic reconstruction of this species complex, with new localities and expanded distribution ranges reported for many species.

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