Dirt-sifting devilfish: winnowing in the geophagine cichlid Satanoperca daemon and evolutionary implications


Winnowing is a foraging strategy common in geophagine cichlids (Cichlidae), in which sediment is sifted for food in the oral cavity. Geophagines have modified pharyngeal structures that function in winnowing, although detailed anatomical and functional information is still needed to clarify the mechanisms by which these fishes obtain food by sifting. With this study, we explore geophagine winnowing kinematics and variability of winnowing phases to test whether this is a highly modulated or stereotyped behavior. Winnowing was characterized with high-speed video of an archetypal winnower, Satanoperca daemon, which employs a three-part feeding behavior involving strike, winnowing, and sediment ejection. Over the course of feeding events, fish exhibited rapid reversal of hydraulic flow within the oral cavity and remarkable versatility during the winnowing stage. We also explored how cranial morphology varies within the clade across a phylogenetic hypothesis for the group. Cranial morphologies were analyzed in 19 species across 12 geophagine genera; principal component analysis suggests a particular winnowing morphospace exploited differentially by species. Central conclusions of this study are that the strike and ejection phases are stereotypic (low variation) but that rhythmic winnowing is highly variable. Winnowing in geophagines is not directly analogous to winnowing in surfperches (Embiotocidae), to which it is often compared. There is substantial morphometric variation across the clade, even between winnowing species. The wide range of anatomical and biomechanical variants in this diverse clade provides intriguing insight into an underexplored feeding strategy.

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We thank K. Conway and H. Prestridge (Texas A&M University Biodiversity Research and Teaching Collection), J. Armbruster and D. Werneke (Auburn University Museum of Natural History), C. Taylor and D. Wylie (Illinois Natural History Survey), and C. Lucena (Museu de Ciencias e Tecnologia PUCRS) for providing loans of geophagine specimens. We thank Susan Mochel for providing assistance with specimen dissection and gill arch preparation at the Field Museum. Andrew George provided training and assistance with use of the high-speed camera for capturing fish feeding events. Aaron Olsen provided extensive help with installation and use of the StereoMorph package in R for digitizing images.


This study was funded by Grants NSF DEB 1447421 and NSF IOS 1425049 to M. Westneat.

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Correspondence to Hannah I. Weller.

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All applicable international, national, and/or institutional guidelines for the care and use of animals were followed, under University of Chicago ACUP 72365. This article does not contain any studies with human participants performed by any of the authors.

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Weller, H.I., McMahan, C.D. & Westneat, M.W. Dirt-sifting devilfish: winnowing in the geophagine cichlid Satanoperca daemon and evolutionary implications. Zoomorphology 136, 45–59 (2017). https://doi.org/10.1007/s00435-016-0335-6

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  • Neotropical cichlids
  • Feeding kinematics
  • Substrate sifting
  • Morphometrics