Editorial Type:
Article Category: Research Article
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Online Publication Date: 08 Dec 2020

Marine Habitat Transitions and Body-Shape Evolution in Lizardfishes and Their Allies (Aulopiformes)

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Page Range: 820 – 832
DOI: 10.1643/CG-19-300
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In this study, we use a geometric morphometric and a character evolution approach to study the evolutionary patterns of body-shape change and habitat transition in the Aulopiformes. Aulopiform fishes (lizardfishes; 289 spp.) inhabit diverse marine habitats from coral reefs to the deep sea and exhibit a wide range of body morphologies. Herein, we examine over 400 aulopiform specimens representing 38 of 44 genera and all families and identify that there are distinct patterns of body-shape change across the aulopiform radiation that coincide with habitat. A fusiform (torpedo-shaped) body is predominant among aulopiforms distributed in inshore-benthic and deep-sea benthic environments (e.g., Aulopidae, Bathysauridae, Synodontidae). There is a trend towards body elongation in taxa distributed in deep-sea pelagic habitats at depths of 200–4,000 meters (e.g., Alepisauridae, Lestidiidae, Notosudidae, Paralepididae) and a trend of body elongation with more centrally positioned dorsal and anal fins in the deep-benthic family Ipnopidae (tripodfishes). Additionally, deep-sea pelagic aulopiforms exhibit the largest variance in body-shape disparity with significant shape disparity compared to aulopiforms found in inshore-benthic and deep-sea environments. Deep-sea benthic lineages also have significantly higher body-shape variance and disparity compared to inshore-benthic lineages. We identify that there are considerable changes in body shape as aulopiform lineages transitioned to differing marine habitats. We infer the common ancestor of aulopiforms to have lived in a deep-sea benthic environment with a single transition to an inshore-benthic environment in the common ancestor of the Aulopoidei (lizardfishes, flagfin fishes) and two independent transitions into deep-sea pelagic environments, once in the common ancestor of Giganturidae, and once in the common ancestor of Alepisauroidea + Notosudoidea. This is the first study to quantitatively investigate changes in the body shape of aulopiform fishes tied to habitat transitions in marine environments from the deep sea to coral reefs. Our findings suggest that aulopiform body plans have broadly diversified in deep-sea pelagic and benthic habitats while remaining comparatively conservative in inshore-benthic habitats.

Copyright: © 2020 by the American Society of Ichthyologists and Herpetologists
Fig. 1. 
Fig. 1. 

Illustrations of aulopiform biodiversity with associated habitats. (A) Aulopidae; (B) Synodontidae: Harpadontinae; (C) Synodontidae: Synodontinae; (D) Paraulopidae; (E) Chlorophthalmidae; (F) Ipnopidae; (G) Evermannellidae; (H) Lestidiidae; (I) Giganturidae; (J) Alepisauridae: Anotopterus; (K) Alepisauridae: Alepisaurus.


Fig. 2. 
Fig. 2. 

Example of fixed landmark (blue circles) and sliding semi-landmark (purple circles) locations on a lizardfish (Synodus variegatus illustrated). Homologous fixed landmarks follow those from McMahan et al. (2011) and include: 1, anterior insertion of the dorsal fin on body; 2, posterior insertion of the dorsal fin on the body; 3, dorsal insertion of caudal fin; 4, ventral insertion of caudal fin; 5, posterior insertion of anal fin on the body; 6, anterior insertion of anal fin on body; 7, the point at which the interopercle meets the ventral body outline; 8, anterior tip of premaxilla. Digital image by R. P. Martin.


Fig. 3. 
Fig. 3. 

Results of principal component analysis depicting principal component 1 and principal component 2 with eight homologous landmarks and 70 semi-landmarks. Colored polygons highlight distribution of specimens within families of lizardfishes.


Fig. 4. 
Fig. 4. 

Phylomorphospace visualization plot of principal component 1 and principal component 2 incorporating the phylogeny from Davis and Fielitz (2010). Circle positions represent the average location in morphospace for each genus.


Fig. 5. 
Fig. 5. 

Results of principal component analysis depicting principal component 1 and principal component 2 with eight homologous landmarks and 70 semi-landmarks. Colored polygons represent distribution by marine habitat.


Fig. 6. 
Fig. 6. 

Marine habitat transitions among Aulopiformes inferred on the time-calibrated phylogeny from Davis and Fielitz (2010). Outgroups trimmed from tree to highlight Aulopiformes. Benthic continental shelf to upper slope may range from approximately 0 to 500 meters depending on region, with most continental shelves breaking at 200 m. Deep-sea pelagic includes the zones below the epipelagic starting with the mesopelagic at 200 m. Maximum-likelihood character reconstruction of habitat shown with probabilities of states represented at nodes.


Contributor Notes

Department of Biological Sciences, 720 Fourth Avenue South, St. Cloud State University, St. Cloud, Minnesota 56301; Email: (AJM) alexjmaile@gmail.com. Send reprint requests to AJM.
School of Marine Sciences, University of Maine, Orono, Maine 04469.
Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131.
Biodiversity Institute, University of Kansas, Lawrence, Kansas 66045.

Associate Editor: W. L. Smith.

Received: 17 Oct 2019
Accepted: 02 Aug 2020
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