# Beyond the sandy bottom: evolutionary and taxonomic insights into lizardfishes (Teleostei: Aulopiformes)

**Authors:** Shih-Yu Wang, Hsin Lee, Barry C. Russell, Wei-Jen Chen

PMC · DOI: 10.7717/peerj.20735 · PeerJ · 2026-03-06

## TL;DR

This study reveals the evolutionary history and hidden diversity of lizardfishes, suggesting taxonomic revisions and clarifying their phylogenetic relationships.

## Contribution

The study provides a robust phylogenetic framework and identifies cryptic species diversity in lizardfishes using molecular data.

## Key findings

- The Synodontidae family is not monophyletic, but its subfamilies are distinct and warrant family status.
- Only Harpadon and Trachinocephalus are monophyletic genera within the family.
- Species delimitation methods identified 108 putative species among ~60 morphospecies, indicating significant cryptic diversity.

## Abstract

Lizardfishes (family Synodontidae) are benthic, carnivorous fishes that primarily inhabit tropical and subtropical sandy seabeds and play an important role in benthic ecosystem functioning. They are characterized by cylindrical bodies and large mouths and currently comprise 84 recognized species across four genera: Synodus, Harpadon, Saurida, and Trachinocephalus. However, the systematics of the Synodontidae—from higher-level classification to species-level taxonomy—has remained contentious since the family was established by Gill in 1861, highlighting the need for a robust phylogenetic framework for further revision. This study aimed to (1) reconstruct the phylogeny of the Synodontidae and related Aulopiformes, (2) test the monophyly of the family and its genera, and (3) assess species diversity and validity using an integrated approach.

We analyzed a multi-nuclear gene dataset (RH, RAG1, ZIC1, and ENC1) and complete mitochondrial genomes to investigate higher-level phylogenetic relationships, evolutionary origins, and divergence times within the Aulopiformes. Intra-familial relationships were examined using combined mitochondrial 12S and COI gene datasets, and species-level taxonomy was assessed using 1,688 COI sequences from public and newly generated data.

Higher-level phylogenetic analyses revealed that the Synodontidae is not monophyletic, however, its two subfamilies, Synodontinae and Harpadontinae, formed well-supported, distinct monophyletic groups, justifying their recognition at the family level. Among the four genera, only Harpadon and Trachinocephalus were monophyletic. Divergence time estimates suggest that the last common ancestor of the Synodontinae (=Synodontidae stat. nov.) originated in the Late Cretaceous, while the Harpadontinae (=Harpadontidae stat. nov.) emerged in the Eocene. Species delimitation based on compiled COI sequences, using Assemble Species by Automatic Partitioning (ASAP) and Bayesian Poisson Tree Processes (bPTP) methods and supported by additional evidence, identified 108 putative species among approximately 60 morphospecies, revealing substantial cryptic diversity.

This study clarifies the evolutionary history and taxonomy of the Synodontidae, supporting the reclassification of its subfamilies as distinct families and revealing extensive hidden species diversity. These findings provide a robust framework for future research on the systematics of lizardfishes and other aulopiform fishes.

## Linked entities

- **Genes:** RHCE (Rh blood group CcEe antigens) [NCBI Gene 6006], RAG1 (recombination activating 1) [NCBI Gene 5896], ZIC1 (Zic family zinc finger 1) [NCBI Gene 7545], ENC1 (ectodermal-neural cortex 1) [NCBI Gene 8507], 12S (s-rRNA) [NCBI Gene 11540003], COX1 (cytochrome c oxidase subunit I) [NCBI Gene 4512]
- **Species:** Synodus (taxon 81391), Harpadon (taxon 94309), Saurida (taxon 143309), Trachinocephalus (taxon 172130)

## Full-text entities

- **Genes:** COI [NCBI Gene 22549028], COI [NCBI Gene 20356534], ENC1 (ectodermal-neural cortex 1) [NCBI Gene 8507] {aka ENC-1, KLHL35, KLHL37, NRPB, PIG10, TP53I10}, ZIC1 (Zic family zinc finger 1) [NCBI Gene 7545] {aka BAIDCS, CRS6, ZIC, ZNF201}, RAG1 (recombination activating 1) [NCBI Gene 5896] {aka RAG-1, RNF74}
- **Diseases:** NC (OMIM:617025)
- **Chemicals:** formalin (MESH:D005557), ethanol (MESH:D000431)
- **Species:** Synodontinae (lizardfishes, subfamily) [taxon 143310], Synodus dermatogenys (banded lizardfish, species) [taxon 1146991], Sinibrama macrops (species) [taxon 194371], Hallaxa translucens (species) [taxon 589672], Teleostei (teleost fishes, infraclass) [taxon 32443], S. fuscus [taxon 202428], Saurida fortis (species) [taxon 2942410], Synodus intermedius (sand diver, species) [taxon 94915], Synodontidae (family) [taxon 81390], Saurida brasiliensis (Brazilian lizardfish, species) [taxon 1226401], Trachinocephalus gauguini (species) [taxon 2480070], Actinopterygii (fishes, superclass) [taxon 7898], Saurida gracilis (gracile lizardfish, species) [taxon 990639], Acanthopagrus schlegelii (black porgy, species) [taxon 72011], Synechococcus sp. (species) [taxon 1131], Harpadon nehereus (Bombay duck, species) [taxon 435155], Hime (genus) [taxon 1266827], Synodus hoshinonis (blackear lizardfish, species) [taxon 197543], Trachinocephalus myops (snakefish, species) [taxon 172131], Saurida wanieso (Waneso lizardfish, species) [taxon 315323], Mochokidae (squeakers, family) [taxon 31000], microlepis [taxon 130519], S. gracilis [taxon 626134], Chlorophthalmidae (greeneyes, family) [taxon 68511], Synodus poeyi (offshore lizardfish, species) [taxon 1226406], Synodus doaki (species) [taxon 1459840], Synodus tectus (Tectus lizardfish, species) [taxon 2578359], Synodus pacificus (species) [taxon 2792404], Bathysaurus (genus) [taxon 172133], Saurida elongata (slender lizardfish, species) [taxon 197541], Saurida umeyoshii (species) [taxon 1537714], Saurida nebulosa (clouded lizardfish, species) [taxon 1074258], Saurida undosquamis (brushtooth lizardfish, species) [taxon 143315], Saurida isarankurai (species) [taxon 1604557], Somniosus pacificus (Pacific sleeper shark, species) [taxon 305516], Saurida tumbil (greater lizardfish, species) [taxon 445361], Harpadon microchir (mizutengu, species) [taxon 143314], Stylatula elongata (slender sea pen, species) [taxon 1214922], Saurida longimanus (species) [taxon 1723705], Synodus saurus (Atlantic lizardfish, species) [taxon 81392], Saurida macrolepis (species) [taxon 1426792], Meleagris gallopavo (common turkey, species) [taxon 9103], Synodus kaianus (Gunther's lizard fish, species) [taxon 172129], Squatina nebulosa (clouded angelshark, species) [taxon 55142], Onychostoma macrolepis (species) [taxon 369639], Synodus macrops (triplecross lizardfish, species) [taxon 1585515], Alepisauroidei (suborder) [taxon 143312], Saurida microlepis (species) [taxon 1481781], Synodus sageneus (peartoothed grinner, species) [taxon 2578358], Ateleopus japonicus (jellynose fish, species) [taxon 143306], Saurida micropectoralis (shortfin lizardfish, species) [taxon 1481875], Synodus variegatus (variegated lizardfish, species) [taxon 261076], Trachinocephalus trachinus (species) [taxon 2681097], Saurida lessepsianus (species) [taxon 1961710], Trachinocephalus (genus) [taxon 172130]
- **Mutations:** M for A, S for G, Y for C, K for G, W for A, R for A

## Full text

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## Figures

8 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12970317/full.md

## References

139 references — full list in the complete paper: https://tomesphere.com/paper/PMC12970317/full.md

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Source: https://tomesphere.com/paper/PMC12970317