Species in Disguise: A New Species of Hornshark from Northern Australia (Heterodontiformes: Heterodontidae)
Abstract
:1. Introduction
2. Materials and Methods
2.1. Morphology
2.2. Vertebral Meristics
2.3. Dentition
2.4. Denticles
2.5. Distribution
2.6. Molecular Analyses
2.7. Institutional Acronyms
2.8. Comparative Material Examined
3. Results
3.1. Comparison with Congeners
3.2. Systematic Account
3.2.1. Synonymy
3.2.2. Type Material
3.2.3. Representative DNA Sequences
3.2.4. Etymology
3.2.5. Diagnosis
3.2.6. Description
3.2.7. Colour
3.2.8. Egg Case
3.2.9. Size
3.2.10. Distribution
3.3. Molecular Analyses
3.4. Intraspecific Variation
3.5. Comparison with Congeners
4. Discussion
4.1. Comparison between Species
4.2. Dentition and Denticles
4.3. Distribution
4.4. Egg Cases
4.5. Key to Species of Heterodontus
- Body and fins: spotted ……………………………………………………………………….. 2Body and fins with stripes, bars or bands: not spotted ………………………………… 5
- Body and fins with white spots in adults and subadults; hatchlings with thin curved parallel lines on body ………………… Heterodontus ramalheira (Western Indian Ocean)Body and fins with dark spots and darker saddles (most evident in juveniles) ….… 3
- First dorsal fin origin behind pectoral fin bases… Heterodontus quoyi (Eastern Pacific)First dorsal fin origin over pectoral fin bases …………………………………………… 4
- Back and sides with small dark spots less than a third of eye diameter in size; no light-coloured bar on interorbital surface of head …………………………………….. ……………………………………………………… Heterodontus francisci (Eastern Pacific)Back and sides with larger dark spots at least half eye diameter in size; a light-coloured bar on interorbital surface of head.…… Heterodontus mexicanus (Eastern Pacific)
- Supraorbital crests: very high ……………… Heterodontus galeatus (eastern Australia)Supraorbital crests: moderate to low ……………………………………………………. 6
- Body with a distinctive harness-like pattern of dark stripes extending from in front of dorsal fin on the dorsal midline to the pectoral and pelvic fin bases …………………..……………………………………… Heterodontus portusjacksoni (southern Australia)Body with vertical dark bands or saddles not arranged in a harness pattern …….… 7
- Body with 22–36 narrow dark bars and saddles from snout to origin of caudal fin ……………………………………………………………………………….…………….… 8Body with 5–14 broad or narrow, diffuse-edged markings from snout to origin of caudal fin ……….…………..……………………………………………………………… 9
- Narrow triangular dark marking extending from snout to midline of head in front of eyes; dark bar extending onto pectoral fin from posterior gills…………………………..……………………………………………………. Heterodontus zebra (Northwest Pacific)Dark marking on snout semicircular, not narrowly triangular; no dark bar extending from onto pectoral fin from posterior gills …….…………………………………………..………………………………………. Heterodontus marshallae n. sp. (northern Australia)
- Dorsal, pectoral and caudal fins without abruptly black tips or white apical spots; body with 11–14 dark markings from snout to origin of caudal fin ……………………………………………………………………….. Heterodontus japonicus (Northwest Pacific)Dorsal and pectoral fins and ventral caudal fin lobe with abruptly black tips, dorsal fins with white apical spots; body with four or five broad dark saddles from snout tip to origin of caudal fin ………………… Heterodontus omanensis (Western Indian Ocean)
Supplementary Materials
Author Contributions
Funding
Institutional Review Board Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
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H. marshallae n. sp. | H. zebra | ||||||||
---|---|---|---|---|---|---|---|---|---|
Paratypes | Mean | sd. | |||||||
Character | Holo | Min | Max | Min | Max | Mean | sd. | ||
Total length (mm) | 541 | 355 | 601 | 514.0 | 98.4 | 212 | 1005 | 479.9 | 271.1 |
Precaudal length | 73.9 | 73.2 | 78.4 | 75.8 | 2.0 | 67.3 | 81.1 | 73.7 | 4.6 |
Pre-second dorsal length | 48.5 | 46.1 | 53.1 | 50.0 | 2.6 | 46.6 | 57.1 | 50.7 | 3.7 |
Pre-first dorsal length | 26.0 | 23.9 | 29.5 | 26.9 | 1.9 | 23.4 | 28.6 | 25.1 | 1.8 |
Pre-first dorsal length (horizontal) | 23.1 | 22.2 | 26.6 | 24.7 | 1.9 | 21.2 | 21.9 | 21.5 | 0.5 |
Head length | 22.3 | 20.3 | 24.1 | 22.7 | 1.5 | 19.5 | 23.1 | 20.6 | 1.3 |
Head length (horizontal) | 21.6 | 20.3 | 23.5 | 21.9 | 1.1 | 18.2 | 22.1 | 19.5 | 1.4 |
Prebranchial length | 17.5 | 15.6 | 19.2 | 17.7 | 1.3 | 15.1 | 17.6 | 16.2 | 1.0 |
Prespiracular length | 12.0 | 10.5 | 13.3 | 12.1 | 1.0 | 10.5 | 13.3 | 11.5 | 0.9 |
Preorbital length | 9.6 | 8.1 | 10.5 | 9.4 | 0.9 | 7.6 | 11.1 | 8.5 | 1.2 |
Preorbital length (horizontal) | 7.9 | 6.5 | 8.9 | 8.1 | 0.9 | 5.4 | 9.6 | 6.8 | 1.4 |
Prenarial length | 3.8 | 3.1 | 4.4 | 3.9 | 0.5 | 3.3 | 3.7 | 3.5 | 0.2 |
Prenarial length (horizontal) | 2.1 | 0.9 | 2.7 | 2.0 | 0.6 | 1.8 | 2.7 | 2.1 | 0.3 |
Preoral length | 3.5 | 2.6 | 3.3 | 3.1 | 0.4 | 2.1 | 3.9 | 3.2 | 0.7 |
Prepectoral length | 21.1 | 21.2 | 24.1 | 22.3 | 1.2 | 17.5 | 21.1 | 19.2 | 1.4 |
Prepelvic length | 39.0 | 36.3 | 41.9 | 40.1 | 2.2 | 34.6 | 44.4 | 38.1 | 3.9 |
Pre-cloacal length | 39.9 | 40.8 | 44.3 | 42.3 | 1.7 | 38.2 | 48.8 | 41.9 | 4.2 |
Preanal length | 58.2 | 55.2 | 62.2 | 59.6 | 2.6 | 57.6 | 58.9 | 58.3 | 0.9 |
Interdorsal space | 17.3 | 15.3 | 17.8 | 16.8 | 0.9 | 15.6 | 20.9 | 18.5 | 2.0 |
Dorsal–caudal space | 17.6 | 16.4 | 18.7 | 17.8 | 0.8 | 13.4 | 16.7 | 15.6 | 1.2 |
Pectoral–pelvic space | 12.4 | 11.6 | 14.7 | 13.4 | 1.2 | 11.3 | 16.6 | 13.7 | 1.8 |
Pelvic–anal space | 14.7 | 11.7 | 15.5 | 14.4 | 1.4 | 15.2 | 17.6 | 16.4 | 1.7 |
Anal–caudal space | 13.2 | 11.0 | 13.5 | 12.5 | 1.0 | 9.4 | 10.5 | 10.0 | 0.8 |
Interorbital space | 7.8 | 7.4 | 7.9 | 7.7 | 0.2 | 6.2 | 8.0 | 6.9 | 0.7 |
Eye length | 3.7 | 3.7 | 3.8 | 3.7 | 0.0 | 2.6 | 3.7 | 3.3 | 0.3 |
Eye height | 2.2 | 2.1 | 2.5 | 2.3 | 0.2 | 1.6 | 2.6 | 2.0 | 0.4 |
Spiracle length | 0.7 | 0.4 | 0.7 | 0.6 | 0.1 | 0.2 | 0.6 | 0.4 | 0.2 |
Nostril width | 1.9 | 1.4 | 2.1 | 1.9 | 0.2 | 1.1 | 1.6 | 1.2 | 0.2 |
Internarial space | 4.9 | 4.9 | 5.9 | 5.4 | 0.4 | 3.9 | 5.8 | 5.2 | 0.8 |
Mouth width | 9.1 | 8.1 | 10.2 | 9.3 | 0.8 | 7.2 | 9.1 | 8.3 | 0.7 |
Upper labial furrow length | 1.5 | 1.3 | 1.7 | 1.5 | 0.2 | 0.8 | 1.4 | 1.1 | 0.2 |
Lower labial furrow length | 3.0 | 3.2 | 3.9 | 3.5 | 0.3 | 3.5 | 3.9 | 3.7 | 0.2 |
Inner nostril to upper labial furrow | 1.9 | 1.8 | 1.9 | 1.9 | 0.1 | 1.3 | 2.5 | 1.9 | 0.5 |
First gill slit height | 4.4 | 4.1 | 5.3 | 4.6 | 0.5 | 3.2 | 5.0 | 3.9 | 0.6 |
Fifth gill slit height | 2.6 | 2.0 | 3.3 | 2.6 | 0.4 | 1.8 | 3.4 | 2.4 | 0.6 |
Intergill length | 4.9 | 4.7 | 5.8 | 5.3 | 0.4 | 4.2 | 5.2 | 4.7 | 0.4 |
Pectoral fin—anterior margin | 20.1 | 18.5 | 22.4 | 20.5 | 1.7 | 18.8 | 26.1 | 22.1 | 2.7 |
Pectoral fin—height | 19.0 | 15.1 | 18.3 | 17.4 | 1.4 | 17.2 | 19.3 | 18.3 | 1.5 |
Pectoral fin—inner margin | 6.5 | 5.7 | 7.0 | 6.3 | 0.5 | 5.5 | 9.6 | 7.6 | 1.4 |
Pectoral fin—base | 7.7 | 7.2 | 8.0 | 7.6 | 0.2 | 7.0 | 8.9 | 8.0 | 0.7 |
Pectoral fin—posterior margin | 15.2 | 14.0 | 17.6 | 15.4 | 1.5 | 13.6 | 17.7 | 15.7 | 1.4 |
Dorsal caudal margin | 25.2 | 21.4 | 25.6 | 23.5 | 1.8 | 17.9 | 28.0 | 24.4 | 3.3 |
Preventral caudal margin | 14.9 | 13.1 | 14.5 | 13.9 | 0.7 | 12.1 | 14.5 | 13.2 | 1.0 |
Upper postventral caudal margin | 5.2 | 3.8 | 5.3 | 4.7 | 0.6 | 4.2 | 6.5 | 5.4 | 1.6 |
Lower postventral caudal margin | 5.2 | 4.7 | 6.1 | 5.2 | 0.5 | 3.2 | 3.5 | 3.3 | 0.3 |
Caudal fork width | 8.5 | 7.5 | 9.3 | 8.1 | 0.7 | 8.8 | 8.9 | 8.8 | 0.1 |
Caudal fork length | 12.0 | 10.6 | 13.3 | 11.5 | 1.0 | 12.4 | 12.5 | 12.5 | 0.1 |
Caudal terminal lobe length | 11.5 | 10.2 | 13.5 | 11.1 | 1.3 | 11.9 | 12.6 | 12.2 | 0.5 |
Caudal terminal margin length | 9.8 | 9.1 | 10.9 | 9.8 | 0.7 | 6.9 | 10.6 | 8.7 | 1.4 |
Caudal subterminal margin | 4.5 | 4.2 | 6.2 | 4.9 | 0.7 | 5.2 | 6.6 | 5.9 | 1.0 |
First dorsal fin—length | 13.4 | 13.7 | 14.6 | 14.1 | 0.5 | 12.0 | 15.4 | 13.5 | 1.1 |
First dorsal fin—anterior margin | 16.9 | 14.0 | 20.0 | 16.3 | 2.2 | 11.7 | 29.5 | 19.5 | 5.8 |
First dorsal fin—base | 9.0 | 9.1 | 9.8 | 9.4 | 0.3 | 8.2 | 9.6 | 8.8 | 0.5 |
First dorsal fin—height | 12.5 | 9.2 | 15.6 | 12.0 | 2.3 | 8.8 | 22.3 | 15.1 | 5.2 |
First dorsal fin—inner margin | 4.6 | 4.3 | 5.7 | 4.8 | 0.5 | 4.0 | 6.2 | 4.9 | 0.8 |
First dorsal fin—posterior margin | 12.4 | 9.0 | 13.4 | 11.5 | 1.6 | 9.2 | 20.0 | 14.0 | 4.3 |
First dorsal fin—soft fin length | 9.4 | 8.3 | 10.4 | 9.5 | 0.8 | 9.3 | 10.3 | 9.8 | 0.7 |
Exposed first dorsal spine length | 4.5 | 3.9 | 4.3 | 4.2 | 0.3 | 1.0 | 3.7 | 2.3 | 1.0 |
Base first dorsal spine | 1.1 | 1.2 | 1.4 | 1.3 | 0.1 | 0.8 | 1.6 | 1.2 | 0.3 |
Second dorsal fin—length | 12.0 | 11.1 | 12.8 | 12.2 | 0.6 | 9.6 | 12.7 | 11.2 | 1.0 |
Second dorsal fin—anterior margin | 12.8 | 10.1 | 14.5 | 12.4 | 1.5 | 8.9 | 20.1 | 14.3 | 3.8 |
Second dorsal fin—base | 8.3 | 6.8 | 8.6 | 7.8 | 0.7 | 6.4 | 8.1 | 7.4 | 0.6 |
Second dorsal fin—height | 8.8 | 6.7 | 11.1 | 8.5 | 1.5 | 6.8 | 16.4 | 10.4 | 3.8 |
Second dorsal fin—inner margin | 4.0 | 3.8 | 5.3 | 4.4 | 0.5 | 3.4 | 4.3 | 3.9 | 0.4 |
Second dorsal fin—posterior margin | 8.7 | 6.8 | 9.2 | 8.2 | 0.8 | 5.7 | 11.8 | 8.5 | 2.1 |
Second dorsal fin—soft fin length | 6.8 | 6.5 | 7.3 | 6.9 | 0.3 | 6.2 | 6.8 | 6.5 | 0.5 |
Exposed second dorsal spine length | 4.2 | 3.9 | 4.7 | 4.2 | 0.3 | 2.0 | 4.0 | 2.8 | 0.7 |
Base second dorsal spine | 1.1 | 1.1 | 1.2 | 1.1 | 0.0 | 0.8 | 1.4 | 1.0 | 0.2 |
Pelvic fin—length | 10.9 | 10.6 | 12.2 | 11.3 | 0.6 | 10.4 | 13.2 | 11.9 | 0.9 |
Pelvic fin—height | 8.2 | 6.5 | 8.0 | 7.6 | 0.6 | 6.6 | 8.1 | 7.2 | 0.5 |
Pelvic fin—inner margin length | 6.3 | 5.9 | 6.5 | 6.2 | 0.2 | 4.4 | 6.8 | 5.8 | 0.9 |
Clasper outer length | 8.3 | 3.1 | 9.9 | 7.6 | 3.1 | 2.5 | 2.7 | 2.6 | 0.2 |
Clasper inner length | 11.7 | 5.9 | 13.6 | 10.8 | 3.4 | 5.0 | 6.1 | 5.6 | 0.8 |
Clasper base width | 2.4 | 1.1 | 3.0 | 2.3 | 0.9 | 1.1 | 1.3 | 1.2 | 0.1 |
Anal fin—length | 8.2 | 7.3 | 7.8 | 7.6 | 0.3 | 7.5 | 10.8 | 8.7 | 1.0 |
Anal fin—base | 4.9 | 4.2 | 5.0 | 4.7 | 0.3 | 5.2 | 7.2 | 5.8 | 0.7 |
Anal fin—height | 4.7 | 3.8 | 5.0 | 4.4 | 0.5 | 4.1 | 5.3 | 4.6 | 0.4 |
Anal fin—anterior margin | 9.4 | 8.4 | 9.2 | 8.9 | 0.4 | 8.8 | 10.8 | 9.9 | 0.8 |
Anal fin—posterior margin | 4.0 | 3.2 | 4.5 | 4.1 | 0.5 | 3.6 | 4.8 | 4.1 | 0.5 |
Anal fin—inner margin | 3.1 | 2.2 | 4.0 | 3.0 | 0.6 | 2.6 | 3.7 | 3.2 | 0.5 |
Head height | 12.9 | 12.0 | 13.8 | 13.0 | 0.6 | 11.1 | 12.0 | 11.6 | 0.3 |
Trunk height | 13.5 | 12.8 | 15.0 | 13.9 | 0.8 | 10.6 | 13.6 | 12.4 | 1.1 |
Abdomen height | 14.0 | 11.2 | 13.8 | 13.0 | 1.1 | 11.2 | 13.1 | 11.9 | 0.8 |
Tail height | 7.6 | 7.1 | 8.5 | 7.8 | 0.5 | 6.5 | 8.6 | 7.6 | 0.7 |
Caudal peduncle height | 3.1 | 3.2 | 3.5 | 3.3 | 0.2 | 2.7 | 3.7 | 3.4 | 0.4 |
Head width | 12.2 | 11.1 | 13.8 | 12.7 | 0.9 | 11.8 | 14.1 | 12.8 | 0.9 |
Head width at anterior nostrils | 7.8 | 6.5 | 7.8 | 7.2 | 0.6 | 7.3 | 9.2 | 8.2 | 0.8 |
Head width at anterior mouth | 8.9 | 7.5 | 9.4 | 8.7 | 0.7 | 9.4 | 10.6 | 10.0 | 0.5 |
Trunk width | 12.1 | 9.7 | 13.0 | 12.1 | 1.2 | 9.5 | 13.5 | 11.4 | 1.5 |
Abdomen width | 11.5 | 9.5 | 13.1 | 11.6 | 1.2 | 8.3 | 12.6 | 10.8 | 1.5 |
Tail width | 7.3 | 6.3 | 8.2 | 7.4 | 0.6 | 4.7 | 7.6 | 6.3 | 1.1 |
Caudal peduncle width | 3.5 | 2.6 | 4.1 | 3.3 | 0.5 | 2.4 | 3.5 | 2.9 | 0.4 |
Second dorsal fin insertion to anal fin origin | 2.2 | 1.5 | 2.9 | 2.2 | 0.5 | 0.5 | 2.7 | 1.3 | 0.8 |
Anal fin insertion to ventral caudal fin origin | 12.4 | 10.4 | 14.0 | 12.4 | 1.2 | 10.3 | 11.9 | 11.0 | 0.6 |
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White, W.T.; Mollen, F.H.; O’Neill, H.L.; Yang, L.; Naylor, G.J.P. Species in Disguise: A New Species of Hornshark from Northern Australia (Heterodontiformes: Heterodontidae). Diversity 2023, 15, 849. https://doi.org/10.3390/d15070849
White WT, Mollen FH, O’Neill HL, Yang L, Naylor GJP. Species in Disguise: A New Species of Hornshark from Northern Australia (Heterodontiformes: Heterodontidae). Diversity. 2023; 15(7):849. https://doi.org/10.3390/d15070849
Chicago/Turabian StyleWhite, William T., Frederik H. Mollen, Helen L. O’Neill, Lei Yang, and Gavin J. P. Naylor. 2023. "Species in Disguise: A New Species of Hornshark from Northern Australia (Heterodontiformes: Heterodontidae)" Diversity 15, no. 7: 849. https://doi.org/10.3390/d15070849