Fig 1.
Anterior margin of nostrils in Cirrhigaleus (A–C, left side) and Squalus (D–E, right side), showing its innervation and internal support. A: C. barbifer, HUMZ 95177, juvenile female, 584 mm TL; B: C. australis, CSIRO H 7042–04, juvenile female, 605 mm TL; C: C. asper, SAIAB 6092, neonate female, 275 mm TL; D: Squalus suckleyi, CAS 21971 (cleared and stained), neonate female, 290 mm TL. Abbreviations: buVII: buccopharyngeal branch of facial nerve; cnab: fleshy core; mnl: medial nasal lobe supported by cartilage (in blue); nab: nasal barbel; nc: nasal capsule; r: rostrum; snf: subnasal fenestra. Scale bars: 5 mm (A); 1 mm (B–D).
Fig 2.
Neurocranium of species of Cirrhigaleus and Squalus in dorsal view.
A: C. barbifer, HUMZ 95177, juvenile female, 584 mm TL; B: C. australis, CSIRO H7042-04, juvenile female, 605 mm TL; C: S. suckleyi, HUMZ 87643, adult male, 665mm TL; D: S. megalops, AMS I46093-001, adult male, 650 mm TL; E: S. mitsukurii, NSMT P-44381, juvenile male, 770 mm TL. Abbreviations: ec: ethmoidal canal of ophthalmicus superficialis nerve; elf: endolymphatic foramen; ep: epiphysial pit; foop: profundus canal for the ophthalmicus profundus nerve; nc: nasal capsule; oc: otic capsule; pep: preorbital process; plf: perilymphatic foramen; poc: preorbital canal of superficial ophthalmic nerve; pop: postorbital process; r: rostrum; sep: supraethmoidal process. Scale bars: 10mm.
Fig 3.
Detail of ethmoidal region of the neurocranium in anterior dorsal view.
A: C. australis, CSIRO H7042-04, juvenile female, 605 mm TL; B: C. asper, SAM 38269, adult female, 1045 mm TL; C: C. asper, UFPB 11864, juvenile female, 970 mm TL; D: S. suckleyi, SAM 38346, adult female, 850 mm TL; E: S. albifrons, MZUSP 121272, adult male, 760 mm TL; F: S. montalbani, MZUSP121270, adult male, 713 mm TL. Abbreviations: ep: epiphyseal pit; sep: supraethmoidal process. Scale bars: 2mm (A,E,F), 5mm (B–D).
Fig 4.
Neurocranium of species of Cirrhigaleus and Squalus in ventral view.
A: C. barbifer, HUMZ 95177, juvenile female, 584 mm TL; B: C. australis, CSIRO H7042-04, juvenile female, 605 mm TL; C: S. suckleyi, HUMZ 87643, adult male, 665mm TL; D: S. megalops, AMS I46093-001, adult male, 650 mm TL; E: S. mitsukurii, NSMT P-44381, juvenile male, 770 mm TL. Abbreviations: aoc: antorbital cartilagem; btp: basitrabecular process; nab: nasal barbel; nc: nasal capsule; pop: postorbital process; r: rostrum; rk: rostral keel: sec: subethmoid chamber; snf: subnasal fenestra; lra: lateral rostral appendage; mra: medial rostral appendage. Scale bars: 10mm.
Fig 5.
Detail of rostral process of Squalus species in ventral view, showing rostral appendages.
A: S. albifrons, MZUSP 121272, adult male, 760 mm TL; B: S. montalbani, MZUSP121270, adult male, 713 mm TL. Abbreviations: lra: lateral rostral appendage; mra: medial rostral appendage; mrp: median rostral prominence. Scale bars: 1 mm.
Fig 6.
Pectoral skeleton of C. australis (CSIRO H7042-04, juvenile female, 605 mm TL), showing pectoral girdle and pectoral basals.
A: anterior view; B: posterior view; C: lateral view; D: dorsal view. Abbreviations: co: coracoid bar; df: diazonal foramen; mes: mesopterygium; mtp: metapterygium; pcf: pectoral fin; pcr: pectoral fin radials; pro: propterygium; ptp: posterior triangular process; scl: scapula; scp: scapular process. Scale bars: 5mm (A–D); 2mm (E,F).
Fig 7.
Detail of coracoid bar of Cirrhigaleus and Squalus species in anterior (A,D,G,J,M), posterior (B,E,H,K,N) and lateral views (C,F,I,L,O), showing the pectoral articular region between the pectoral girdle and pectoral basals. A–C: C. australis, CSIRO H7042-04, juvenile female, 605 mm TL; D–F: C. asper, SAM 38269, adult female, 1045 mm TL; G–I: S. acanthias, SAM 38276, adult male, 670mmTL; J–L: S. albifrons, MZUSP 121272, adult male, 760 mm TL; M–O: S. montalbani, MZUSP121270, adult male, 713 mm TL. Abbreviations: bse: barrel-shaped elements; df: diazonal foramen; fpr: facet for propterygium; msc: mesocondyle; mtc: metacondyle; pc: procondyle; scl: scapula. Scale bars: 2 mm.
Fig 8.
Puboischiadic bar (right side) in dorsal (A,D,G,J), ventral (B,E,H), posterior (C,F,I) and medial (K) views showing the pelvic articular region with the pelvic fin. A–C: C. australis, CSIRO H7042-04, juvenile female, 605 mm TL; D–F: C. asper, SAM 38269, adult female, 1045 mm TL; G–I: S. montalbani, MZUSP121270, adult male, 713 mm TL; J,K: S. albifrons, MZUSP 121272, adult male, 760 mm TL. Abbreviations: abc: condyle for the anterior pelvic basal; abv: anterior pelvic basal; bp: basipterygium; cbp: condyle for the basipterygium; fpb: facet for the basipterygium; fvn: foramen for ventral fin nerve; lpp: lateral prepelvic proces; plp: posterior-lateral process; pub: puboischiadic bar. Scale bars: 1mm.
Fig 9.
Detail of cartilages of the clasper in Squalus species in dorsal (A,D,G,J), ventral (B,E,H,K) and terminal dorsal (C,F,I,L) views. A–C: S. acanthias, SAM 38276, adult male, 670mmTL; D–F: S. brevirostris, HUMZ 189762, adult male, 433 mm TL; G–I: S. albifrons, MZUSP 121272, adult male, 760 mm TL; J–L: S. montalbani, MZUSP121270, adult male, 713 mm TL. Abbreviations: ax: axial cartilage; bp: basipterygium; b1: first intermediate segment; b2: second intermediate segment; mp: mesial process; rd: dorsal marginal cartilage; rl: pelvic radials; td: dorsal terminal cartilage; td2: dorsal terminal 2 cartilage; tv: ventral terminal cartilage; t3: accessory terminal 3 cartilage; β: beta cartilage. Scale bars: 1 mm.
Fig 10.
Cartilages of the clasper (left side) of C. australis, NMNZ P 38074, adult male, 1020 mm TL in dorsal view (A) and detail of terminal cartilages in ventral view (B). Abbreviations: ax: axial cartilage; b1: first intermediate segment; b2: second intermediate segment; rd: dorsal marginal cartilage; rv: ventral marginal cartilage; td: dorsal terminal cartilage; td2: dorsal terminal 2 cartilage; tv: ventral terminal cartilage; tv2: ventral terminal 2 cartilage; t3: accessory terminal 3 cartilage; β: beta cartilage. Scale bar: 10 mm.
Fig 11.
Ventral view of the head (right side) in Cirrhigaleus species showing nasal barbels and upper labial furrows.
A: C. barbifer, HUMZ 197852 (neotype), adult female, 870 mm TL; B: C. australis, AMS I 27022–001 (paratype), adult female, 1205 mm TL; C: C. asper, uncatalogued specimen, adult female, 1068 mm TL.
Fig 12.
Scanning electron microscopy of dermal denticles of species of Cirrhigaleus in coronal (A–L, except H) and lateral (H) views. C. barbifer (A–C): A: HUMZ 197852 (neotype), adult female, 870 mm TL; B: NMW 98257, adult female, 960 mm TL; C: CSIRO H 5875–09, adult female, 978 mm TL; C. australis (D–H): D: NMNZ P 38074, adult male, 1020 mm TL; E: NMNZ P 42489, juvenile male, 710 mm TL; F: CSIRO H 7048–01, adult male, 993 mm TL; G–H AMS I 45670–001, juvenile male, 630 mm TL. C. asper (I–L): I: SAIAB, 6092, juvenile female, 275 mm TL; J: SAIAB 31890, adult female, 1090 mm TL; K: SAM 39879, adult female, 1023 mm TL; L: NUPEC uncatalogued, adult female, 1270 mm TL. Scale bars: 100 μm (I); 200 μm (A,D,G,J–L); 500 μm (B,C,E,F,H).
Fig 13.
Strict consensus cladogram of the two equally most-parsimonious cladograms of 69.9 steps resulting from the analysis of 51 morphological characters (CI = 0.80; RI = 0.88).
Relative Bremer support values are given below each node.
Fig 14.
Simplified cladograms of previous phylogenetic hypotheses for Squalidae, highlighting the relationships of Cirrhigaleus species (thick red branches).
Interrelationships between the three major groups of species within Squalus are also shown. A: Shirai [22]; B: Naylor et al. [15]; C: present study. Blue rectangle: S. acanthias group; Green rectangle: S. megalops group; Yellow rectangle: S. mitsukurii group.
Fig 15.
Map of the geographical distribution of Cirrhigaleus species.
Stars: type material; dots: other material; red symbols: C. barbifer; green symbols: C. australis; yellow symbols: C. asper.
Fig 16.
Cirrhigaleus barbifer in lateral (A,B) and dorsal (C) views. A: HUMZ 197852 (neotype of C. barbifer), adult female, 870 mm TL; B,C: HUMZ 231872, adult female, 899 mm TL (photo credits: Fisheries Science Center, The Hokkaido University Museum).
Table 1.
External measurements of C. barbifer expressed as percentage of total length (%TL).
Fig 17.
Upper (A) and lower (B) teeth of C. barbifer, NMW 98257, adult female, 960 mm TL (labial view). Scale bars: 1 mm.
Fig 18.
First (a, b) and second (c, d) dorsal fins of C. barbifer, showing intraspecific regional variations. a,c: HUMZ 197852 (neotype), adult female, 870 mm TL, Japan; b,d: CSIRO H5875-09, adult female, 978 mm TL, Indonesia. Scale bars: 20 mm.
Fig 19.
Pectoral (A), pelvic (B) and caudal (C) fins of the neotype of C. barbifer, HUMZ 197852, adult female, 870 mm TL. A,B: ventral view; C: lateral view. Scale bars: 20 mm.
Fig 20.
Specimens of C. barbifer in lateral view.
A: SU 13901 (holotype of Phaenopogon barbulifer Herre, 1935), juvenile female, 730 mm TL, Japan; B: MNHN 1997–3568, adult female, 800 mm TL, New Caledonia.
Fig 21.
Specimens of C. asper in lateral view.
A: BMNH 1972.10.10.1 (holotype), adult male, 880 mm TL, Seychelles; B: SAIAB 6092, juvenile female, 275 mm TL, South Africa; C: USNM 220585, juvenile male, 245 mm TL, the U.S.A. Scale bars: 50 mm.
Table 2.
External measurements of C. asper expressed as percentage of total length (%TL).
Fig 22.
Upper teeth (labial view) of the paratype of C. asper, BMNH 1972.10.10.2, adult male, 847 mm TL.
Fig 23.
First (A) and second (B) dorsal fins of C. asper. A,B: BMNH 1972.10.10.1, adult male, 880 mm TL. Scale bars: 20 mm.
Fig 24.
Pectoral (A), pelvic (B) and caudal (C) fins of the paratype of C. asper in ventral (A,B) and lateral views (C). A,B: BMNH 1972.10.10.3, adult female, 865 mm TL; C: BMNH 1972.10.10.1, adult male, 880 mm TL. Scale bars: 20 mm.
Fig 25.
Clasper (right side) of the paratype of C. asper, BMNH 1972.10.10.2, adult male, 847 mm TL in dorsal view.
Abbreviations: ap: apopyle; cg: clasper groove; hp: hypopyle; p2: pelvic fin; rh: rhipidion.
Fig 26.
Fresh specimen of C. asper in lateral (A) and ventral (B) views, SAIAB 186460, juvenile female, 456 mm TL, South Africa. Reprinted from NRF-SAIAB under a CC BY license, with permission NRF-SAIAB, 2010.
Fig 27.
Cirrhigaleus australis in lateral view.
A: CSIRO H 7042–04, juvenile female, 605 mm TL, Australia; B: NMNZ P 34821, adult male, 971 mm TL, New Zealand.
Table 3.
External measurements of C. australis expressed as percentage of total length (%TL).
Fig 28.
Upper (A) and lower (B) teeth (labial view) of C. australis, AMS I 45670–001, juvenile male, 630 mm TL. Scale bars: 1 mm.
Fig 29.
First (A,B) and second (C,D) dorsal fins of C. australis. A,C: AMS I 42891–001 (paratype), adult female, 1085 mm TL, Australia; B,D: NMNZ P 28732, adult female, 1189 mm TL, New Zealand. Scale bars: 20 mm.
Fig 30.
Pectoral (A), pelvic (B) and caudal (C) fins of C. australis (NMNZ P 28732, adult female, 1189 mm TL) in ventral (A,B) and lateral (C) views. Scale bars: 20 mm.
Fig 31.
Clasper (left side) of C. australis, NMNZ P 38074, adult male, 1020 mm TL in dorsal view.
Abbreviations: ap: apopyle; cg: clasper groove; hp: hypopyle; p2: pelvic fin; rh: rhipidion. Scale bar: 20 mm.
Table 4.
Cranial measurements of C. barbifer, C. asper and C. australis expressed as percentage of total length of the neurocranium (% CL).