Platax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

First specimen-based and genetic record of the little gulper shark
Centrophorus uyato (Rafinesque, 1810) (Squaliformes:
Centrophoridae) from Southwestern Taiwan
Shing-Lai Ng1, Kwang-Ming Liu2,3,4 and Shoou-Jeng Joung1,3*
1

Department of Environmental Biology and Fisheries Science, National Taiwan Ocean University, Keelung, Taiwan

2

Center of Excellence for the Oceans, National Taiwan Ocean University, Keelung, Taiwan

3

George Chen Shark Research Center, National Taiwan Ocean University, Keelung, Taiwan

4

Institute of Marine Affairs and Resource Management, National Taiwan Ocean University, Keelung, Taiwan*

Corresponding author. Email: f0010@mail.ntou.edu.tw

Abstract
The little gulper shark Centrophorus uyato was first described off the coast of Italy>200 years ago by Rafinesque (1810). While having a broad distribution, including Taiwan,no reliable specimen-based or genetic records are known in-country. Herein we haveprovided a morphological description of a juvenile specimen collected off Southwestern
Taiwan and successfully amplified a 1,044 bp fragment of the mitochondrial NADH2 gene.
The resulting morphometric and genetic characteristics agree well with specimens
described elsewhere except for that the Taiwanese juvenile possessed only 112 total centra(vs. 113–120).
Keywords: centrophorid shark, diversity, elasmobranch, mitochondrial DNA, taxonomy

uncommon in deep-sea fisheries’ bycatch,and their oil-rich livers are sometimes
harvested (Cheng et al., 1979; Bañón et al.,2006). Owing to their presumably
conserved life history traits and possibly
high susceptibility to fishing pressure,most species are listed as vulnerable tocritically endangered on the IUCN “red list”(available at: https://www.iucnredlist.org/
search?query=centrophorsu&searchType=species).

Introduction
The gulper shark genus Centrophorus
is a group of medium-sized (100–180 cm
total length [TL]), deep-sea sharks, withmost species hypothesized to be widelydispersed across the global oceans (Ebert
et al., 2021). One species, C. squamosus,
was recorded in the abyss (Priede et al.,2006), though they are typically found at
depths of 200 to 1,000 m. They are not
31

FPlatax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Gulper sharks present a challenge totaxonomists due to 1) their superficiallysimilar appearance, 2) the scarcity of intact
type specimens, 3) brief originaldescriptions, and 4) their complicatednomenclatural history (White, 2008).
Recently, these taxonomic uncertainties
were partially addressed (White et al.,2013, 2017), with Centrophorus uyato(Rafinesque, 1810) being redescribed by
White et al. (2022) based on a neotype
captured near the type locality. The authors
also reported a nearly circum-global
distribution for this species based on
Compagno (1984) and Serena (2005).
However, no voucher specimens from
Taiwan are known, and White et al. (2022)
did not describe specimens from theNorthwestern Pacific. Herein we describe
a presumed neonate centrophorid recently
collected off of Southwestern Taiwan. Its
morphology and genetic structure suggestthat it is C. uyato, which would represent
the first confirmed record from Taiwan (&possibly the entirety of the Northwestern
Pacific). We have described its
morphology and genetics in detail in thisarticle.

ervation in 70% ethanol. Methods formorphological characterization followed
White et al. (2022), with measurements directly taken by either digital calipers or a tape ruler (precision= 0.1mm).
Proportional measurements are expressed as percentage of TL. Morphometriccomparisons, including the neotype aswell as 16 small specimens (<700 mm
TL), were made with data from Whiteet al. (2022). X-rays were taken in Na
tional Taiwan Ocean University’s (NTOU) Department of Aquaculture, andthe specimen was deposited in the Pis
ces collection of the National Museumof Marine Biology and Aquarium (NM
MBA).
DNA
barcodingwiththemitochondrial NADH dehydrogenasesubunit 2 (NADH2) marker was conducted
for species confirmation. A small piece ofwhite muscle below the second dorsal-finbase was removed from the specimen andpreserved in 95% ethanol at -20˚C until
date of DNA extraction. DNA wasextracted with the EasyPure genomic DNA
mini-kit EP500 (Bioman, Taiwan) per the
manufacturer’s recommendations. A~1,200 bp fragment was PCR-amplified
using the primers ILEM-Mustelus (5'AAG-GAC-CAC-TTT-GAT-AGA-GT-3')
and ASN-Mustelus (5'-AAC-GCT-TAGCTG-TTA-ATT-AA-3') of Naylor et al.
(2005). Each 15 μl mixture contained 7.5
μl of PowerAmp 2X PCRmix-Green

Materials and methods
The sole specimen was collected f
rom a commercial bottom trawler oper
ated off Southwestern Taiwan (approxi
mate coordinates mentioned below). It
was fixed in 10% formalin before pres32

FPlatax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

buffer (Bioman), 0.6 μl of each primer(stock=10 μM), 1 μl of template DNA, and5.3 μl of ddH20. The mixture wasdenatured at 94˚C for 3 min, thensubjected to 35 cycles of denaturation at
94°C for 30 s, annealing at 48°C for 30 s,
and extension at 72°C for 90 s, and finallya single extension at 72˚C for 10 min. The
PCR product was sequenced by Genomics(Xizhi, Taiwan).
The resulting DNA sequence wasassembled and trimmed using Bioedit v.
7.2.5. (USA). The sequences from C.
atromarginatus (GN14669), C. granulosus(GN11780), C. longipinnis (GN10190), C.
squamosus (GN6514, GN11780), C.
tessellatus (GN10527, NG184), C. uyato(GN13664, GN14988, & GN17096),
Deania calcea (GN1709), and Etmopterus
spinax (GN5164) were downloaded from
Genbank(www.ncbi.nlm.nih.gov/genbank/) for construction of a phylogenetic tree.
The sequences were aligned, and genetic
distances were calculated; MEGA 11 wasused to identify the best-fitting substitution
model for phylogenetic inference(minimum AICc). The length of thealigned sequences was 1,044 bp after
trimming. A maximum likelihood
phylogenetic tree based on the best-fitting
Tamura-Nei model with + Γ + I
substitution (Tamura & Nei, 1993) with1,000 bootstraps (i.e., pseudo-replicates)
was constructed, and Deania calcea andEtmopterus spinax were the outgroups.

The sequence was deposited in GenBank
under accession number OQ909542.

Results
Centrophoridae
Centrophorus uyato (Rafinesque, 1810)
Figs. 1–3
Specimen examined. 1 specimen.
NMMB-P37946, tissue collection number
NG319 (Genbank accession number
OQ909542.1), 393 mm TL neonate female,off Donggang, Southwestern Taiwan, ca.
22°N, 120°E, 12 July 2022.
Description of the Taiwanese specimen(see Table 1 for morphometric data.). Body
slightly elongate, fusiform, and deepest
near pectoral fin insertion. Headmoderately long, length 26% of TL,
somewhat robust, width graduallydepressed in front of spiracles, oval incross section towards pectoral fin origin,and height 69% of width. Snout very short,narrowly rounded laterally, bluntly
rounded dorsally (Fig. 2A), and preorbital
length 97% of interorbital space (prenarial
length 42% of preoral length). Eye 77% of
preorbital length, 304% of height, andnotched posteriorly (though not extending
to spiracle). Spiracle rather large and teardrop shaped. Gill openings directed
antero-ventrally from top to head, subequal in size, with the 5th slightly longer
than the rest: height of 5th gill slit 106% of
orbit height.

33

FPlatax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Fig. 1. Centrophorus uyato NMMB-P37946, 393 mm TL neonate female: A. fresh; B. after
preservation. Scale bar=50 mm.
bottom, moderate, elliptical, length 25% of
Nostrils small, nearly transverse, andlength 51% of internarial space; anterior
nasal flap narrowly subtriangular, with arudimentary mesial lobe (internarial space
34% of preoral length).

teeth serrations very poorly defined on

Mouth very weakly arched (Fig. 2B),

3B); medial cusp relatively pointed; nowidth 71% of preoral length; upper labial

lateral cusps; ridges present on anterior

furrows slightly longer than lower ones;

crown, not extending onto cusp (except for

post-oral groove present, more than twice

the central ridge, which extends to the tip

the length of the upper labial furrows andof the cusp). First dorsal fin relatively

extending postero-laterally from angle ofsmall, with apex broadly rounded (Fig.

jaws. Upper teeth with two straight,

2C); anterior margin slightly convex;

symmetrical teeth at the symphyseal area;

posterior margin slightly concave; free

upper tooth shape conical and somewhat

rear tip thicker basally and long; inner

narrow, with peg-shaped roots and amargin straight; insertion of base about

distinct apron along basal face; anterior

midway between pectoral and pelvic fin

and lateral teeth with oblique cusps,

origin; fin-spine origin just posterior to

becoming more oblique posteriorly. Lower

pectoral-fin base’s end; spine exposed

mesial cutting edge; lower jaw teeth
interlocked, with strongly oblique cups
forming a cutting edge; cusps notcheddistally (Fig. 3A). Dermal denticles small,slightly pointed, and not overlapping (Fig.

34

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Platax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Fig. 2. Details of Centrophorus uyato specimen NMMB-P37946, a 393-mm (TL) neonate
female: A. dorsal view of the head; B. ventral view of the head; C. 1st dorsal fin; D.
2nd dorsal fin.

Fig. 3. Details of Centrophorus uyato specimen NMMB-P37946, a 393-mm TL neonate
female: A. teeth from upper and lower jaw; B. dermal denticles. Scale bar=1 mm.

35FPlatax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

anteriorly just below junction of spine andanterior margin straight, posterior marginsoft portion of fin; soft portion of fin

slightly concave, apex broadly rounded,

connected at about the level of two thirds

and free rear tip acute. Caudal peduncleof the total spine length; exposed spine

rather long, laterally compressed, tapering

length 63% of fin height and 72% of

towards caudal fin; pelvic-caudal space

exposed 2nd dorsal fin spine length.

47% of pectoral-pelvic space and 52% ofSecond dorsal fin slightly smaller than first,

pre-pectoral length; ventral groove poorly

its height 81% of 1st dorsal fin height (Fig.

developed; lateral keels absent; dorsal-

2D); anterior margin slightly convex andcaudal space 42% of inter-dorsal length;

apex somewhat rounded; posterior margin

precaudal pits absent. Caudal fin fairly

weakly concave; free rear tip considerably

long, caudal fork distinct, with upper post-

elongated, with its inner margin length

ventral margin slightly convex; terminal

151% of fin height; exposed spine length

lobe deep; apex of lower lobe angular;

109% of fin height; fin spine origin about

dorsal-caudal margin 83% of head length;

midway to pelvic fin free rear tip; 2ndlength of caudal pre-ventral margin 96% ofdorsal-spine base broad and tapering

caudal fork length.

distally; spine exposed anteriorly justMonospondylous pre-caudal centra

below connection of the spine and the soft

57, diplospondylous precaudal centra 27,

portion of the fin; soft portion of fin

precaudal centra 84, caudal centra 28, andconnected at about the level of half of thetotal vertebral centra 112. Tooth file count:total spine length; inter-dorsal space 57%

upper jaw 18+19, 37 and lower jaw

of pre-first dorsal fin length; inter-dorsal

14+16,30.

groove not prominent.

Coloration. When fresh dorsal and

Pectoral fin quite large, with anterior

dorsolateral surfaces uniformly purplish

and inner margins almost straight; anterior

grey;

margin length 74% of pectoral fin length;

dorsolateral and ventral color demarcatedapex rounded; posterior margin slightly

on head and lateral side before pelvic-fin

concave;

substantiallybase, extending below lower margin of eye,elongated, inner margin length 73% of

below upper margin of gill slits, and apectoral fin length, almost reaching apex

horizontal line through pectoral-fin origin.

of 1st dorsal fin; base very short, length

A dark grey blotch present above gill slits.

52% of anterior margin length. Pelvic fins

Pectoral- and pelvic-fin edges mostly

moderately

triangular,translucent, dorsal fin-tips with prominent,

length 86% of 2nd dorsal fin length,

narrow black edges, caudal-fin postventral

free

rear

large,

tip

broadly

36

F

ventralsidesomehow

paler;

Platax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

margin dark distally, terminal lobe with aembryonic specimen identified as C. uyato

black edge. (in preservative) body color

was found in the NMMBA (NMMB-

turns slightly darker.

P005590; Fig. 5), yet it has beenSize. Largest reliable specimen was 1,125

reidentified herein as a Squalus species

mm TL (330-461 mm TL at birth & mature

based on the absence of a caudal terminal

at 745–850 & 850–994 mm TL for males

lobe. Therefore, our specimen represents& females, respectively; White et al.,the first specimen-based record from2022). Our specimen is possibly a neonate

Taiwan.

given its size (393 mm TL) and presenceIn

Taiwanseven

centrophouridof umbilical scar.

species are recognized: C. atromarginatus

Distribution. Widely distributed fromGarman, 1913, C. granulosus Bloch &tropical to sub-boreal waters (except in the

Schneider, 1801, C. longipinnis White,

Eastern Pacific), at a depth range of 100 to

Ebert, & Naylor, 2017, C. moluccensis

1,490 m (White et al., 2022). The present

Bleeker, 1860, C. squamosus Bonnaterre,

specimen was captured at around 400 m.

1788, C. uyato, and an unresolved

Molecular

molecular“isodon/tessellatus” species (Ebert et al.,analysis based on a NADH2 marker

2013; White et al., 2022). C. uyato is

confirmed the identity of our specimen,

distinguished from C. granulosus and C.

and the genetic distance with other C.

squamosus by its flat, pointed (juveniles),uyato specimens (including the neotype)or pavement-like (adults) dermal denticles

ranged from 0.6 to 0.8%. Based on our(vs. raised & tear drop-shaped), making its

results, C. uyato is most closely related toskin appear smooth (vs. rather rough); its

C. atromarginatus, with a 4.1% genetic

pectoral fin’s free rear tip is also greatly

distance (98% bootstrap support; Fig. 4).

elongated (vs. barely produced). It differs

Remarks. For details of synonymies of C.

from C. longipinnis by having a muchuyato, please see White et al. (2022). The

shorter 1st dorsal fin base: length 10-14%

occurrence of C. uyato in Taiwan was

TL (vs. very long: length 16-23% TL). It

reported previously (e.g., Compagno,

differs from C. moluccensis by having a

1984; Hsu & Joung, 2004; Serena, 2005),

taller 2nd dorsal fin: height 4-6% TL (vs.

yet no specimen was examined. Shen &3-4% TL; White et al., 2013, 2017, 2022).

Wu (2011: 83) provided a photo of a gulper

Our specimen is clearly distinct from C.

shark identified as C. uyato, yet it likely

atromarginatus and the isodon/tessellatus

represents C. moluccensis based on thespecies (as C. tessellatus in Fig. 4)

analysis.

The

reduced size of its 2nd dorsal fin. An
37F

Platax 20: 31-42, 2023
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Fig. 4. Maximum likelihood phylogenetic tree generated by the best-fitting Tamura-Nei
model (with + Γ + I substitution), based on NADH2 sequences of selected
Centrophorus species. Numbers at nodes represent bootstrap values in percentages.

Bold characters represent sequences generated in the present study.

Fig. 5. Preserved Squalus sp. specimen, NMMB-P05590, a 93-mm TL embryonic malethat was previously misidentified as Centrophorus uyato. Scale bar = 10 mm.

38FPlatax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Tab. 1. Proportional measurements (as percentages of total length [TL]) for theCentrophorus uyato specimen of the present study, and measurements of the
neotype and 16 other specimens obtained from White et al. (2022).
Reference

Present study

White et al. (2022)

Specimen

NMMB-P37946

BMNH 2021.10.4.1 (neotype)

Characteristic
TL (mm)Precaudal length
Prenarial length
Preoral length
Preorbital length
Prespiracle length
Prebranchial length
Head length
Prepectoral length
Prepelvic length
Snout-anterior vent length
Pre-D1 length
Pre-D2 lengthInter-dorsal space
D2-caudal space
Pectoral-pelvic space
Pelvic-caudal space
Orbit length
Orbit height
Interorbital space
Nostril widthInternarial length
Spiracle length
Mouth width
Upper labial furrow length
1st gill slit height
Head height
Head width
Head width nostril
Abdomen width
Trunk height
Trunk width
Tail width
Caudal peduncle height
Caudal peduncle width
Pectoral fin anterior margin length
Pectoral fin base length
Pectoral fin height
Pectoral fin inner margin length
Pelvic fin length
Pelvic fin heightPelvic fin inner margin length
D1 length
D1 anterior margin length
D1 base length
D1 height
D1 inner margin length
D1 posterior margin length
D2 length
D2 anterior margin length

n=1
393
74
3.7
11.5
6.3
15.6
22
26.1
25.4
56.9
59.3
35.7
63.4
20.5
8.5
28.1
13.1
6.4
2.1
8.5
2
4
1.4
8.2
1.7
2.1
8.4
12.2
8.1
7
12
7.1
4.1
4
2.2
11.2
5.8
8.8
11
10.3
4.9
5.9
16.3
12.6
10.7
4.3
6
8.4
12
9.6

n=1
983
79.3
3.7
9.5
5.3
12.1
18.5
22.3
22.1
57.1
28.7
64.6
23.2
6.4
31.3
13.9
5.3
1.4
8.3
1.6
3.8
1.2
8.3
1.8
2
9.6
11.1
6.1
6
3.7
2.9
12.3
5.8
8.4
12.3
11.2
5.6
5.8
19
12.6
13.5
5.8
5.8
9.3
12
8.6

39

F

Min
394
75.9
3.7
9.4
5.7
12.9
18.8
20.9
21.1
53
55.1
28.2
59.3
18.2
6.6
24.4
12.4
6
1.6
7.5
1.6
3
1.5
7.9
1.7
1.7
8.9
10.4
6.1
6.6
9.2
6.4
3.8
3.3
2
11.2
3.8
7.8
10.6
9.8
4.5
5
16.8
11.5
10.9
5.7
5.7
8
11.8
9

Other
specimens
Max
666
79.6
4.7
11.8
7.2
14.9
21.9
25.1
24.9
56.6
58.6
31.5
63.8
22.4
8.7
30.6
14.7
7.3
2.7
10.8
2.3
4.4
2.3
10.1
2.6
2.8
10.8
13.5
7.8
10.6
12
11.7
5.2
4.1
2.9
13.3
5.5
9.2
13.4
11.7
5.8
6.7
20.8
14.9
13.9
7.3
7
10.6
14.9
11.8

Head height
Head width
Head width nostril
Abdomen width
Trunk height
Trunk width
Tail width
Caudal peduncle height
Caudal peduncle width
Pectoral fin anterior margin length
Pectoral fin base length
Pectoral fin height
Pectoral fin inner margin length
Pelvic fin length
Pelvic fin heightPelvic fin inner margin length
D1 length
D1 anterior margin length
D1 base length
D1 height
D1 inner margin length
D1 posterior margin length
D2 length
D2 anterior margin length
D2 base length
D2 height
D2 inner margin length
D2 posterior margin length
Caudal dorsal margin
Caudal fork width
Caudal fork length
Caudal preventral margin length
Caudal lower post-ventral margin
Caudal upper post-ventral margin
Caudal fin subterminal margin
Caudal fin terminal lobe length
D1 midpoint-pectoral fin insertion
D1 midpoint-pelvic fin origin
Pelvic fin midpoint-D1 insertion
Pelvic fin midpoint-D2 origin
D1 exposed spine length
D2 exposed spine length

8.4
9.6
12.2
11.1
8.1
6.1
7
12
7.1
4.1
6
4
3.7
2.2
2.9
11.2
12.3
5.8
5.8
Platax 20: 31-42, 2023
8.8
8.4
doi: 10.29926/platax.202312_18.0003
11
12.3
10.3
11.2
4.9
5.6
5.9
5.8
16.3
19
12.6
12.6
10.7
13.5
4.3
5.8
6
5.8
8.4
9.3
12
12
9.6
8.6
8.2
8.3
4.5
4.7
5.3
4.1
4.9
6.3
21.6
20.1
10.3
7
12.8
12.2
12.3
11.9
3.9
4.6
8.9
8.2
4.3
2.9
8.6
8.9
12.3
9.9
19.5
20.9
18.9
17.7
7.2
5.7
2.7
2.4
3.8
2.1

8.9
10.4
6.1
6.6
9.2
6.4
3.8
3.3
2
11.2
3.8
7.8
10.6
9.8
4.5
5
16.8
11.5
10.9
5.7
5.7
8
11.8
9
7.6
4.2
3.5
5.7
19.8
6.4
11.9
12
3.3
7.4
1.7
7.4
5.9
14.8
13.2
1.8
2.1
2.7

10.8
13.5
7.8
10.6
12
11.7
5.2
4.1
2.9
13.3
5.5
9.2
13.4
11.7
5.8
6.7
20.8
14.9
13.9
7.3
7
10.6
14.9
11.8
10.2
6.1
5.2
8.7
22.8
8
14.1
14.7
5.2
9.7
4.5
9.9
10.5
21.1
17.5
5.7
3.2
4

based on the molecular data: genetic dis

buyer in Donggang) for kindly providing

tances of 4.1 and 6%, respectively. Ho

the specimen. We thank Q.-Y. Wu (NTOU)

wever, morphological differences are nofor her assistance in the moleculart pronounced (White et al., 2022), andexperiment, as well as Dr. H.-M. Chen anda taxonomic revision is needed to fully

S.-H. Yeh for their assistances in taking thecharacterize the within-genus morpholog

x-rays. We also thank P.-N. Lee (NMMBA)

ical variability.

for curatorial assistance.

Comparative material. Squalus sp.1
specimen (Fig. 5). NMMB-P05590, 93

References

mm TL embryonic male, off Daxi,

Bañón, R., C. Piñeiro & M. Casas. 2006.

Northeastern

Taiwan,ca.

24°90'N,

Biological aspects of deep-water sharks

122°00'E, 1 February 1983, coll. H.-Y.

Centroscymnus

Wang.

Centrophorus squamosus in Galician

coelolepis

andwaters (north-western Spain). Journal of

Acknowledgements

the Marine Biological Association of the

We are grateful to Z.-L. Fang (fishUnited Kingdom 86(4):843-846.

40

F

Platax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Cheng, T.H., K.S. Yeh & C.T. Chen. 1979.

FAO, Rome.

Deep-sea shark fishery in Taiwan. JCRR

Shen, S.C. & K.Y. Wu. 2011. Fishes of Taiwan.

Fisheries Series 36:122-130.
Compagno,

L.J.V.

1984.

FAO

National Museum of Marine Biology andSpeciesAquarium, Pingtung, Taiwan. 896 pp.

Catalogue. Vol. 4, Sharks of the World.

Tamura, K. & M. Nei. 1993. Estimation of the

An annotated and illustrated catalogue of

number of nucleotide substitutions in the

shark species known to date. FAO

control region of mitochondrial DNA in

Fisheries Synopsis No. 125. vol. 4, pt. 1

humans and chimpanzees. Molecular(noncarcharhinoids), pp. viii, 1-250, pt. 2

Biology and Evolution 10:512-526.

(carcharhiniformes), pp. 251-655.

White, W.T. 2008. Family Centrophoridae,

Hsu, H.H. & S.J. Joung. 2004. Four new

Gulper sharks. In: Gomon, M., D. Bray &records of cartilaginous fishes fromR. Kuiter(Eds.), Fishes of Australia’s

Taiwan. Journal of the Fisheries SocietySouthern Coast. New Holland Publishersof Taiwan 31:183-189.

(Australia), Victoria, pp. 85-88.

Naylor, G.J.P., J.A. Ryburn, O. Fedrigo & A.

White, W.T., D.A. Ebert & G.J.P. Naylor. 2017.

Lopez. 2005. Phylogenetic relationships

Revision of the genus Centrophorus

among the major lineages of modern(Squaliformes: Centrophoridae): Part 2-elasmobranchs. In: Hamlett, W.C. (Ed.),

Description of two new species of

Reproductive Biology and Phylogeny ofCentrophorus and clarification of the

Chondrichthyes - Sharks, Batoids andstatusChimaeras.

Barbosa du Bocage & de Brito Capello,

SciencePublisher,Inc.,

Enfield (NH), USA, pp. 1-25.

ofCentrophorus

lusitanicus

1864. Zootaxa 4344(1):86-114.

Priede, I.G., R. Froese, D.M. Bailey, O.A.

White, W.T., D.A. Ebert, G.J.P. Naylor, H.C.

Bergstad, M.A. Collins, J.E. Dyb, C.

Ho, P. Clerkin, A. Verissimo & C. Cotton.

Henriques, E.G. Jones & N. King. 2006.

2013.

The absence of sharks from abyssal

Centrophorus

regions

Centrophoridae): Part 1-Redescription of

of

the

world'soceans.

Revision

of

the

genus

(Squaliformes:

Proceedings of the Royal Society B:

Centrophorus

Biological

Schneider), a senior synonym of C. acus

Sciences273(1592):1435-

1441.

(Bloch&Garman and C. niaukang Teng. Zootaxa

Serena, F. 2005. Field identification guide to3752(1):35-72.

the sharks and rays of the Mediterranean
andgranulosus

Black

Sea.

FAO

White, W.T., J. Guallart, D.A. Ebert, G.J.

SpeciesNaylor, A. Verissimo, C.F. Cotton, M.

Identification Guide for Fishery Purposes.

Harris, F. Serena, & S. P. Iglésias. 2022.
41

F

Platax 20: 31-42, 2023
doi: 10.29926/platax.202312_18.0003

Revision of the genus Centrophorus(Squaliformes: Centrophoridae): Part 3Redescription of Centrophorus uyato(Rafinesque) with a discussion of its
complicatednomenclatural

history.

Zootaxa 5155(1):1-51.

42

F