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Squatina caillieti sp. nov., a new species of angel shark (Chondrichthyes: Squatiniformes: Squatinidae) from the Philippine Islands

February 2011
Zootaxa 2759(2759):49-59

DOI:10.11646/zootaxa.2759.1.2

Project: Looking for Lost Sharks: An Exploration of Discovery

Authors:

Jon Walsh

Pacific Gas and Electric Company

David A. Ebert

Moss Landing Marine Labs

A new species of angel shark, Squatina caillieti sp. nov., is described from a single specimen collected in deepwater off Luzon in the Philippines. The new species is closest to S. formosa and S. nebulosa, but differs from its congeners based on the following characters: unfringed barbels with rod-like tips, upper lip arch semi-oval in shape, large papillae present on the inside posterior margin of the spiracles, a greater interspiracle space than interorbital space, pelvic fin-tips which reach the first dorsal origin, a short pelvic fin base, short pelvic inner margin very short, and a short pelvic posterior margin; pelvic girdle span more than 1.4 times greater than head length; dorsal fins angular, greater interdorsal space than dorsal caudal space; caudal fin lobed, very short upper postventral caudal margin. The new species is the only Squatina confirmed as occurring in the Philippines. We also comment on the biogeography of western North Pacific Squatina and provide a revised regional key to this group.

. Western North Pacific squatinid vertebral and spiral valve counts.

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Accepted by M.R. de Carvalho: 21 Dec. 2010; published: 7 Feb. 2011

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)

Zootaxa 2759: 49–59 (2011)

www.mapress.com/zootaxa/Article

Squatina caillieti sp. nov., a new species of angel shark (Chondrichthyes:

Squatiniformes: Squatinidae) from the Philippine Islands

JONATHAN H. WALSH1, DAVID A. EBERT1, 2, 3 & LEONARD J.V. COMPAGNO4

1Pacific Shark Research Center, Moss Landing Marine Laboratories, 8272 Moss Landing Road, Moss Landing, CA, 95039, U.S.A.; ph.

831-771-4400; fax 831-632-4403. E-mail: jwalsh@mlml.calstate.edu

2Research Associate, Department of Ichthyology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA.

94118, USA

3Research Associate, South African Institute for Aquatic Biodiversity, Private bag 1015, Grahamstown 6140, South Africa

4Shark Research Centre, Iziko – South African Museum, P.O. Box 91, Cape Town, 8000, South Africa

Abstract

A new species of angel shark, Squatina caillieti sp. nov., is described from a single specimen collected in deepwater off

Luzon in the Philippines. The new species is closest to S. formosa and S. nebulosa, but differs from its congeners based

on the following characters: unfringed barbels with rod-like tips, upper lip arch semi-oval in shape, large papillae present

on the inside posterior margin of the spiracles, a greater interspiracle space than interorbital space, pelvic fin-tips which

reach the first dorsal origin, a short pelvic fin base, short pelvic inner margin very short, and a short pelvic posterior mar-

gin; pelvic girdle span more than 1.4 times greater than head length; dorsal fins angular, greater interdorsal space than

dorsal caudal space; caudal fin lobed, very short upper postventral caudal margin. The new species is the only Squatina

confirmed as occurring in the Philippines. We also comment on the biogeography of western North Pacific Squatina and

provide a revised regional key to this group.

Key words: Squatinidae, Squatina caillieti, new species, Philippines

Introduction

The family Squatinidae Bonaparte, 1838, consists of a single genus and approximately 19 valid species (Ebert &

Compagno, 2011). The genus is wide ranging occurring mostly on continental shelves and upper slopes, mainly in

temperate waters, but with some species penetrating into the tropics (Compagno et al., 2005a). These ray-like,

dorso-flattened, sharks are medium-sized with most being less than 1.6 m in total length (TL). These sharks are

taken in both targeted fisheries and as by-catch in many areas of the world. Unfortunately, the differences among

this group are poorly known and until recently have received very little attention.

In the western Pacific, at least seven Squatina species are known to occur. Four valid species are recognized in

the western North Pacific (Walsh & Ebert, 2007), S. formosa Shen & Ting, 1972, S. japonica Bleeker, 1858, S. neb-

ulosa Regan, 1906, and S. tergocellatoides Chen, 1963, and three species from tropical Australian waters (Last &

White, 2008), S. albipunctata Last & White, 2008, S. australis Regan, 1906, and S. legnota Last & White, 2008. A

Squatina species tentatively identified as S. formosa, from Philippine waters (Compagno et al., 2005b) was exam-

ined and compared to Squatina material, including S. formosa, as part of a regional revision on this group (Walsh &

Ebert, 2007). Based on specimens, including type material, examined by the authors and compared to the Philip-

pines specimen it was concluded that the Squatina species previously identified as S. formosa (Compagno et al.,

2005a, b) in fact represents an undescribed species. Here we describe this new species of Squatina.

Methods

Morphometric measurements were taken following Compagno (2001), but with the addition of measurements for

the upper lip arch and the caudal fin following Walsh & Ebert (2007). Measurements were taken to the nearest 0.1

WA LS H ET AL.

mm, and are presented as a proportion of total length (TL) to facilitate direct comparison with other species. Verte-

bral counts were taken directly from digital radiographs, while tooth counts, and spiral valve counts were taken

directly from specimens. Radiographs were taken from more than one angle to limit obscurement of vertebrae from

other anatomical structures. Abbreviations for institutions and field numbers are as follows: British Museum of

Natural History, London, United Kingdom (BMNH); California Academy of Sciences, San Francisco, California,

U.S.A. (CAS); Australian National Fish Collection, Hobart, Australia (CSIRO); Hokkaido University, Laboratory

of Marine Zoology, Faculty of Fisheries, Hokkaido, Japan (HUMZ); National Taiwan University, Department of

Zoology, Taipei, Taiwan (NTUM); David A. Ebert field numbers (DAE).

Squatina caillieti sp. nov.

(Figures 1, 2, Table 1)

Holotype, CAS 226473, immature female, 328 mm total length, Taiwan Fisheries Research Institute Fishery

Researcher 1 sta. FR1-PHI-02-95, 23 September 1995, 363–385 m, Philippines, Luzon, 13°08.98–09.84'N,

124°04.72–00.01'E, collected by L.J.V. Compagno and P.R. Last.

Squatina formosa: Compagno et al., 2005a, Sharks of the World, p 142, fig, pl. 18. Compagno et al., 2005b,

Checklist of Philippine Chondrichthyes, p 56.

Diagnosis. A squatinid distinct from other western North Pacific squatinids based on the following character-

istics: unfringed barbels with rod-like tips; upper lip arch semi-oval in shape, upper lip arch height (1.2), upper lip

arch width (4.3); large papillae present on the inside posterior margin of the spiracles; a greater interspiracle space

(8.9) than interorbital space (8.6); lacking midback row of thorny tubercles; large prominent ocelli not present on

pectoral fins; pelvic fintips reaching the first dorsal origin, pelvic fin base short (12.9), pelvic inner margin very

short (8.6) and short pelvic posterior margin (16.3); pelvic girdle width (27.9) more than 1.4 times greater than

head length (18.7); dorsal fins angular, greater interdorsal space (7.4) than dorsal caudal space (6.4), subdorsal sad-

dles present; caudal fin lobed, very short upper postventral caudal margin (2.5).

Description. Dorsal surface covered with interspersed denticles of moderate roughness; ventral surface com-

paratively smooth, except for narrow bands of denticles along anterior margins of both pectoral and pelvic fins.

Head rounded, length slightly less than 0.2 times total length, with a maximum width occurring anterior of gill

openings. Moderate tubercles interspersed above mouth and eye crests, smooth oval patch above midpoint of

mouth in between eyes. Eyes are almond-shaped, closely set, with an interorbital space of 8.6; eye-spiracle distance

short (1.5). Spiracles are crescent shaped with pronounced papillae on interior along posterior margin. Interspiracle

space (8.9) is slightly greater than interorbital space (8.6). Center of upper lip arch exposed at midpoint of upper

jaw, exposure semi-oval in shape, extending dorsally approximately 0.6–0.7 of upper jaw space, upper lip height

(1.2), and upper lip arch width (4.3). Labial furrows, roughly equal in length, extending from corners of mouth

medially, with upper labial furrow partially covered with dermal folds. Distinct nasal flaps protruding from dermal

folds above mouth, two barbels protruding from each flap; inner nasal barbel rod-like with small branch protruding

ventrally near tip, inner basal portion contains little if any fringe; outer nasal barbel rod-like. Nostrils large, pro-

truding slightly, and tear-shaped. Dermal folds along exterior of head, one small lobe present at corners of mouth

extending ventrally. Mouth length about 0.3 times as long as mouth width. Dentition consisting of small, dagger-

like teeth, conical without cusplets on a broad base, in 2 orderly longitudinal rows on upper jaw, 3 rows on bottom

jaw, no teeth at symphysis, teeth by row .

Pectoral fins large and broadly rounded, originating just behind gills. Anterior margin of pectoral fin mostly

straight and about three quarters as long as pectoral length, extending to a lateral apex. Angle of lateral apex is

slightly more obtuse than 120°. Margin from lateral apex to posterior-most lobe slightly concave. Posterior lobe

broadly rounded. Pectoral inner margin is slightly less than one half of pectoral length, convex, and with a small

lobe near pectoral base.

Overall pelvic fin shape is somewhat triangular with rounded fintips. Pelvic fins originating anterior to pectoral

fin free rear tips. Pelvic fin length approximately three quarters as long as pectoral fin length. Pelvic fin base is

approximately equal to pectoral fin base. Anterior margin slightly curvilinear, extending at roughly a 45° angle

from trunk to rounded apex lateral of body, anterior margin 0.4 times as long as pelvic fin length. Pelvic girdle span

(27.9) between pelvic fin apices moderately broad, about 1.5 times head length. Posterior margin of pelvic fin

99 1010

−

SQUATINA CAILLIETI SP. NOV., A NEW SPECIES OF ANGEL SHARK

straight to posterior free tip approximately 0.7 times fin length. Pelvic inner margin is straight and short, approxi-

mately 0.4 times as long as pectoral fin length. Pelvic fin insertion furrows on ventral extend in a narrow ellipse to

anterior apogee of vent, vent is within ellipse. One pelvic fin tip extends to first dorsal origin, the other does not.

Dorsal fins slightly angular, second dorsal fin slightly smaller than first dorsal fin, with denticles covering the

whole of fins. Interdorsal space is about 1.1 times longer than dorsal caudal space. Anterior margin of both dorsal

fins straight, nearly equidistant. Dorsal bases equal, first dorsal base (4.6), second dorsal base (4.6). Apex of first

and second dorsal fins both lobed. Posterior margins straight, about 0.5–0.6 times as long as anterior margins. Inner

margin of dorsal fins straight, approximately 0.4 times as long as anterior margins.

Caudal peduncle compressed dorso-ventrally with lateral longitudinal ridges, tapering posteriorly. Caudal fin

lobe-like, markedly at dorsal apex, dorsal margin broadly rounded, about 0.8 times as long as preventral caudal fin

margin. Subterminal caudal fin margin is approximately 1.4 times longer than caudal upper post ventral margin.

Caudal lower postventral margin is slightly convex, approximately 1.5 times longer than caudal upper post ventral

margin.

Total vertebrae 137; total precaudal vertebrae 111, monospondylous vertebrae 51, diplospondylous vertebrae

60, caudal vertebrae 26. Spiral valve count: 7.

Coloration. Dorsal surface of holotype was greenish in color throughout with numerous brown spots, outlined

in white; pelvic fins with white margins; black subdorsal saddles were also present. Upon preservation, the speci-

men faded to a deep uniform brown with numerous white spots; subdorsal saddles were lost with preservation as

was the white edging on the pelvic fins; ventral side is uniformly white.

Distribution. Holotype and only known specimen caught off Luzon (13°08.98–09.84'N, 124°04.72–00.01'E),

Philippine Islands (Figure 3) at a depth of 363–385 m in a trawl.

Etymology. The new species is named in honor of Dr. Gregor Cailliet for his outstanding contributions in the

field of ichthyology, especially in the area of chondrichthyan age and growth. Pronunciation of caillieti: [kai-yā-

i]. Remarks. Although similar to other western North Pacific (WNP) squatinids, there are several characters that

distinguish S. caillieti from other regional squatinids. In our previous work on WNP squatinids (Walsh & Ebert

2007), we showed that a major character in identification of this group was the pelvic fin tips in relation to the first

dorsal fin origin. In S. caillieti the fin tips reach the dorsal origin, negating the possibility that the specimen is either

S. japonica or S. tergocellatoides. In addition, the specimen lacks midback spines (S. japonica), ocelli or fringed

barbels (S. tergocellatoides), has a lower spiral count than other WNP squatinids, and differs dramatically from

both in coloration.

Squatina caillieti is also distinct from the recently described Indonesian Squatina legnota (Last & White

2008). The pelvic fin tips of S. legnota do not reach the first dorsal fin origin, which is not the case in S. caillieti.

Also, S. legnota has a greater dorsal-caudal space (DCS) than inter-dorsal space (IDS), which is the convers condi-

tion found in S. caillieti. Additionally, S. caillieti differs slightly from S. legnota as it is a lighter brown color and

has more noticeable ocelli after preservation when the mucous layer is not present. Unfortunately, no photos of

fresh S. legnota specimens were available for comparison.

Squatina caillieti more closely resembles the other two known WNP species, S. formosa and S. nebulosa, but

several morphometrics differences are present by which these three species can be distinguished from each other.

The pelvic fin measurements for S. caillieti are smaller at the posterior margin (P2P) than S. formosa, and smaller

at the inner margin (P2I) and base (P2B) than both S. formosa and S. nebulosa. Squatina caillieti also has a mark-

edly smaller upper postventral caudal margin than both S. formosa and S. nebulosa. In addition, Squatina caillieti

also has a greater interspiracle distance (ISP) than interorbital distance (INO), a character unique among all speci-

mens examined. Among the other two species, S. formosa INO is equal to ISP, and S. nebulosa, INO is greater than

ISP (Figure 4). Squatina caillieti is also the only known WNP squatinid to have a greater interdorsal space (IDS)

than dorsal-caudal space (DCS) (Figure 5). Squatina caillieti also appears to be the only WNP squatinid with large

papillae in the inner posterior margin of the spiracles, but more specimens should be examined to validate this char-

acter. Finally, although possessing similar vertebral counts to S. formosa and S. nebulosa, S. caillieti has less caudal

vertebrae and a lower spiral valve count than those species (Table 2).

WA LS H ET AL.

FIGURE 1. Squatina caillieti sp. nov. holotype, immature female, 326 mm, photo taken of fresh specimen (A) dorsal view, (B)

ventral view, and (C) dorsal view of specimen after preservation.

SQUATINA CAILLIETI SP. NOV., A NEW SPECIES OF ANGEL SHARK

FIGURE 2. Squatina caillieti sp. nov. holotype (A) Head, (B) Dorsal fins, and (C) Caudal fin.

WA LS H ET AL.

TABLE 1. Squatina caillieti sp. nov. measurements. Total Length (TL) is given in millimeters (mm), all other measurements

are percent TL.

The observed characteristics distinguishing S. caillieti from other western Pacific squatinids were consistent

among all individuals of each species examined. Furthermore, individuals of each comparative species were com-

pared between similar sized individuals reducing variation due to ontogenetic changes in size. Although additional

specimens, particularly of S. caillieti would benefit character analysis, the study did benefit from comparison

between the holotypes of S. formosa and S. nebulosa, the new species, and additional comparative specimens of

each of these and other western Pacific squatinids.

The genus Squatina is highly diverse in the WNP with at least five species now recognized (Figure 6). All five

species based on the current knowledge of their distribution appear to be endemic to the WNP. At the present time

Measure (abbreviation) Measure (abbreviation)

Total Length (mm) (TL) 326

Pre-Caudal Length (PRC) 84.7 Pectoral Fin Inner Margin (P1I) 14.7

Pre-Orbital Length (POB) 2.5 Pectoral Fin Posterior Margin (P1P) 13.8

Pre-Spiracle Length (PSP) 6.7 Pectoral Fin Span (P1S) 43.6

Pre-Branchial Length (PG1) 15.3 Pectoral-Pelvic Space (PPS) 11.3

Head Length (HDL) 18.7 Pelvic Fin Length (P2L) 24.5

Pre-Pectoral Length (PP1) 18.7 Pelvic Fin Anterior Margin (P2A) 10.1

Pre-Pelvic Length (PP2) 41.4 Pelvic Fin Base (P2B) 12.9

Snout-Vent Length (SVL) 47.2 Pelvic Fin Height (P2H) 9.8

Pre-1st Dorsal Length (PD1) 63.5 Pelvic Fin Inner Margin (P2I) 8.6

Pre-2nd Dorsal Length (PD2) 74.8 Pelvic Fin Posterior Margin (P2P) 16.3

Mouth Length (MOL) 4.9 Pelvic Fin Span (P2S) 28.2

Mouth Width (MOW) 12.9 Pelvic-Caudal Space (PCS) 30.4

Upper Labial Furrow Length (ULA) 4.3 1st Dorsal Fin Length (D1L) 8.0

Lower Labial Furrow Length (LLA) 2.8 1st Dorsal Fin Anterior Margin (D1A) 7.4

Internarial Width (INW) 5.2 1st Dorsal Fin Base (D1B) 4.6

Outernarial Width (ONW) 5.8 1st Dorsal Fin Height (D1H) 5.5

Nostril Width (NOW) 1.8 1st Dorsal Fin Inner Margin (D1I) 2.8

Anterior Nasal Flap Length (ANF) 1.8 1st Dorsal Fin Posterior Margin (D1P) 4.6

Upper Lip Arch Width (UAW) 4.3 2nd Dorsal Fin Length (D2L) 6.4

Upper Lip Arch Height (UAH) 1.2 2nd Dorsal Fin Anterior Margin (D2A) 8.0

Eye Length (EYL) 3.1 2nd Dorsal Fin Base (D2B) 4.6

Eye Height (EYH) 1.2 2nd Dorsal Fin Height (D2H) 4.0

Interorbital Space (INO) 8.6 2nd Dorsal Fin Inner Margin (D2I) 3.1

Spiracle Length (SPL) 1.5 2nd Dorsal Fin Posterior Margin (D2P) 4.3

Eye-Spiracle Space (ESL) 1.5 Interdorsal Space (IDS) 7.4

Interspiracle Space (ISP) 8.9 Dorsal-Caudal Space (DCS) 6.4

Head Height (HDH) 4.6 Caudal Peduncle Height (CPH) 2.1

Head Width (HDW) 16.6 Caudal Peduncle Width (CPW) 2.1

Trunk Height (TRH) 6.1 Dorsal-Caudal Fin Margin (CDM) 12.3

Trunk Width (TRW) 16.6 Preventral Caudal Fin Margin (CPV) 15.3

Pectoral Fin Length (P1L) 32.2 Lower Postventral Caudal Margin (CPL) 3.7

Pectoral Fin Anterior Margin (P1A) 24.5 Upper Postventral Caudal Margin (CPU) 2.5

Pectoral Fin Base (P1B) 12.3 Subterminal Caudal Fin Margin (CST) 3.4

Pectoral Fin Height (P1H) 14.1

SQUATINA CAILLIETI SP. NOV., A NEW SPECIES OF ANGEL SHARK

the only known capture of a S. caillieti is from off Luzon in the Philippines Islands. However, to the north in the

waters surrounding Taiwan at least four other species are known to occur; S. formosa, S. japonica, S. nebulosa, and

S. tergocellatoides (Walsh & Ebert, 2007). Interestingly, of the four species known to occur around Taiwanese

waters two of them tend to be sympatric in more northerly, cooler waters north of Taiwan. Both S. japonica and S.

nebulosa are known to occur from Taiwan northwards to the main Japanese island of Honshu, including the Sea of

Japan, and as far north as Hokkaido, also they occur along the Chinese mainland into the Yellow Sea, and along the

Korean Peninsula (Compagno et al., 2005a). The poorly known S. tergocellatoides is known primarily from off

Taiwan and Malaysia (Yano et al., 2005). It is likely that this species is wider ranging and may occur in the Philip-

pines. The poorly known S. formosa at the present time has only been reported from Taiwan, but as identification of

WNP squatinids improves its distribution will likely broaden. Finally, a sixth species, the recently described S.

legnota, is known only from off Cilacap, Central Java, Indonesia, Bali, and Lombok in the western South Pacific

(Last & White, 2008) and from discarded specimens from Bali and Lombock, but may be more wide-ranging once

its distribution is better understood.

With the addition of the new species we herein provide a revised and updated key to the WNP squatinids fol-

lows.

FIGURE 3. Distribution of western North Pacific Squatina species. Red star indicates where S. caillieti sp. nov. holotype was

caught in relation to other known ranges of western North Pacific species.

WA LS H ET AL.

FIGURE 4. Comparison of interspiracle distances (ISP) and interorbital distances (INO) of (A) S. caillieti sp. nov. holotype,

(B) S. formosa, and (C) S. nebulosa.

SQUATINA CAILLIETI SP. NOV., A NEW SPECIES OF ANGEL SHARK

FIGURE 5. Comparison of interdorsal distances (IDS) and dorsal-caudal distances (DCS) of (A) S. caillieti sp. nov. holotype,

(B) S. formosa, and (C) S. nebulosa.

WA LS H ET AL.

TABLE 2. Western North Pacific squatinid vertebral and spiral valve counts.

Key to Western North Pacific Squatina Species (revised from Walsh & Ebert, 2007)

1a. Pelvic fin tips do not extend to origin of first dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2

1b. Pelvic fin tips extend to or surpass origin of first dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2a. A prominent row of thorn-like denticles extending from mid-back to caudal peduncle; no distinct ocelli on posterior lobes of

the pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. japonica

2b. No row of thorn-like denticles extending from the mid-back to the caudal peduncle, distinct paired ocelli on the posterior lobes

of the pectoral fins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tergocellatoides

3a. Dorsal-caudal space greater than inter-dorsal space; interorbital space greater than interspiracle space, pelvic inner margin

greater than 9 % total length, pelvic base length greater than 13% total length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4

3b. Dorsal-caudal space less than interdorsal space; interorbital space less than interspiracle space, pelvic inner margin less than

9% total length, pelvic base length less than 13% . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. caillieti sp. nov.

4a. Upper lip arch semi-circular in shape (>1.5% TL in height); dorsals are lobed with a curvilinear anterior margin; pelvic girdle

distance 1.4 times or less head length; caudal fin is lobed, especially dorsally, with a curvilinear postventral caudal margin. . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. formosa

4b. Upper lip arch is not semi-circular in shape (<1.5% TL in height); dorsals are not lobed (angular) without a curvilinear anterior

margin (straight); pelvic girdle distance greater than 1.4 times head length; caudal fin is not lobed (angular), especially dor-

sally, without a curvilinear postventral caudal margin (straight) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. nebulosa

Material examined

Squatina formosa: (4 specimens) Holotype NTT7213130 (now labeled as NTUM 01329), immature female, Tung-

Kang, Pingtung, Taiwan, 31 January 1972; DAE 881805, immature male, Tahsi, Taiwan, May 1988, collected by

David A. Ebert; DAE 052105, immature male, Tahsi, Taiwan, May 2005, collected by David A. Ebert; DAE

052305-2, immature female, Tahsi, Taiwan, May 2005, collected by David A. Ebert.

Squatina japonica: (4 specimens) HUMZ 107395, immature female, off Shirahama, Shimoda City, Shizuoka

prefecture, Japan, 27 January 1986, caught with a bottom gill net at 30 m; HUMZ 105913 immature male, off

Itado, Shimoda, Shizuoka prefecture, Japan, 19 Nov 1985, caught with a bottom gill net at 10m; CAS 20955:

immature male, Japan, Tokyo market, collected by R/V Snyder and Sindo as part of the “Albatross 1906 cruise;”

CAS 20956, immature male, Japan, Tokyo market, collected by R/V Snyder and Sindo as part of the “Albatross

1906 cruise.”

Squatina legnota: (1 specimen) Paratype CSIRO H 6565-01, adult male, 1252 mm TL, Tanjung Luar fish

landing site, Lombok, Indonesia, 08o 45’S, 116o 35’ E, 9 September 2004.

Squatina nebulosa: (8 specimens) Holotype BMNH 1862.11.1.89, immature female, Japan; Paratypes of S.

formosa, three specimens, were examined and reidentified as S. nebulosa by Walsh & Ebert (2007). These speci-

mens are NTU7222433 (now labeled as NTUM 01327), and NTU7041632 (now labeled as NTUM 01328), imma-

ture female, Tahsi, Taiwan (24o 56.5’N, 121o 53.0’E) in single trawling net at 100–120 fathoms, 16 April 1970,

collected by W.H. Ting; NTU7041632 (now labeled as NTUM 01328), immature female, Tahsi, Taiwan, 24 Febru-

ary 1972, collected by W.H. Ting; DAE 882105, immature female, Tahsi, Taiwan, May 1988, collected by David

A. Ebert; DAE 052305-1, immature male, Tahsi, Taiwan, May 2005, collected by David A. Ebert; DAE 052505,

S. caillieti sp. nov. S. formosa S. nebulosa

Vertebral counts n = 1 n = 2 n = 4

Total 137 137–139 138–139

Pre-caudal 111 107–110 108

Monospondylous 51 48–52 49

Diplospondylous 60 58–59 59

Caudal 26 29–30 30–31

n = 1 n = 1 n = 1

Spiral Valve count 7 9 12

SQUATINA CAILLIETI SP. NOV., A NEW SPECIES OF ANGEL SHARK

immature male, May 2005, collected by David A. Ebert; HUMZ 149422, immature female, caught in trawl net in

Okinawa Trough, 25° 37.28’N, 126° 05.35’E to 25° 38.12’N, 126° 07.83’E, 2 August 1994; HUMZ 149423,

immature male, caught in trawl net in Okinawa Trough, 25° 37.28’N, 126° 05.35’E to 25° 38.12’N, 126° 07.83’E,

Japan, 2 August 1994.

Acknowledgements

We thank S.-J. Joung (National Taiwan Ocean University), Peter Last, William White (CSIRO, Hobart, Tasmania,

Australia), George Burgess (Florida Center for Shark Research) and Wade Smith (Oregon State University) for aid

in obtaining comparative WNP squatinid specimens used for this project, Kazuhiro Nakaya (Hokkaido University,

Hokkaido, Japan) for his generous assistance in loaning S. nebulosa specimens, David Catania, Tomio Iwamoto

and Mysi Hoang (California Academy of Sciences) for help with S. japonica specimens and references on squat-

inds, and David Jessup and Eva Berberich (California Department of Fish and Game) for their help with digital

radiographs, Charlie Endris for his expertise in constructing the map depicting WNP squatinid home ranges, and

Enrique Cendejas with his help taking photographs of S. formosa, S. nebulosa, and S. caillieti sp. nov. specimens.

This study was supported by funds from NOAA/NMFS to the National Shark Research Consortium, the Pacific

Shark Research Center, and by The David and Lucile Packard Foundation.

Literature cited

Bleeker, P. (1858) Vierde bijdrage tot de kennis der icthyologische fauna van Japan. Acta Societatis Scientiarum Indo-Neerlan-

dicae, 3(art. 10), 1–46.

Bonaparte, C.L. (1838) Iconografia della fauna italica per le quattro classi degli animali vertebrati. Tomo III. Pesci. Roma.

Iconografia, volume 3: Fasc. 22–23, puntata 104, 110–120, 2 pls.

Chen, J.T.F. (1963) A review of the sharks of Taiwan. Biology Bulletin of Tunghai University Ichthyology Series, No. 1, 1–102.

Compagno, L.J.V. (2001) Sharks of the World: an annotated and illustrated catalogue of shark species known to date. Vol. 2.

Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes and Orectolobiformes). Food and Agriculture

Organization of the United Nations, Rome, 269 pp.

Compagno, L.J.V. (2005) Checklist of living chondrichthyes. In W. Hamlett, ed. Reproductive Biology and Phylogeny of Chon-

drichthyes: Sharks, Batoids, and Chimaeras. Science Publishers, Inc., Enfield, New Hampshire, USA, pp. 503–548.

Compagno, L.J.V., Dando, M. & Fowler, S. (2005a) Sharks of the world. Princeton University Press, Princeton and Oxford,

368pp.

Compagno L.J.V., Last, P.R., Stevens, J.D. & Alava, MNR. (2005b) Checklist of Philippine Chondrichthyes, CSIRO Marine

Laboratories, Report 243, Hobart, Australia, 109 pp.

Ebert, D.A. & Compagno, L.J.V. (2011) Sharks of the World: An annotated and illustrated catalogue of shark species known to

date. Vol. 1. Cow and frilled, dogfish, angel, and saw sharks (Hexanchiformes, Squaliformes, Squatiniformes, and Pristio-

phoriformes). Food and Agriculture Organization of the United Nations, Rome, 411 pp.

Last, P.R. & White, W.T. (2008) Three new angel sharks (Chondrichthyes: Squatinidae) from the Indo-Australian region. Zoot-

axa, 1743,1–26.

Regan, C.T. (1906) Descriptions of some new sharks in the British Museum Collection. Annals and Magazine of Natural His-

tory (Series 7), 18, 435–440.

Shen, S-C. & Ting, W-H. (1972) Ecological and morphological study on fish fauna from the waters around Taiwan and its adja-

cent islands. Notes on some continental shelf fishes and descriptions of two new species. Bulletin of the Institute of Zoo-

logic Academy – Sinica, 11, 13–31.

Walsh, J.H. & Ebert, D.A. (2007) A review of the systematics of western North Pacific angel sharks, genus Squatina, with rede-

scriptions of Squatina formosa, S. japonica, and S. nebulosa (Chondrichthyes: Squatiniformes, Squatinidae). Zootaxa,

1551, 31–47.

Yano, K., Ahmad, A., Gambang, A.C., Idris, A.H., Solahuddin, A.R. & Aznan, Z. (2005) Sharks and rays of Malaysia and Bru-

nei Darussalam. SEAFDEC-MFRDMD/SP/12. Kuala Terengganu, 557 pp.

First Record of African Angel Shark, Squatina africana (Chondricthyes: Squatinidae) in Indian Waters, Confirmed by DNA Barcoding

Article

Full-text available

May 2018

A single specimen of African angel shark, Squatina africana (Regan, 1908) was caught off Lakshadweep (11°5′47″ N; 72°2′21″ E), India in September 2016. The present study is a new report of the above species from Indian waters. In addition to classical methodologies, DNA barcoding was also adopted for species identification. The 650 bp-long region of mitochondrial Cytochrome Oxidase subunit I was sequenced to obtain the DNA barcode for the species under study. The sequence divergence value within species and between species was calculated using MEGA V.7.0, where Kimura 2 parameter (k2p) model was chosen as a distance model. The average k2p distance separating individuals within species was 1.76% and inter specific divergence was 8–10%. A neighbour joining network was constructed to provide a graphical representation of divergence between the species. Using the maximum identity with Gen Bank database, K2P divergence distance, NJ-network and traditional morphological approach, we could identify the given specimen as a mature male African angel shark.

Angel sharks (Squatinidae): A review of biological knowledge and exploitation

Article

Nov 2020

Angel sharks (Squatina spp.) are distributed in warm temperate to tropical waters around the world. Many species occur in shelf seas and exhibit seasonal inshore‐offshore migrations, moving inshore to give birth. Consequently, there can be high spatial overlap between angel shark populations with fisheries and other human activities. Their dorso‐ventrally flattened body shape, large size (most species attain >100 cm total length, LT) and demersal nature means that they may be taken in a variety of demersal fishing gears from birth. Available data indicate that angel sharks typically have a biennial reproductive cycle, with litter sizes generally <20 and the young born at ca. 20–30 cm. The biological characteristics of angel sharks render them susceptible to overexploitation, as exemplified by the decline of Squatina squatina from many parts of its former range in the North‐east Atlantic and Mediterranean Sea. Currently, half of the 22 recognised extant species of angel shark are classed as Threatened on the International Union for Conservation of Nature (IUCN) Red List (with a further three classified as Data Deficient). Given the biological vulnerability of angel sharks, and that many species are data‐limited, the current paper provides a review of available biological information and fisheries data pertaining to this family. This article is protected by copyright. All rights reserved.

Evolutionary trends of the conserved neurocranium shape in angel sharks (Squatiniformes, Elasmobranchii)

Article

Full-text available

Jul 2020

elasmobranchii (i.e., sharks, skates, and rays) forms one of the most diverse groups of marine predators. With a fossil record extending back into the Devonian, several modifications in their body plan illustrate their body shape diversity through time. the angel sharks, whose fossil record dates back to the Late Jurassic, some 160 Ma, have a dorsoventrally flattened body, similar to skates and rays. fossil skeletons of this group show that the overall morphology was well established earlier in its history. By examining the skull shape of well-preserved fossil material compared to extant angel sharks using geometric morphometric methods, within a phylogenetic framework, we were able to determine the conservative skull shape among angel sharks with a high degree of integration. the morphospace occupation of extant angel sharks is rather restricted, with extensive overlap. Most of the differences in skull shape are related to their geographic distribution patterns. We found higher levels of disparity in extinct forms, but lower ones in extant species. Since angel sharks display a highly specialized prey capture behaviour, we suggest that the morphological integration and biogeographic processes are the main drivers of their diversity, which might limit their capacity to display higher disparities since their origin.

New Species of Squatina (Squatiniformes: Squatinidae) from Brazil, with Comments on the Taxonomy of Angel Sharks from the Central and Northwestern Atlantic

Article

Full-text available

Mar 2018
COPEIA

Morphological analysis of South- and Northwestern Atlantic specimens of Squatina revealed an undescribed species of angel shark occurring on the continental slope of Brazil between latitudes 11 and 22S. The new species of Squatina is distinguished from western Atlantic congeners by a unique combination of dorsal color pattern, vertebral counts, morphology of lateral dermal folds, dermal denticles, female reproductive tract, clasper, and pectoral fin characters. A preliminary assessment of the taxonomy of angel shark species from the Central and Northwestern Atlantic is also provided.

Bibliography database of living/fossil sharks, rays and chimaeras (Chondrichthyes: Elasmobranchii, Holocephali) List of Valid Extant Species List of Described Extant Species Statistic

Book

Full-text available

Mar 2021

The table and provided download links below are intended for informational use in Chondrichthyan research. The allocation aims for faciliating to find species numbers and most recent information on taxonomic changes. We will regularly update the table and download links at lest twice annually. The updates will be announced on facebook (https://www.facebook.com/sharkreferences) and in our monthly newsletter (sign up here: https://eepurl.com/sJNGb). The Excel sheet allows for the application of individual filter- and sorting options. The list of described spsecies complements taxonomic information for the list of valid species by providing synonyms and / or new taxonomic combinations.

Species Delimitation of Southeast Pacific Angel Sharks (Squatina spp.) Reveals Hidden Diversity through DNA Barcoding

Article

Full-text available

Apr 2021
Diversity

Angel sharks are distributed worldwide in tropical to subtropical waters. Across the Eastern Pacific Ocean (EPO), two valid species are reported: The Pacific angelshark Squatina californica and the Chilean angelshark Squatina armata; however, there is still uncertainty about their geographic distribution, mainly along the northern Peru coast where the species have been reported to be sympatric. The aim of this study is to describe the genetic differences between the genus Squatina from the EPO, including samples from northern Peru, and using DNA barcoding and three species delimitation models: Poisson tree processes (PTP) model, Bayesian implementation of the PTP (bPTP) model and the general mixed Yule coalescent (GMYC) model. The three approaches summarized 19 nominal Squatina species in 23 consensus Molecular Taxonomic Units (MOTU). Only 16 of them were in accordance with taxonomic identifications. From the EPO, four Squatina MOTUs were identified, one from North America (S. californica USA/Mexico) and three sampled in northern Peru, S. californica Peru, S. armata and Squatina sp. (a potential new species). This study contributes to the management and conservation policies of angel sharks in Peru, suggesting the presence of an undescribed species inhabiting the northern Peruvian coast. The use of molecular approaches, such as DNA barcoding, has the potential to quickly flag undescribed species in poorly studied regions, including the Southeast Pacific, within groups of ecologically and economically important groups like angel sharks.

Squatina japonica. THE IUCN RED LIST OF THREATENED SPECIES™

Technical Report

Full-text available

Mar 2021

Squatina nebulosa. THE IUCN RED LIST OF THREATENED SPECIES™

Technical Report

Full-text available

Mar 2021

An identification key to elasmobranch genera based on dental morphological characters Part A: Squalomorph sharks (Superorder Squalomorphii).

Article

Full-text available

Mar 2019

In addition to articulated, mostly formaldehyde-fixed and ethanol-preserved, taxidermy or anatomical specimens, sharks and rays are represented in scientific collections by numerous jaws and isolated teeth. These specimens often source from historical collections where existing informations about species, sex or geographic origin in many cases are scarce, incomplete or incorrect. The identification key for jaws and teeth presented herein focuses on squalomorph sharks, which comprise almost 34 % of all sharks with 179 species in 31 genera and 11 families. The key is essentially based on the following characters: vascularisation stage, labial apron, number of cusplets, distal heel, lingual ornamentation, cutting edge, and dentition kind. The key allows the identification to genus level. It is further supplemented by a comprehensive glossary of tooth morphological terms as well as an updated checklist of all currently described squalomorph sharks with indication of the distribution and the dental formula. Zusammenfassung: In zahlreichen wissenschaftlichen Sammlungen befinden sich neben formaldehyd-oder alkoholkonservierten Ganzkörper-und anatomischen Präparaten oftmals Kiefer und Zähne von Haien und Rochen. Diese Exemplare stammen häufig aus historischen Sammlungen und die vorhandenen Informati-onen über die Art, das Geschlecht oder die geografische Herkunft sind oftmals lückenhaft oder falsch. Mit der vorliegenden Arbeit wird ein Bestimmungsschlüssel für Kiefer und Zähne von squalomorphen Haien vorgestellt. Diese Überordnung beinhaltet derzeit 179 Arten in 31 Gattungen und 11 Familien und stellt damit annähernd fast 34 % aller heute bekannten Haiarten dar. Zur Identifizierung wird dabei im Wesentlichen auf folgende Merkmale zurückgegriffen: Art des Vaskularisationssystems, labiales Apron, Anzahl der Zahn-spitzen, Talon, linguale Ornamentik, Form der Schneide und Art der Bezahnung. Der Schlüssel ermöglicht die Bestimmung bis auf Gattungsebene. Ergänzt wird er durch ein umfangreiches Glossar der wichtigsten zahnmorphologischen Begriffe sowie eine aktualisierte Checkliste aller derzeit bekannten squalomorphen Haie mit Angabe der geographischen Verbreitung und der Zahnformel.

Pating Ka Ba? An Identification Guide to Sharks, Batoids and Chimaeras of the Philippines

Book

Full-text available

Jan 2014

FOREWORD Sharks and their relatives, the rays, skates and chimaeras, were previously considered to be of low economic value to large-scale commercial fisheries. In recent years, sharks have gained commercial importance to large-scale commercial and artisanal fisheries. They are increasingly targeted for their meat, skin, cartilage, teeth, jaws, liver/liver oil, and even other internal organs. Demand for sharks and sharks products in the past 30-40 years has resulted in an increasing number of species threatened with extinction due to a combination of factors from over-exploitation, degradation of important habitats (i.e., nursery and breeding grounds) from development and pollution compounded by the effects of a changing climate. Consequently, there is now increasing concern about the rise of shark catches and the impact this has for some shark species populations. There is a need to balance fishing efforts and resource exploitation with conservation and management measures to attain sustainability of our marine fishery resources. As reported in the Philippine National Plan of Action for the Conservation and Management of Sharks (NPOA-Sharks), most if not all of possibly 160 or more cartilaginous fish species reported to occur in our waters factor either in directed/targeted catch or incidental by-catch of commercial and/or municipal fisheries. Yet, there is limited reporting record on each of these species nationwide, particularly in terms of species-specific catch and effort, landing volume and trade data. There is also a serious lack of information on the biological parameters of shark species, on the identification of most species, and on the capacity to manage the resource as a whole. Such limitation on our knowledge of sharks and of the practices employed in shark fisheries in many of our fishing grounds is hindering us in providing appropriate management measures on the ground. To improve knowledge on the state of shark stocks and facilitate the collection of the necessary information, adequate resources and support system are required for research and management. Sharks and its relatives are characterized by K-selected life history traits such as slow growth, late sexual maturity, low fecundity, low natural mortality, and long life resulting in low rates of population increase. Due to these character traits and many other complex spatial structures such as size/sex segregation and seasonal migration, sharks often have a low stock-to-recruitment relationship and long stock recovery times when overfished. Thus, these fishes are highly vulnerable to overexploitation. This book “Pating Ka Ba? An Identification Guide to Sharks, Batoids and Chimaeras of the Philippines” is long overdue. There is a need to improve the management of directed fisheries of sharks and its relatives, as well as certain multispecies fisheries in which they constitute as by-catch. This identification guide is a useful reference to enhance our understanding on these species and our capacity to manage them. Atty. Asis G. Perez Director Bureau of Fisheries and Aquatic Resources Foreword

A Review Of The Systematics Of Western North Pacific Angel Sharks, Genus Squatina, With Redescriptions Of Squatina Formosa, S. Japonica, And S. Nebulosa (Chondrichthyes: Squatiniformes, Squatinidae)

Article

Full-text available

Aug 2007

Squatinids are quite distinct from other shark-like fishes, but individual species are difficult to differentiate. Four of the 16 known, valid squatinid species occur in the western North Pacific (WNP). Differences among the WNP species complex have traditionally relied upon the nasal barbel shape, interorbital and interspiracle distances, ocellus patterns, number of dermal folds about the mouth, and the presence of midback thorns. Unfortunately, many of these characters are difficult to distinguish, hindering identification of individuals. Using WNP squatinid specimens and photographs, both from field expeditions and museums, we confirm the validity of four species in the area. Additionally, we correct mistakes made in the literature on S. formosa type material, clarify differences in the particularly challenging distinction between S. formosa and S. nebulosa, and provide the basis for a revised dichotomous key for the region that includes all four known valid WNP squatinid species.

Bijdrage tot de kennis der ichthyologische fauna van Ternate

Article

Jan 1853

P Bleeker

Three New Angel Sharks (Chondrichthyes: Squatinidae) From The Indo-Australian Region

Article

Mar 2008

Four species of angel sharks (family Squatinidae) occur in temperate and subtropical Australian waters. Two of these, Squatina albipunctata sp. nov. and S. pseudocellata sp. nov., which occur mainly off subtropical eastern and western Australia respectively, are formally described and illustrated. The new species differ from temperate Australian species, S. australis and S. tergocellata, in morphometrics, meristics, squamation, and coloration. Another new angel shark, S. legnota sp. nov. from eastern Indonesia, is compared to these species. Unlike Australian Squatina, it has unfringed (rather than fringed) barbels on its nasal flap.

FAO Species Catalogue. Vol. 4, Part 2, Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date

Article

Nov 1983

Job number added acc. to DOCREP assignment

Iconografia della fauna italica per le quattro classi degli animali vertebrati

Jan 1838
110-120

C L Bonaparte

Bonaparte, C.L. (1838) Iconografia della fauna italica per le quattro classi degli animali vertebrati. Tomo III. Pesci. Roma. Iconografia, volume 3: Fasc. 22-23, puntata 104, 110-120, 2 pls.

A review of the sharks of Taiwan

Jan 1963
1-102

J T F Chen

Chen, J.T.F. (1963) A review of the sharks of Taiwan. Biology Bulletin of Tunghai University Ichthyology Series, No. 1, 1-102.

Checklist of Philippine Chondrichthyes

Jan 2005
pp

L J V Compagno
P R Last
J D Stevens
Mnr Alava

Compagno L.J.V., Last, P.R., Stevens, J.D. & Alava, MNR. (2005b) Checklist of Philippine Chondrichthyes, CSIRO Marine Laboratories, Report 243, Hobart, Australia, 109 pp.

Checklist of living chondrichthyes

Jan 2005
503-548

L J V Compagno

Compagno, L.J.V. (2005) Checklist of living chondrichthyes. In W. Hamlett, ed. Reproductive Biology and Phylogeny of Chondrichthyes: Sharks, Batoids, and Chimaeras. Science Publishers, Inc., Enfield, New Hampshire, USA, pp. 503-548.

Descriptions of some new sharks in the British Museum Collection

Jan 1906
435-440

C T Regan

Regan, C.T. (1906) Descriptions of some new sharks in the British Museum Collection. Annals and Magazine of Natural History (Series 7), 18, 435-440.

Sharks and rays of Malaysia and Brunei Darussalam

Jan 2005
pp

K Yano
A Ahmad
A C Gambang
A H Idris
A R Solahuddin
Z Aznan

Yano, K., Ahmad, A., Gambang, A.C., Idris, A.H., Solahuddin, A.R. & Aznan, Z. (2005) Sharks and rays of Malaysia and Brunei Darussalam. SEAFDEC-MFRDMD/SP/12. Kuala Terengganu, 557 pp.