Cartilaginous fish diversity in the Western Visayas, Philippines, including two putative unidentified species and the first record of Carcharhinus plumbeus (Elasmobranchii: Carcharhiniformes: Carcharhinidae)

Roxanne Cabebe-Barnuevo; Dianne Frances Penuela; Emmanuel S. Delloro Jr.; Ricardo P. Babaran; Hiroyuki Motomura; Maria Celia D. Malay

doi:10.3897/aiep.55.139721

Abstract

This annotated checklist documents the diversity of cartilaginous fishes, including sharks, batoids (rays and skates), and chimaeras, from Panay and Guimaras islands, Western Visayas, Philippines, through surveys at fish markets and ports. A total of fourteen species were sampled and identified, including one chimaera, six sharks, and seven batoids from ten families (Chimaeridae, Hemiscylliidae, Scyliorhinidae, Carcharhinidae, Pristiophoridae, Rhinobatidae, Plesiobatidae, Urolophidae, Dasyatidae, and Aetobatidae). Among these, the presence of the sandbar shark, Carcharhinus plumbeus (Nardo, 1827) in Philippine waters is confirmed for the first time, whereas a potential unidentified batoid species within the genera Himantura Müller et Henle, 1837 and Urolophus Müller et Henle, 1837 are reported. The twelve nominal taxa described herein include three species classified as Least Concern, five are Near Threatened, two Vulnerable, one Endangered, and one is Critically Endangered as per the IUCN Red List assessments. These documented species supplement recent inventories, expanding our knowledge on the diversity and distribution of cartilaginous fishes in the region and country, while also providing comprehensive morphological descriptions, validated molecular data, and curated voucher specimens for future reference.

Keywords

checklist, Chondrichthyes, COI gene, morphology, new record, taxonomy

Introduction

Members of the class Chondrichthyes, commonly known as the cartilaginous fishes, are divided into two subclasses: Holocephali (chimaeras) and Elasmobranchii (including rays, skates, and sharks). In southeast Asia, a total of 393 cartilaginous species have been reported, consisting of seven chimaeras, 196 sharks, and 190 batoids (SEAFDEC 2022). Compagno et al. (2005) reported 164 cartilaginous fish species from the Philippines, comprising three chimaeras, 95 sharks, and 66 batoids. Of these, the field guide of Alava et al. (2014) provided identification resources for 91 species. In a more recent review conducted by Santos et al. (2017), previous reports were updated, resulting in a nominal list of about 206 cartilaginous species from Philippine waters. Recent taxonomic additions from the Philippines include Squatina caillieti Walsh, Ebert et Compagno, 2011 (from Luzon), Pristiophorus lanae Ebert et Wilms, 2013 (Marinduque), Rhinobatos whitei Last, Corrigan et Naylor, 2014 (Dipolog), and Okamejei panayensis Misawa, Babaran et Motomura, 2022 (Panay Island, where it derives its name). The life history traits of cartilaginous fishes render them highly vulnerable to overexploitation and population collapse, and many species are of particular ecological importance and conservation concern (Stevens et al. 2000; Simpfendorfer et al. 2002; Baum et al. 2003; Field et al. 2009; Ferretti et al. 2010; Dulvy et al. 2014; Cortés et al. 2015). Thus, there is increasing interest in documenting the diversity, distribution, and population status of cartilaginous fish species to facilitate science-based conservation and management decision-making.

Seventeen species of cartilaginous fishes have previously been reported from the Western Visayas (Koeda and Manjaji-Matsumoto 2017; Manjaji-Matsumoto 2017), a region of the central Philippines with highly intensive coastal fisheries. As part of a larger project to re-survey the diversity of marine fishes in the Western Visayas, this study is focused on collecting and DNA-barcoding vouchered museum specimens of cartilaginous fishes from the region. The research aims to provide a comprehensive inventory of chimaeras, sharks, rays, and skates, while utilizing DNA barcodes to confirm species identifications and enhance the accuracy of taxonomic records. By providing validated molecular data and ensuring accessibility of voucher specimens, this research contributes to the knowledge of the diversity and distribution of these species in the Philippines, providing a valuable resource for future research and conservation efforts.

Materials and methods

Samples were purchased directly from local fish markets, landing sites, and fish ports by the project researchers, in collaboration with trained local enumerators, in the provinces of Aklan, Antique, Capiz, Iloilo, and Guimaras (Fig. 1) from August 2020 to February 2022. Whenever possible, the fishing gear used in catching the specimens was recorded. Upon collection, each specimen was wrapped in moist tissue paper and frozen. Frozen specimens were then transported to the University of the Philippines Visayas (UPV), Miagao, Iloilo. Curatorial protocols by Motomura and Ishikawa (2013) were followed. Measurements of the specimens were taken using digital calipers, following Serena (2005) and expressed as a percentage of disc width (DW) for the majority of batoids and of total length (TL) for sharks and chimaeras. All measurements were recorded to the nearest 1 mm and are presented in Tables 1, 2. All comparable metrics included in the description were given as a percentage of TL or DW. All specimens were deposited at the University of the Philippines Visayas-Museum of Natural Sciences, Miagao, Iloilo, Philippines (UPVMI).

Table 1.

Download as

CSV

XLSX

Morphological measurements for the six shark species collected from Western Visayas, Philippines used in the study. All measurements are expressed as a percentage of total length (TL).

Species	Chiloscyllium plagiosum	Chiloscyllium punctatum	Chiloscyllium punctatum	Chiloscyllium punctatum	Atelomycterus marmoratus	Carcharhinus plumbeus	Carcharhinus plumbeus	Carcharhinus plumbeus	Carcharhinus sorrah	Pristiophorus lanae
Institutional code	UPVMI 3123	UPVMI 3124	UPVMI 3125	UPVMI 3122	UPVMI 3129	UPVMI 3120	UPVMI 3183	UPVMI 3119	UPVMI 3121	UPVMI 3126
Total length	441	302	536	553	479	363	370	391	582	700
In %TL
Head length	15	17	17	16	17	24	23	23	20	36
Trunk length	18	16	19	19	25	26	25	28	28	21
Tail length	67	67	64	65	58	51	52	49	52	43
Precaudal-tail length	46	42	40	47	35	22	22	21	18	22
Interdorsal space	9	13	11	14	13	19	19	20	23	14
Preoral length	5	4	3	4	4	8	8	8	8	28
Mouth width	6	5	5	7	7	9	9	9	7	4
Eye diameter	2	2	2	2	3	3	3	3	2	3
Interorbital space	6	5	5	5	5	11	11	11	9	4
Internasal distance	4	4	3	4	5	6	8	7	6	3
1^st dorsal-fin base	9	8	8	13	8	11	13	13	10	5
2^nd dorsal-fin base	9	9	10	12	9	5	5	5	3	5
1^st dorsal-fin height	14	12	12	16	13	16	16	16	13	10
2^nd dorsal-fin height	13	11	11	15	12	5	5	6	3	10
Pectoral-fin base	5	5	5	6	5	6	6	6	6	3
Pectoral-fin height	13	13	13	15	12	16	16	16	16	12
Pelvic-fin base	7	6	7	7	6	5	5	5	4	4
Pelvic-fin height	11	10	10	10	9	6	6	6	5	6
Anal-fin base	9	9	10	9	8	5	5	5	5	-

Table 2.

Download as

CSV

XLSX

Morphological measurements for the six batoid species collected from Western Visayas, Philippines used in the study. All measurements are expressed as a percentage of disc width (DW), except for Rhinobatos whitei, which are expressed as a percentage of total length (TL).

Species	Rhinobatos whitei	Rhinobatos whitei	Plesiobatis daviesi	Himantura sp.	Neotrygon orientalis	Taeniura lymma	Aetobatus ocellatus
Institutional codes	UPVMI 3128	UPVMI 3127	UPVMI 3112	UPVMI 3111	UPVMI 3113	UPVMI 3114	UPVMI 3115
Total length	135	694	-	-	506	261	1556
Disc Width	32	34	368	377	256	104	574
Tail length	57	56	-	-	127	149	219
Disc length	41	40	99	92	71	105	44
Interorbital space	5	6	14	19	15	15	14
Snout length (preorbital)	14	16	37	21	16	26	7
Snout length (preoral)	17	18	34	21	14	17	7
Internasal space	4	6	15	8	7	9	4
Pelvic-fin length	4	7	14	20	16	23	15
Mouth width	7	6	13	9	7	12	7

Figure 1.

Map of the Western Visayas region, Philippines, highlighting the provinces of Aklan, Antique, Capiz, Iloilo (Panay Island), and Guimaras (Guimaras Island) where the samples for this study were collected.

Muscle tissue samples were collected from the ventral area of the fish and preserved in absolute ethanol. Extraction of DNA was carried out following the instructions provided in the GF-1 Nucleic Acid Extraction Kit (Vivantis Technologies Sdn. Bhd, Malaysia) and PureLinkTM Genomic DNA Mini Kit (Invitrogen). Polymerase chain reaction (PCR) amplification was performed using the forward and reverse primers FishF1, FishR1, VF2_t1, FishF2_tl, FishR2_t1, and FR1d_t1, designed to amplify the mitochondrial cytochrome c oxidase subunit I (COI) gene (Ward et al. 2005; Ivanova et al. 2007).

Molecular laboratory work was performed at the Philippine Genome Center – Visayas Satellite Facility or the Marine Biodiversity and Evolution Lab at UPV. The PCR mixture (25 μL) was composed of nuclease-free water (18.4 μL), 10x PCR buffer (2.25 μL), MgCl₂ (1.25 μL), 10 mM dNTP mix (0.5 μL), Taq DNA polymerase (0.1 μL), forward and reverse primers (0.25 μL each), and DNA template (2 μL). The thermocycler conditions were as follows: 2 min at 95°C (initial step); 35 cycles of 0.5 min at 94°C (denaturation); 0.5 min at 54°C (annealing), and 1 min at 72°C (extension); and 10 min at 72°C (final extension). The PCR products were visualized using 1% agarose gel and subsequently purified using the GF-1 PCR Clean-up Kit and PureLinkTM PCR Purification Kit. The amplified DNA was quantified using a Genova nano spectrophotometer (Jenway, USA) or MultiSkan Skyhigh Microplate Spectrophotometer (Thermo Fisher Scientific). PCR products were sent to Macrogen Inc. (South Korea) for sequencing. The forward and reverse sequences were examined, trimmed, and realigned using Unipro UGENE software (Okonechnikov et al. 2012). The generated sequences were deposited in the Genetic Sequence Bank (GenBank) database of the National Center for Biotechnology Information (NCBI), with accession numbers OR614465 to OR614481.

Phylogenetic trees were constructed using the maximum likelihood (ML) method, based on Kimura 2-parameter (K2P) model (Kimura 1980), with 1000 bootstrap replications. All analyses were performed using MEGA X software (Kumar et al. 2018). When constructing the phylogenetic tree, priority was given to sequences from vouchered specimens available in GenBank and BOLD, sequences from species which are closely related to the collected specimens as mentioned in the remarks, and sequences referenced in other published studies. Sequences used in the analysis are provided in Suppl. material 1.

Results

This study documents 14 cartilaginous fish species, including one chimaera, six sharks, and seven batoids. Each species profile includes a morphological description, distribution details, GenBank accession number, standard name, and conservation status based on the IUCN (2023) Red List, and additional relevant information (e.g., comparisons with closely related species). Species are organized by family following the systematic arrangement outlined by Nelson et. al (2016), with genera and species names listed in alphabetical order.

Family Chimaeridae Rafinesque, 1815

Genus Chimaera Linnaeus, 1758

Chimaera phantasma Jordan et Snyder, 1900

IUCN standard name:: Silver Chimaera Fig. 2, Table 1

Material examined.

UPVMI 3118, 889 mm TL (tail broken), female, fish market, Culasi, Antique Province, Philippines, 27 July 2020, coll. E.A. Obar.

Figure 2.

Chimaera phantasma from Western Visayas, Philippines, UPVMI 3118, 889 mm TL (tail broken) A body in lateral view B head in lateral view C oronasal region.

Morphological description.

Body large, compressed, elongated, and tapering towards tail; lateral-line canals prominent and wavy. Head large; lateral-line canals straight rather than wavy. Eyes large, horizontally circular; spiracles absent. Snout short, rounded and very soft; tips gelatinous. Mouth small, connected to nostrils. Nostrils large and rounded. Tooth plates on upper jaw two (obviously separated), broad; tooth plate on lower jaw single, pointed, and curved. First dorsal fin tall with a single, erectile spine; first dorsal spine slender, smooth in front, sharply serrated behind; second dorsal fin short and spineless; dorsal-fin base long, extended to tail area. Anal fin short, deeply notched; anal-fin base small. Pectoral fin large, leaf-shaped; apex bluntly rounded. Pelvic fin small, broadly ovoid. Caudal fin elongated, tapering, lanceolate with distinct upper and lower lobes. Most of posterior portion of tail cut off, but remaining parts of tail narrow, rounded, with smooth edges, and whip-like.

Color of fresh specimen. Body surface silver, underside white; snout black; fin edges black.

Color of preserved specimen. Body surface and fins brown; underside dusky white to gray; snout surface area darker brown, underside white; dorsal-, anal-, and caudal-fin edges black.

Distribution.

western Pacific. Japan, Philippines, Borneo, and Taiwan (Last et al. 2010; Ebert et al. 2013).

DNA barcode.

The COI sequence of 650 base pairs (bp) was submitted to GenBank under accession number OR614464.

Conservation status.

Vulnerable.

Remarks.

Chimaera phantasma can reach a maximum total length of 1 m and typically inhabits depths ranging from 90–540 m (Compagno 1999). There have been reports of occurrences in Philippine waters, including Batangas, Cebu, Negros Oriental (Dumaguete), Palawan, and Zamboanga Del Norte (Dapitan and Dipolog cities) (Compagno et al. 2005; Aquino et al. 2023).

The specimen collected in this study was identified as C. phantasma based on its COI data (see Fig. 3) and morphological characteristics, including a silver, tapering body with a blunt, fleshy snout; a wavy lateral line; a long first dorsal spine extending beyond the fin apex; an anal fin separated from the caudal fin by a shallow notch; and grayish to blackish fins. These features match the description of C. phantasma provided by Jordan and Snyder (1900), Last et al. (2008), and Finucci et al. (2018). Molecular data analysis suggests that Chimaera argiloba Last, White et Pogonoski, 2008, Chimaera ogilbyi Waite, 1898, and C. phantasma are closely related species (Finucci et al. 2018). This relationship is also observed in the phylogenetic tree generated in this study (Fig. 3). However, according to Last et al. (2008), C. phantasma can be distinguished from C. argiloba by its uniformly gray to black first dorsal fin (vs. pale gray with a distinct white posterior margin in C. argiloba), and black pectoral fins (vs. gray to brown). Furthermore, C. phantasma can be easily distinguished from C. ogilbyi by the presence of an anal fin (vs. absent in C. ogilbyi) (Finucci et al. 2018).

Figure 3.

Maximum likelihood phylogenetic tree of partial COI sequences from chimaeras and sharks, constructed using the Kimura 2-parameter model. Philippine sequences are identified by UPVMI voucher codes, with related genera color-coded. Bootstrap support values (1000 replicates) are shown at nodes, and the scale bar represents evolutionary distance in substitutions per site.

Family Hemiscylliidae Gill, 1862

Genus Chiloscyllium Müller et Henle, 1837

Chiloscyllium plagiosum (Bennett, 1830)

IUCN standard name:: Whitespotted Bambooshark Fig. 4, Table 1

Material examined.

UPVMI 3123, 441 mm TL, male, fish landing site, Cabalagnan, Nueva Valencia, Guimaras Province, Philippines, October 2021, coll. J. Tubillara.

Morphological description.

Body small, elongated, moderately slender, with lateral dermal ridges. Eyes relatively small without nictitating eyelids; spiracles large and located below eyes. Snout rounded and broad. Mouth small, positioned closer to eyes than to snout tip. Nostrils small, partially covered by broad nasal flap; nasal flap reaching mouth; sensory barbels on each nostril very short. Teeth on both jaws small with single small cusp. Both dorsal fins fully erect, equal in size; posterior margin straight; first dorsal fin originating behind pelvic-fin origin. Anal fin short, originating behind free rear tip of second dorsal fin; positioned just anterior to caudal-fin origin; anal-fin base length shorter than base of lower caudal-fin lobe (9% vs. 16% TL, respectively). Pectoral fin slightly larger than dorsal fins, originating just below third gill opening; posterior margin straight. Pelvic fin approximately similar in size to dorsal fins, located just behind pectoral fin and front of first dorsal fin insertion. Caudal fin asymmetrical; subterminal notch well developed; ventral lobe absent. Precaudal pits absent. Gill slits on both sides five, small; fourth and fifth gill slits situated extremely close to one another.

Color of fresh specimen. Body and head surface area light brown with darker transverse bands and covered of numerous white and black spots; underside light brown with numerous white spots and few black spots. Fins brown with numerous white spots.

Color of preserved specimen. Transverse dark brown bands, white and black spots still visible on the body, head, and fins.

Distribution.

Indo–Pacific. western Indian Ocean: Madagascar, India, and Sri Lanka; western Pacific: Indonesia, Philippines, Japan, and New Guinea (Human 2022).

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank under accession number OR614480.

Conservation status.

Near Threatened

Remarks.

Chiloscyllium plagiosum has a maximum recorded length of 950 mm (Compagno and Niem 1998a). This species can be found at depths ranging from 0–50 m (Weigmann 2016). Sightings have been reported in Philippine waters, particularly in Cavite, Cebu, Guimaras Island, Manila Bay, Negros Occidental, Negros Oriental, Oriental Mindoro (Calapan), and Palawan (Compagno et al. 2005; Gonzales 2013).

The specimen was identified as C. plagiosum based on its slender body and tail, anal-fin base shorter than the base length of lower caudal-fin lobe (9% vs. 16% TL, respectively), presence of lateral dermal ridges, transverse dark bands, and small white spots. According to Compagno and Niem (1998a), C. plagiosum is frequently misidentified as Chiloscyllium griseum Müller et Henle, 1838, Chiloscyllium hasselti Bleeker, 1852, Chiloscyllium indicum (Gmelin, 1789), or Chiloscyllium punctatum Müller et Henle, 1838. These species all exhibit transverse dark bands on their bodies, especially when young, which often leads to misidentifications. However, C. plagiosum can be distinguished from C. indicum by its shorter anal-fin base compared to the base length of lower caudal-fin lobe (vs. base lengths of anal-fin and lower caudal fin lobe equal in C. indicum) (Compagno and Niem 1998a). Additionally, C. plagiosum is differentiated from C. griseum, C. hasselti, and C. punctatum by the presence of lateral dermal ridges (Fig. 4B), which are absent in the latter three species (Compagno and Niem 1998a). The identity of the collected specimen was further verified using COI data, which showed it forming a distinct clade with a sequence of C. plagiosum, separate from the clades of the aforementioned closely related species (Fig. 3).

Figure 4.

Chiloscyllium plagiosum from Western Visayas, Philippines, UPVMI 3123, 441 mm TL. A body in lateral view B head in dorsal view C oronasal region.

Chiloscyllium punctatum Müller et Henle, 1838

IUCN standard name:: Grey Carpetshark Fig. 5, Table 1

Material examined.

UPVMI 3124, 302 mm TL, male, fish landing site, Roxas City, Capiz Province, Philippines, trawl, November 2021, coll. M. Batay. UPVMI 3125, 536 mm TL female, same data as preceding. UPVMI 3122, 553 mm TL, male, fish market, Tangalan, Aklan Province, Philippines, 23 November 2020, coll. M. Macavinta.

Figure 5.

Chiloscyllium punctatum from Western Visayas, Philippines, UPVMI 3124, 302 mm TL (A body in lateral view B head in dorsal view C oronasal region); UPVMI 3125, 536 mm TL (D body in lateral view E head in dorsal view F oronasal region); UPVMI 3122, 553 mm TL (G body in lateral view H head in dorsal view I oronasal region).

Morphological description.

(based on UPVMI 3124): Body small and slender without lateral dermal ridges. Eyes relatively small and oval, without nictitating eyelids; spiracles moderately small, located below eyes. Snout rounded and broad. Mouth small, located closer to eyes than snout tip. Nostrils small and subterminal, partially covered by nasal flap; nasal flap broad just reaching mouth; sensory barbels on each nostril very short. Teeth on both jaws small with single small cusp. Dorsal fins fully erect, equal in size; first dorsal fin originating behind pelvic-fin origin; posterior margin concave. Anal fin moderately long, originating behind free rear tip of second dorsal fin; anal-fin base length shorter than base of lower caudal-fin lobe (9% vs. 16% of TL, respectively). Pectoral fin angular with rounded free rear tips, originating just below third gill opening. Pelvic fin slightly smaller than dorsal fins, located just behind pectoral fin. Caudal fin asymmetrical; subterminal notch well developed; ventral lobe absent; precaudal pits absent. Gill slits on both sides five, small; fourth and fifth gill slits situated extremely closed to one another, making fifth gill slit hardly noticeable.

Color of fresh specimens . UPVMI 3124 (smallest specimen): Body and head white with brown to black transverse bands, underside white; bands extending to anal, dorsal, and pectoral fins; pelvic fin white. UPVMI 3125 (medium sized): Body, head and fins brown with faint brown bands. UPVMI 3122 (largest specimen): Body, head, and fins uniformly brown, colored bands absent.

Color of preserved specimen. UPVMI 3124 and UPVMI 3125: Transverse dark brown bands still visible on the body, head, and fins; underside white.

Distribution.

Indo–Pacific. western Indian Ocean: Arabian Sea and southern India; western Pacific: Thailand, Indonesia, Philippines, Japan, New Guinea, and Australia (Human 2022).

DNA barcode.

COI sequences of 617 bp were submitted to GenBank with the following accession numbers: OR614477 (UPVMI 3124), OR614474 (UPVMI 3125), and OR614472 (UPVMI 3122).

Conservation status.

Near Threatened

Remarks.

Chiloscyllium punctatum can reach a maximum total length of 1050 mm and has been found in the intertidal zone at depths exceeding 85 m (Compagno 2001). There have been reports of occurrences in Philippine waters, specifically in Cebu (Cebu City, and Lapulapu), Kalinga (Pasil), Mindoro Island, Negros Oriental (Dumaguete), Palawan, and Zamboanga (Compagno et al. 2005; Gonzales 2013; Aquino et al. 2023).

The specimen was identified as C. punctatum based on several combined characteristics, including a slender body and tail, absence of lateral dermal ridges on the body, a concave posterior margin of dorsal fins, and an anal-fin base shorter than the base length of the lower caudal-fin lobe (9% of TL vs. 16% of TL, respectively). As mentioned in the remarks under C. plagiosum, C. punctatum is often confused with this species, as well as with C. griseum and C. hasselti. In addition to the previous points of comparison, C. punctatum can be distinguished from C. griseum and C. hasselti by the relative size of its dorsal fins, which are larger than the pelvic fins in C. punctatum, but smaller in the other two species (Compagno and Niem 1998a). The three specimens collected in this study consistently have first dorsal-fin heights (12% or 16% of TL, mean 13% TL) and second dorsal-fin heights (11% or 15% TL, mean 12% TL) larger than the pelvic-fin height (10% TL). Additionally, the base lengths of the first (8% or 12% TL, mean 9% TL) and second dorsal fins (9–12% TL, mean 10% TL) exceeded the pelvic-fin base (6% or 7% TL, mean 7% TL) (see Table 1). The identification of these three specimens was further confirmed using COI data, which showed them forming a distinct clade with the sequence of C. punctatum, separate from those of closely related species (Fig. 3).

Family Scyliorhinidae Gill, 1862

Genus Atelomycterus Garman, 1913

Atelomycterus marmoratus (Bennett, 1830)

IUCN standard name:: Coral Catshark Fig. 6, Table 1

Material examined.

UPVMI 3129, 479 mm TL, female, fish landing sites, Sibunag, Guimaras Province, Philippines, longline, 23 November 2020, coll. M. Asgar.

Morphological description.

Body elongated, slender, and cylindrical. Head short, narrow, and slightly flattened from above. Eyes horizontally oval; spiracles small, situated just behind eyes. Snout short; tip blunt and slightly flattened from above. Mouth long, broadly ovate, with labial furrows extending on both upper and lower jaws. Nostrils large, usually covered by broad, triangular nasal flap extending to mouth. Teeth short, very sharp, tricuspid; central cusp twice length of cusplets on either side. Skin thick, covered with dermal denticles and very sharp to touch. Dorsal fins moderately small with concave posterior margins; first dorsal fin originating behind pelvic fins; second dorsal fin originating from midpoint of anal-fin base. Anal fins similar in shape to dorsal fins but smaller in size. Pectoral fins moderately large; posterior margin slightly convex. Pelvic fin smaller than pectoral fins. Caudal fin relatively short; posterior tip of upper lobe broad; lower lobe indistinct; ventral notch located near tip of upper lobe. Five small gill slits on both sides; last two gill slits situated just above pectoral-fin base.

Color of fresh specimen. Body brown, covered with white spots outlined in black; some spots merged into horizontal bars, notably around head area; underside white with small, faint, brown-black spots extending from the trunk area to caudal fin. Fin edges white with scattered, irregular white and brown-black spots.

Color of preserved specimen. Body brown; underside white. White spots scattered on the body and fins are still visible, but the black outlines have faded.

Distribution.

Indo–Pacific. Western Indian Ocean: Pakistan to southern India; western Pacific: Malaysia, Indonesia, New Guinea, Philippines, Taiwan, and southern China (Human and Ebert 2022).

DNA barcode.

The COI sequence of 607 bp was submitted to GenBank under accession number OR614478.

Conservation status.

Near Threatened

Remarks.

Atelomycterus marmoratus can reach a maximum total length of 700 mm (Compagno and Niem 1998b) and is commonly found in coral reef habitats. The depth range of A. marmoratus remains unclear, however, a specimen was recorded at a minimum depth of 40 m near Bolinao, Philippines (Compagno and Niem 1998b; Compagno et al. 2005). There have been reports of this species in Philippine waters, including Cavite, Cebu, Manila, Negros Occidental (Bacolod), Negros Oriental (Dumaguete), Palawan (Busuanga), Pangasinan (Bolinao), and Sulu (Jolo) (Compagno et al. 2005; Gonzales 2013; Aquino et al. 2023).

The specimen collected for this study was identified as A. marmoratus based on its combined characteristics, including a slender body covered with bordered white spots, two equal-sized dorsal fins, and a long labial furrow. Atelomycterus erdmanni Fahmi et White, 2015 share similar characteristics with A. marmoratus in having white spots on dorsal and lateral surfaces. However, A. marmoratus can be distinguished by having more spots (vs. fewer spots in A. erdmanni), as noted by Fahmi and White (2015). Although direct counts comparing the number of white spots between these two species were not provided, this study observed that the Philippine specimen has approximately five rows of white spots on dorsal view of the head (see Fig. 6B), while the head spots of A. erdmanni consist of about two rows (Fahmi and White 2015, figs. 1, 2). This feature should be confirmed in a larger number of specimens, including those of varying sizes, to verify its consistency across the species. The morphological identification of the Philippine specimen as A. marmoratus is further supported by its COI data, as shown in the phylogenetic tree (Fig. 3), where the Philippine sequence clusters with that of A. marmoratus and is clearly separated from the sequence of A. erdmanni.

Figure 6.

Atelomycterus marmoratus from Western Visayas, Philippines, UPVMI 3129, 479 mm TL. A body in lateral view B head in dorsal view C oronasal region.

Family Carcharhinidae Jordan et Evermann, 1896

Genus Carcharhinus Blainville, 1816

Carcharhinus plumbeus (Nardo, 1827)

IUCN standard name:: Sandbar Shark Fig. 7, Table 1

Material examined.

UPVMI 3119, 391 mm TL, male, Iloilo Fish Port Complex, Iloilo City, Philippines, 15 July 2020, coll. R. Cabebe-Barnuevo, E. Delloro Jr., D. Mediodia, E. Obar, and N. Ylaron. UPVMI 3120, 363 mm TL, male, same data as preceding. UPVMI 3183, male, 370 mm TL, same data as preceding.

Figure 7.

Carcharhinus plumbeus from Western Visayas, Philippines, UPVMI 3119, 391 mm TL (A body in lateral view B oronasal region); UPVMI 3120, 363 mm TL (C body in lateral view D oronasal region).

Morphological description.

Body moderately small and cylindrical. Head slightly depressed. Eyes large and rounded, spiracles absent. Snout short and relatively blunt; length of snout shorter than mouth width (8% vs. 9% TL, respectively). Mouth short and transversely curved. Nostril long with short nasal flap; nostril distance shorter than mouth width (6% vs. 9% TL, respectively). Teeth on upper jaw broadly triangular; teeth on lower jaw still not visible. First dorsal fin tall, height more than half of predorsal length (16% vs. 27% TL, respectively), originating over pectoral-fin insertion; second dorsal fin considerably smaller than first dorsal fin, originating slightly anterior to anal-fin insertion. Anal fin small, considerably same size with second dorsal fin; posterior margin concave. Pectoral fin large; posterior margin concave. Pelvic fin relatively smaller than pectoral fin. Caudal fin relatively long; posterior tip of upper lobe slightly narrow; lower lobe distinct, slightly triangular; ventral notch located near tip of upper lobe. Interdorsal-fin ridge present. Precaudal pits transverse.

Color of fresh specimen. Body and head dark gray; underside white; fins plain with dusky tip.

Color of preserved specimen. Body, head, and fins plain gray; underside portion of body white.

Distribution.

Wide-ranging in tropical through warm temperate areas, including Mediterranean Sea, but possibly not eastern Pacific. Extends from western central Pacific to Hawaii (Compagno and Niem 1998c), and across western Indian Ocean from South Africa (eastern Cape), to the Red Sea, Gulf of Oman, Persian/Arabian Gulf, Pakistan, India, Socotra, Madagascar, Seychelles, Mauritius, and Lakshadweep (Winton and Ebert 2022).

DNA barcode.

COI sequences of 617 bp were submitted to GenBank with accession numbers OR614473 (UPVMI 3119) and OR614471 (UPVMI 3120).

Conservation status.

Endangered

Remarks.

This is the first verified report of Carcharhinus plumbeus from Philippine waters, as previous studies only reported a possible set of jaws that could not be confirmed (Compagno et al. 2005). Globally, populations of C. plumbeus have declined by more than 50% due to fisheries overexploitation and coastal habitat degradation (Rigby et al. 2021). The confirmed presence of small C. plumbeus individuals (smaller than published size-at-birth estimates; Rigby et al. 2021) in Iloilo’s largest fish market indicates an urgent need to enact monitoring and management policies. The largest known specimen of C. plumbeus measures 3 m TL (Soufi-Kechaou et al. 2018). Typically, this species is found at depths ranging from 20–60 m, although it can occasionally inhabit deeper waters up to 280 m (Compagno and Niem 1998c; Last et al. 2010).

The specimens were identified as C. plumbeus based on several morphological characteristics, including the presence of an interdorsal ridge, triangular upper jaw teeth, and first dorsal fin originating above pectoral fin, with its height exceeding half of the predorsal length (16% vs. 27% of TL, respectively). Serena (2005) noted that C. plumbeus is frequently confused with Carcharhinus altimus (Springer, 1950) and Carcharhinus obscurus (Lesueur, 1818). These three species exhibit similarities such as rounded snout, serrated upper teeth, presence of an interdorsal ridge, and fins with plain or dusky tips rather than conspicuously black-tipped fins. However, C. plumbeus is distinguished from these two species by its tall first dorsal fin (vs. short dorsal-fin height, less than half of predorsal length) (Compagno and Niem 1998c). Additionally, C. plumbeus can be distinguished from C. altimus by having a short snout, which is shorter than the mouth width (8% vs. 9% TL, respectively) (vs. long snout with a length equal to or greater than the mouth width in C. altimus). Phylogenetic analysis further supports this identification, showing that C. plumbeus is closely related to C. altimus and C. obscurus, with these three species forming a distinct clade separate from Carcharhinus sorrah (Valenciennes, 1839) (Fig. 3). Moreover, COI barcoding results confirm the identification of the Philippine specimens as C. plumbeus, as they cluster with other sequences of C. plumbeus. The extended distribution in Philippine waters confirmed by this study will support efforts to assess and protect this species.

Carcharhinus sorrah (Valenciennes, 1839)

IUCN standard name:: Spottail Shark Fig. 8, Table 1

Material examined.

UPVMI 3121, 582 mm TL, female, fish landing site, Roxas City, Capiz Province, Philippines, longline, 25 May 2021, coll. M. Batay.

Figure 8.

Carcharhinus sorrah from Western Visayas, Philippines, UPVMI 3121, 582 mm TL. A body in lateral view B head in dorsal view C oronasal region.

Morphological description.

Body moderately stout and spindle shaped. Eyes moderately large and rounded. Snout long and rounded. Mouth short and transversely curved. Nostrils narrow; nasal flaps short. Teeth on both jaws sharp with strongly serrated cusps. First dorsal fin moderately large, originating behind pectoral free rear tip; apex rounded; posterior margin concave; free rear tip long; second dorsal fin very short and small, originating over anal-fin base; free rear tip long. Anal fin small and short, rounded apex; posterior margin concave; free rear tip long. Pectoral fin moderately large, falcate; apex narrow and rounded; posterior margin concave. Pelvic fin moderately small; anterior margin straight. Caudal fin long and smooth; posterior tip of upper caudal-fin lobe narrow; posterior margin of upper lobe concave; ventral notch deep; lower caudal-fin lobe short and triangular; posterior tip of lower lobe rounded. Interdorsal-fin ridge present. Precaudal pits transverse. Gill slits short, with five slits on both sides.

Color of fresh specimen. Body and head gray dorsally; white ventrally. Second dorsal fin, pectoral fins, and lower caudal-fin lobe with conspicuous black tips. First dorsal fin and upper caudal-fin lobe with black margins. Pelvic fins and upper caudal-fin lobe plain color. Posterior margin along apex of anal fin with black margins.

Color of preserved specimen. Body and head gray dorsally; white ventrally. Conspicuous spots, and black edges on fins still distinct.

Distribution.

Found in Indo–Pacific Region including western central Pacific from China southward to Indonesia, Philippines, and Australia (Compagno and Niem 1998c; this study), also in Western Indian Ocean: Persian Gulf, Red Sea to South Africa (KwaZulu-Natal), Madagascar, Seychelles, Mascarenes, and Maldives (Winton and Ebert 2022).

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank with accession number OR614467.

Conservation status.

Near Threatened

Remarks.

Carcharhinus sorrah can grow to a maximum total length of 1600 mm (Allen and Erdmann 2012). This species is commonly found in coastal areas, including continental and insular shelves, both near the shore and at depths of up to 140 m (Compagno and Niem 1998c), as well as in coral reef environments from 1 to 73 m deep (Allen and Erdmann 2012). There have been reports of its presence in Philippine waters, specifically in Cavite, Cebu, Manila Bay, Negros Occidental (Cadiz City, and Sagay), and Palawan (Compagno et al. 2005; Gonzales 2013; Aquino et al. 2023). The specimen was identified as C. sorrah based on the presence of an interdorsal ridge between the dorsal-fin bases, a long and parabolic-shaped snout, strongly black-tipped fins (second dorsal, pectoral, and lower caudal fins), and an extremely low second dorsal fin with a long, attenuated rear tip (second dorsal-fin height 3% TL vs. first dorsal-fin height 13% TL, see Table 1). These features are consistent with the descriptions provided by Compagno and Niem (1998c), and Choi et al. (1998). Additionally, COI barcoding confirms its identification as it forms a single clade with other sequences of C. sorrah (Fig. 3).

Family Pristiophoridae Bleeker, 1859

Genus Pristiophorus Müller et Henle, 1837

Pristiophorus lanae Ebert et Wilms, 2013

IUCN standard name:: Lana’s Sawshark Fig. 9, Table 1

Material examined.

UPVMI 3126, 700 mm TL, male, fish landing site, Roxas City, Capiz Province, Philippines, gillnet, 29 March 2021, coll. M. Batay.

Figure 9.

Pristiophorus lanae from Western Visayas, Philippines, UPVMI 3126, 700 mm TL. A body in dorsal view B oronasal region.

Morphological description.

Body elongated and cylindrical. Head considerably depressed, not elongated. Eyes large, horizontally oval; spiracle large, oblique, located just behind eye. Snout long, flattened, surrounded with saw-like teeth. Mouth short and transversely curved. Pair of barbels present, located closer to mouth than rostral tip. Teeth on rostrum sharp with three distinct sizes; teeth on both jaws small; rows of teeth on upper jaw 35. Dorsal fins moderately large, fully erect, and of similar sizes; first dorsal fin originating posterior to rear tip of pectoral fin; second dorsal fin originating far beyond rear tip of pelvic fin. Anal fin absent. Pectoral fins extremely enlarged; posterior margin slightly concave. Pelvic fin same size as dorsal fins, originating posterior to rear tip of first dorsal fin; posterior margin straight. Caudal fin long, asymmetrical; subterminal notch on upper lobe deep; posterior tip angular. Claspers short and broad. Gill openings small, similar in sizes, with five gill slits on both sides.

Color of fresh specimen. Body and head above uniformly dark brown; underside lighter brown; rostrum brown with two longitudinal lines of darker color; pectoral- and pelvic- fin bases relatively darker brown, lightening towards rear end; no spots or distinct markings.

Color of preserved specimen. Body uniformly brown; underside light brown; longitudinal lines on rostrum still visible.

Distribution.

Philippines (Ebert and Wilms 2013; Koeda and Manjaji-Matsumoto 2017; Weigmann et al. 2020; this study).

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank under accession number OR614466.

Conservation status.

Near Threatened

Remarks.

Pristiophorus lanae was originally described based on specimens collected from various locations in the Philippines, including Batangas, Camarines Sur, Cebu, and Marinduque, at depths ranging from 229 to 593 m (Ebert and Wilms 2013). Additional reports have since been made from Panay Island (Koeda and Manjaji-Matsumoto 2017; Weigmann et al. 2020; Leeney et al. 2024; this study). This species is endemic to the Philippines, with the largest known specimen measuring 900 mm TL, as reported by Weigmann et al. (2020). The identification of the collected specimen was verified through its COI sequence data, which is identical with the other sequence of P. lanae (Fig. 3).

Family Rhinobatidae Müller et Henle, 1837

Genus Rhinobatos Linck, 1790

Rhinobatos whitei Last, Corrigan et Naylor, 2014

IUCN standard name:: Philippine Guitarfish Fig. 10, Table 2

Material examined.

UPVMI 3127, 234 mm DW, 702 mm TL, male, fish landing site, Roxas, City, Capiz Province, Philippines, trawl, September 2021, coll. M. Batay. UPVMI 3128, 43 mm DW, 136 mm TL, juvenile, same data as preceding.

Figure 10.

Rhinobatos whitei from Western Visayas, Philippines, UPVMI 3127, 234 mm DW (A body in dorsal view B head in ventral side C oronasal region); UPVMI 3128, 43 mm DW (D body in dorsal view E head in ventral side F oronasal region).

Morphological description.

Body moderately elongated and medium-sized, covered with tiny dermal denticles; trunk depressed and flattened. Disc broad and wedge-shaped. Head large and depressed. Eyes large, oval and protruding; spiracles very large, located just behind eye. Snout relatively long; tip bluntly pointed; anterior margin concave; rostral cartilages well separated and converging towards front area. Mouth moderately wide and slightly arched. Nostrils large, elongated, oval rather than rounded; nasal flap very small, narrow, not covering nostrils. Teeth very small, rhomboid, very compact; upper jaw with 83 rows; lower jaw with 77 rows. Dorsal fins falcate, almost equal in size, posterior margin straight. Pectoral fins flat and very large. Pelvic fin short, angular and firm, originating just behind free rear tip of pectoral fins (in juvenile specimen, free rear tip of pectoral fins not reaching insertion of pelvic fins); apex bluntly rounded. Caudal fin moderately large, asymmetrical; lower caudal-fin lobe absent. Gill slits five on each side; fifth slit obviously smallest.

Color of fresh specimen. Adult specimen (UPVMI 3127): Body with dark-brown spots; underside white; dorsal fins with black free rear tip. Juvenile (UPVMI 3128): Body dorsal surface brown with numerous small, white and brown-black spots; underside white; dorsal fins with black free rear tip; pectoral- and pelvic-fin edges creamy white.

Color of preserved specimen. Adult: Body brown with numerous tiny pale white to gray spots, and large brown to light brown spots (spots more pronounced than in fresh specimen); dorsal and caudal fins with white free rear tip. Juvenile: Body light brown with numerous tiny white and brown spots; dorsal fins with black free rear tip.

Distribution.

Philippines (Last et al. 2014; Manjaji-Matsumoto 2017; this study).

DNA barcode.

COI sequences of 617 bp were submitted to GenBank under accession numbers OR614481 (UPVMI 3127) and OR614470 (UPVMI 3128).

Conservation status.

Critically Endangered

Remarks.

Rhinobatos whitei was originally described based on specimens collected from the markets in central and southern Philippines (specifically in Bacolod, Cebu, Dipolog, and Dumaguete), with a size range of 317–844 mm TL (Last et al. 2014). Subsequent records have identified this species from Panay Island, Philippines (Manjaji-Matsumoto 2017; this study), and to date, there have been no reports of its occurrence in other countries, affirming its status as a Philippine endemic. Rhinobatos whitei has been assessed as critically endangered due to severe declines in ray populations throughout the Philippines, coupled with the highly restricted geographic range of the species (Dulvy et al. 2021). This study reports the first collection of a juvenile specimen, and knowledge on juveniles may be important in protecting the species.

Rhinobatos whitei and Rhinobatos manai White, Last et Naylor, 2016 share the characteristic of having whitish spots, which distinguishes them from other congeners (White et al. 2016). However, the collected specimen was identified as R. whitei based on its poorly defined white spots and presence of large, diffuse dusky and orange blotches as described by Last et al. (2014). In contrast, R. manai is characterized by well-defined rusty brown spots and faint white spots with grayish edges, as described by White et al. (2016). Although no COI data are available for either species, the identification of the collected specimen was confirmed based on the morphological descriptions and coloration comparisons provided above.

Family Plesiobatidae Nishida, 1990

Genus Plesiobatis Nishida, 1990

Plesiobatis daviesi (Wallace, 1967)

IUCN standard name:: Giant Stingaree Fig. 11, Table 2

Material examined.

UPVMI 3112, 368 mm DW, male, fish landing site, Roxas City, Capiz Province, Philippines, trawl, September 2021, coll. M. Batay.

Figure 11.

Plesiobatis daviesi from Western Visayas, Philippines, UPVMI 3112, 368 mm DW. A body in ventral side B body in dorsal view C oronasal region.

Morphological description.

Body large and depressed, upper surface covered with small dermal denticles; trunk depressed; disc large. Head broad and depressed. Eyes horizontally enlarged, situated dorsolaterally on head; spiracles wide, bigger than eyes, and located just behind them. Snout moderately elongated and broadly pointed; smooth to touch. Mouth straight, and moderately broad. Nostrils extremely wide, circular, and located near mouth; nasal flap short, partially covering nostrils, and fused with broad, short nasal curtains extending to anterior corners of mouth. Teeth small, rounded oval, with 34 rows on both jaws. Pectoral fins very large, smooth, forming almost rounded disc. Pelvic fins small, angular rather than rounded. Gill slits located on underside small, with five on each side. Claspers short and broad. Tail was cut off upon purchase.

Color of fresh specimen. Dorsal area brown, underside white; rear edges of disc and pelvic fins black-brown; no spots or distinct markings observed.

Color of preserved specimen. Dorsal area brown; underside white; black edges on disc and pelvic fins still visible.

Distribution.

Indo–Pacific. western Indian Ocean: South Africa (KwaZulu-Natal) to southern Mozambique, and southern India to Sri Lanka; western Pacific: Philippines, Taiwan, Kyushu–Palau Ridge (Philippine Sea), Ryukyu Islands, Australia, and Hawaii (Ebert and Aschliman 2022).

DNA barcode.

The COI sequence of 613 bp was submitted to GenBank under accession number OR614468.

Conservation status.

Least Concern

Remarks.

Plesiobatis daviesi is currently the only recognized valid species in the family Plesiobatidae. This study confirms the occurrence of P. daviesi in the Western Visayas region, adding to its previous records in Negros Oriental (Dumaguete), northern Luzon, and Zamboanga del Norte (Dapitan City) (Compagno and Last 1999; Compagno et al. 2005). Typically, this species is found on continental slopes at depths ranging from 275 to 680 m (Compagno and Last 1999; Last et al. 2010). It can reach a maximum total length of 2700 mm (Compagno and Last 1999), while Philippine records indicate a size range of 507–1550 mm in TL, and 288–800 mm in DW (Compagno et al. 2005). The collected specimen was identified as P. daviesi based on its large body, pointed snout, soft and fleshy disc, and short, non-whiplike tail. The generated COI sequence formed a single clade with other sequences of P. daviesi, supporting the morphological identification, as shown in Fig. 12.

Figure 12.

Maximum likelihood phylogenetic tree of partial COI sequences from batoids, constructed using the Kimura 2-parameter model. Philippine sequences are identified by UPVMI voucher codes, with related genera color-coded. Bootstrap support values (1,000 replicates) are shown at nodes, and the scale bar represents evolutionary distance in substitutions per site.

Family Urolophidae Müller et Henle, 1841

Genus Urolophus Müller et Henle, 1837

Urolophus sp.

Fig. 13

Material examined.

UPVMI 3116, 191 mm DW, female, Iloilo Fish Port Complex, Iloilo City, Philippines, 8 October 2020, coll. R.P. Babaran.

DNA barcode.

COI sequences were submitted to GenBank under accession numbers OR614469 (UPVMI 3116, 617 bp) and PQ461195 (KAUM–I. 62938, 642 bp).

Remarks.

The collected specimen (UPVMI 3116) was initially identified as Urolophus aurantiacus (Müller et Henle, 1841) based on the color photographs and morphological characteristics similar to a specimen (KAUM–I. 62938) also collected from Panay Island, which was identified as U. aurantiacus by Manjaji-Matsumoto (2017). Both specimens from Panay Island exhibit rounded disc apices, a short and pointed snout, and a dorsal spine on the short tail. However, COI gene sequence data from both specimens (KAUM–I. 62938 and UPVMI 3116) showed significant divergence from the available sequences of U. aurantiacus. Instead, the sequences from both specimens cluster more closely with Urolophus expansus McCulloch, 1916 (see Fig. 13).

Figure 13.

Maximum likelihood phylogenetic tree of partial COI sequences based on the Kimura 2-parameter model, showing the relationship of Urolophus sp. (UPVMI 3116 and KAUM–I. 62938) with related sequences of Urolophus spp. from BOLD and GenBank databases, with Chimaera phantasma as the outgroup. Bootstrap support values (1,000 replicates) are shown at nodes, and the scale bar represents evolutionary distance in substitutions per site.

However, U. expansus is characterized by blotches on the tail area and faint colored bars in front of and behind the eyes (McCulloch 1916; Ebert 2014), which are absent in both specimens from Panay Island. Moreover, U. expansus has only been reported from western and south Australia, in the southeastern Indian Ocean (Fricke et al. 2024). Despite the observed genetic similarity, morphological and biogeographical evidence suggests that the Panay specimens represent a putative unidentified species. Further investigation and comparisons are required to confirm the identification of these specimens.

Family Dasyatidae Jordan, 1888

Genus Himantura Müller et Henle, 1837

Himantura sp.

Figs 14, 15, Table 2

Material examined.

UPVMI 3111, 377 mm DW, male, fish market, Carles, Iloilo Province, Philippines, February 2022, coll. R.P. Babaran.

Figure 14.

Himantura sp. from Western Visayas, Philippines, UPVMI 3111, 377 mm DW. A body in dorsal view B oronasal region.

Morphological description.

Body large; dorsal surface with median row of denticles extending from eye area to tail base, and two heart-shaped thorns located in center; disc wide, rhomboidal. Eyes moderately large; spiracles very wide with few denticles. Snout triangular with pointed tip. Mouth narrow; labial furrows not developed. Nostrils extremely wide, located just above mouth; nasal flap skirt-shaped, very short, not reaching mouth. Pectoral-fin apex rounded to angular; front margins slightly concave; posterior margins convex. Pelvic fin short, triangular with broad apex. Tail was cut off upon purchase. Gill slits located on underside, moderately small, with five slits on each side.

Color of fresh specimen. Dorsal surface and fins light brown with numerous, irregular brown spots; ventral area white.

Color of preserved specimen. Dorsal surface dark brown with light brown margins; spots still visible; ventral area pale white.

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank with accession number OR614479.

Remarks.

Borsa et al. (2021) identified seven distinct species within the genus Himantura, including Himantura australis Last, White et Naylor, 2016a, Himantura leoparda Manjaji-Matsumoto et Last, 2008, Himantura tutul Borsa, Durand, Shen, Alyza, Solihin et Berrebi, 2013, Himantura undulata (Bleeker, 1852), Himantura uarnak (Gmelin, 1789), Himantura sp. 1, and Himantura sp. 2. Sequences representative of each species were selected to construct a phylogenetic tree aimed at determining the identity of the specimen from Panay Island, with H. tutul used as outgroup (Fig. 15). The results indicate that the collected specimen clusters within the clade corresponding to Himantura sp. 1 sensu Borsa et al. (2021). The identification of this specimen remains provisional, pending the formal description of Himantura sp. 1 as outlined by Borsa et al. (2021).

Figure 15.

Maximum likelihood phylogenetic tree based on the Kimura 2-parameter model, constructed using COI gene sequence of UPVMI 3111 (this study) and verified sequences of Himantura spp. from Borsa et al. (2021). Bootstrap support values (1,000 replicates) are shown at nodes, and the scale bar represents evolutionary distance in substitutions per site.

Genus Neotrygon Castelnau, 1873

Neotrygon orientalis Last, White et Serét, 2016

IUCN standard name:: Oriental Bluespotted Maskray Fig. 16, Table 2

Material examined.

UPVMI 3113, 256 mm DW, female, fish market, Aklan Province, Philippines, 22 November 2020, coll. M. Macavinta.

Figure 16.

Neotrygon orientalis from Western Visayas, Philippines, UPVMI 3113, 256 mm DW. A body in dorsal view B body in ventral side C oronasal region.

Morphological description.

Body moderate in size; dorsal area with median row of small thorns located on center region; disc not greatly expanded, rhomboid. Eyes moderately large and elongated; spiracles very wide. Snout short, slightly angular with rounded tip. Mouth small. Nostrils moderately oval, partially covered with nasal flap; nasal flap skirt-shaped, slightly reaching mouth. Pectoral-fin apex angular; front margins convex; posterior margins straight. Pelvic fin short, subtriangular. Tail tapering, not whip-like; dorsal and ventral skin folds present; spine on caudal fin already removed upon purchase. Gill slits located on underside moderately short, with five slits on each side.

Color of fresh specimen. Dorsal area pale brown with light blue spots outlined with darker blue of various sizes; ventral area white, becoming brown on edges; faint brown band present across the eyes; eyes and spiracles surrounded with numerous, tiny black spots, with a few similar black spots widely scattered on the dorsal area; tail dark brown with irregular black and white bands.

Color of preserved specimen. Central region of the dorsal area dark brown to black; disc area light brown; blue spots transformed into dark brown; ventral area pale white, becoming brown on edges; tiny black spots still visible; black and white bands on tail still distinct.

Distribution.

Australia, New Guinea, Indonesia, Malaysia, Borneo, Philippines, and Taiwan (Last et al. 2016b; Manjaji-Matsumoto 2017; Borsa et al. 2018)

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank under accession number OR614465.

Conservation status.

Least Concern

Remarks.

Neotrygon orientalis was originally described based on 12 specimens collected from Indonesia, with a size range of 145–343 mm DW, although it can reach a maximum DW of 380 mm (Last et al. 2016b; Manjaji-Matsumoto 2017). This species is part of the broader Neotrygon kuhlii (Müller et Henle, 1841) complex. In this study, the sequence generated from the Philippine specimen was compared to verified COI gene sequences of Neotrygon spp. provided by Last et al. (2016b) and Hata and Motomura (2024). Phylogenetic tree analysis showed that the Philippine specimen clustered with the verified COI sequences of N. orientalis (Fig. 12). Therefore, based on both morphological characteristics and COI data, the collected specimen was identified as N. orientalis.

The precise distribution of N. orientalis in Philippine waters remains unclear, as Last et al. (2016b) noted that specimens from the Philippines previously identified as N. kuhlii by Naylor et al. (2012) and Compagno et al. (2005) might, in fact, be N. orientalis. Nonetheless, reports by Manjaji-Matsumoto (2017), and this study provide confirmed evidence of N. orientalis from Panay Island, Philippines.

Genus Taeniura Müller et Henle, 1837

Taeniura lymma (Forsskål, 1775)

IUCN standard name:: Bluespotted Lagoon Ray Fig. 17, Table 2

Material examined.

UPVMI 3114, 104 mm DW, female, fish landing site, Sibunag, Guimaras Province, Philippines, 17 October 2020, coll. J. Fernandez.

Figure 17.

Taeniura lymma from Western Visayas, Philippines, UPVMI 3114, 104 mm DW. A body in dorsal view B body in ventral side C oronasal region.

Morphological description.

Body small; disc oval. Eyes moderately large and protruding; spiracles large, located just behind eyes. Snout short and broadly angular. Mouth very small. Nostrils moderately short and narrow; nasal flap short, partially covering nostrils; posterior margin fringed, extending beyond mouth. Tooth rows on both jaws, pavement-like; 15 rows on upper jaw, 16 on lower jaw. Pectoral fin horizontally elongated with smooth, rounded margins. Pelvic fin angular. Tail short and stout; lower caudal-fin fold pronounced, reaching tail tip; single strong spine along midsection. Gill slits on underside small, with five slits on each side.

Color of fresh specimen. Dorsal body surface brown to olive-green; underside white; disc covered with bright, large blue spots. Fins brown; pectoral and pelvic fins with a few smaller, denser blue pots. Tail with a pair of black, parallel lines on its side; underside of tail white; caudal-fin fold edges brown.

Color of preserved specimen. Dorsal body surface brown; underside white. Blue spots on disc area and fins transformed into dark brown to black. Pair of black, parallel lines on tail still visible; caudal-fin fold edges dark brown.

Distribution.

Widely distributed in tropical to warm-temperate seas, including the Mediterranean Sea (Manjaji-Matsumoto et al. 2022).

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank with accession number OR614476.

Conservation status.

Least Concern

Remarks.

Taeniura lymma, known for its vibrant blue spots, can grow up to 750 mm in TL and 350 mm DW (Last et al. 2010). This coastal species is widely found throughout Philippine waters, inhabiting depths down to 20 m. It has a broad distribution and is the most common stingray in coral reef areas (Last and Compagno 1999; Manjaji-Matsumoto 2017). Reports of this species within the country include Cebu, Guimaras Island, Negros Oriental (Siaton), Panay Island, Palawan, Siquijor Island, Sulu (Jolo, Sibutu, Sitankai), and Zamboanga (Compagno et al. 2005; Gonzales 2013; Manjaji-Matsumoto 2017; Aquino et al. 2023). The collected specimen was easily identified as T. lymma based on its broad tail with two stings, a pair of parallel black lines, and its smooth, oval disc profile covered with numerous vivid blue spots. This identification was confirmed by its COI data, which clustered with other sequences of T. lymma (see Fig. 12).

Family Aetobatidae Agassiz, 1858

Genus Aetobatus Blainville, 1816

Aetobatus ocellatus (Kuhl in Kuhl and van Hasselt, 1823)

IUCN standard name:: Spotted Eagle Ray Fig. 18, Table 2

Material examined.

UPVMI 3115, 574 mm DW, female, fish landing site, Sibunag, Guimaras Province, Philippines, 15 October 2020, coll. J. Fernandez.

Figure 18.

Aetobatus ocellatus from Western Visayas, Philippines, UPVMI 3115, 574 mm DW. A body in dorsal view B body in ventral side C oronasal region.

Morphological description.

Body large; dorsal surface smooth; denticles and thorn absent from middle area; disc rhomboidal and broad, rather wide than long. Eyes moderately small and rounded; spiracles very large, located behind eye rather than below, and completely visible from dorsal view. Snout soft, large, protruding with blunt tip. Mouth moderately large. Nostrils large, partially covered by nasal flap; nasal flap moderately large, V-shaped, with posterior tips reaching lower jaw. Teeth plate on upper jaw rather wide than long in comparison to teeth plate on lower jaw; teeth on lower jaw elongated, extending beyond jaw, making it exposed when mouth is closed. Dorsal fin small, subtriangular with rounded apex. Pectoral fin very large, triangular with rounded apex, originating behind eye; anterior margin concave basally, becoming straight towards middle area, then convex towards rear; posterior margins concave in front, becoming straight towards rear; anterior edge entire or smooth and thick; posterior edge crenate and thin. Pelvic fin moderately elongated with rounded and crenate posterior margin. Tail tapering, very long, whip-like; dorsal and ventral folds absent; two spines positioned just behind dorsal fin; first spine was already removed upon purchase; second spine long, tapering with sharp, serrated margins. Gill slits located on underside short, with five on each side.

Color of fresh specimen. Dorsal area gray to black covered with small white spots; ventral area white with black fin edges; spots mostly concentrated on the pectoral and pelvic fins; few similar spots on the nape area; tail black.

Color of preserved specimen. Dorsal area dark brown; ventral area pale white; white spots still visible; tail black.

Distribution.

Tropical and warm temperate seas of the Indo-West Pacific. Red Sea, South Africa, Pakistan, India, Sri Lanka, Hawaii, Japan, Philippines, and Australia (Last et al. 2010; de Carvalho and Soares 2022; this study).

DNA barcode.

The COI sequence of 617 bp was submitted to GenBank under accession number OR614475.

Conservation status.

Vulnerable.

Remarks.

Aetobatus ocellatus can grow to a disc width of 3000 mm (White et al. 2010) and is often found at depths of 20–25 m (Randall and Cea 2011). Aetobatus ocellatus was categorized as “uncertain (?)” (Alava et al. 2014; Santos et al. 2017); however, the presently reported study confirms its occurrence in the country (see Table 3). The distribution of this species within Philippine waters should be verified, as previous reports on Aetobatus narinari (Euphrasen, 1790) from the Philippines may have referred to A. ocellatus.

Table 3.

Download as

CSV

XLSX

Additional confirmed cartilaginous fish species in Philippine waters.

Species	Status based on Santos et al. (2017)	Remarks
Carcharhinus plumbeus	Not included in the original list	New record from the country with institutional codes UPVMI 3119 (391 mm TL), UPVMI 3120 (363 mm TL), and UPVMI 3183 (370 mm TL) (this study).
Pristiophorus lanae	“Confirmed, NO species account (N)”	Specimens deposited in the UPV-Museum of Natural Sciences includes UPVMI 1648 (786.4 mm TL) (Koeda and Manjaji-Matsumoto 2017), and UPVMI 3126 (700 mm TL) (this study).
Rhinobatos whitei	“Confirmed, NO species account (N)”	Specimens deposited in the UPV-Museum of Natural Sciences include UPVMI 576 (361.1 mm TL) (Manjaji-Matsumoto 2017), UPVMI 3127 (702 mm TL), and UPVMI 3128 (136 mm TL) (this study).
Urolophus sp.	Not included in the original list	One specimen (UPVMI 3116) was deposited in the UPV-Museum of Natural Sciences, and the other specimen was reported as Urolophus aurantiacus by Manjaji-Matsumoto (2017) and has been deposited in the Kagoshima University Museum (KAUM–I. 62938). However, these specimens may represent an undescribed species, as suggested in this study (see remarks under Urolophus sp.).
Himantura sp. 1 sensu Borsa et al. (2021)	Not included in the original list	Borsa et al. (2017) identified two unknown species of Himantura, and the specimen from this study clustered with Himantura sp. 1 (see remarks under Himantura sp.).
Aetobatus ocellatus	“Uncertain (?)”	A specimen was described in this study and deposited in the UPV-Museum of Natural Sciences (UPVMI 3115).

The presently reported specimen was identified as A. ocellatus based primarily on its dark green to black dorsal surface, and the white spots, which were more restricted to the dorsal disc surface and did not extend to the head or eye regions. This identification was supported by molecular evidence (Fig. 12). On the other hand, color photographs provided by White et al. (2010), show that the white spots on the dorsal surface of A. narinari are evenly distributed throughout the eye, head, and disc areas.

White et al. (2010) identified the major distinction between A. ocellatus and A. narinari as the background coloration on dorsal surface: A. ocellatus exhibit dark greenish, grayish, or nearly blackish color, sometimes with a pinkish tinge, whereas A. narinari is much paler yellowish-brown. Moreover, these two species show minor morphometric differences: A. ocellatus has a mean total length (measured from snout tip to tip of tail) of 281% DW and a mean tail length (from anteriormost edge of the cloaca to the tail tip) of 230% DW, whereas A. narinari measures a total length of 263% and a tail length of 208% DW. The Philippine specimen, with a total length of 271% and a tail length of 219% DW, is closer to A. ocellatus (4% difference) in total length, compared to A. narinari (3%). Both species show a similar 5% difference in tail length relative to the Philippine specimen. Therefore, based on its dark green to black dorsal coloration, COI data, and total length, the Philippine specimen aligns more closely with A. ocellatus.

Discussion

This study provides insights into the taxonomy and diversity of cartilaginous fishes in the Western Visayas, particularly in Panay and Guimaras islands, an important fishery and biodiversity area of the Philippines. A key aspect of this research is the provision of new vouchered specimens, which are accessible for further research and are stored at the University of the Philippines Visayas, Museum of Natural Sciences in Iloilo. These specimens, along with the morphological descriptions and generated COI sequence data, offer valuable resources for supporting future research, ensuring progress in understanding the cartilaginous fish diversity in the region.

DNA barcoding has been widely used to confirm species identifications among cartilaginous fishes, resulting in the discovery of new species, differentiation of cryptic species, recognition of putative species, and other applications such as monitoring biodiversity, exploring evolutionary relationships, and supporting conservation efforts by identifying species at risk (Ward et al. 2008; Last et al. 2008, 2014, 2016a, 2016b; Richards et al. 2009; Griffiths et al. 2010; Fahmi and White 2015; White et al. 2016; Finucci et al. 2018; Borsa et al. 2021; Carugati et al. 2022; Loh et al. 2023; Hata and Motomura 2024; Lee et al. 2024). In this study, the COI sequences provide critical verification of cartilaginous fishes collected from Panay and Guimaras islands, serving as a complementary tool to traditional morphological descriptions and photographic documentation to enhance the accuracy and reliability of species identification.

A phylogenetic tree was constructed to illustrate the placement of the sequences generated in this study relative to similar and closely related species, as highlighted in the remarks for each species profile. As shown in Figs 3, 12, the generated sequences cluster with those of similar species, supporting the accuracy of their identification. Although COI sequences for Rhinobatos whitei are currently not available in public databases, the identity of the R. whitei specimen collected in this study was confirmed based on its distinct morphological features (see Remarks). Furthermore, separate phylogenetic trees were constructed for the two putative new species identified in this study (Urolophus sp. and Himantura sp.), highlighting their phylogenetic relationships and distinctiveness from other species within their respective genera (Figs 13, 15, respectively; see also Remarks).

By integrating molecular and morphological techniques, this study offers a comprehensive and accurate inventory of chimaeras, sharks, and batoids in the region, revealing a new record and two unidentified, putatively novel species. Consequently, this study expands upon previous inventories, including Santos et al. (2017) listing, by documenting a total of fourteen cartilaginous fish species. The list includes the sandbar shark, Carcharhinus plumbeus, which is reported here as a new record from Philippine waters. Additionally, the discovery of potential undescribed species within the genera Himantura and Urolophus highlights the importance of continued research. This results in a total of 209 cartilaginous fishes in the country, an increase from the previous record of 206 species (Santos et al. 2017; see Table 3).

In line with this inventory, Pristiophorus lanae and Rhinobatos whitei are both endemic to the Philippines and were categorized as “Confirmed, NO species account (N)” (Santos et al. 2017). However, these two species were also reported by Koeda and Manjaji-Matsumoto (2017), as well as by Manjaji-Matsumoto (2017), and are documented in this study from Panay Island (see Table 3). On the other hand, the occurrence of Aetobatus ocellatus in the Philippines was categorized as “uncertain (?)” (Santos et al. 2017), nevertheless, a specimen of this species was collected during this study. The voucher specimens deposited in UPV-Museum of Natural Sciences, along with the COI barcodes, are valuable for confirming the species accounts of these three species.

Conclusions

This study contributes to our understanding of cartilaginous fish diversity in the Western Visayas, specifically in Panay and Guimaras islands, through the integration of molecular (COI) and morphological data. This approach increases the likelihood of detecting undescribed species and cryptic species (e.g., Himantura sp. 1, Urolophus sp.), and also aids in rapid confirmation of new country records (e.g., Carcharhinus plumbeus). Moreover, documentation of endemic species (Pristiophorus lanae and Rhinobatos whitei) also plays a crucial role in conservation efforts by contributing to the accurate cataloging of species at risk.

By providing newly vouchered specimens, along with relevant information such as morphological descriptions, COI data, and color photographs, this study lays a strong foundation for future research, ensuring that these findings are robust and accessible for future studies. Phylogenetic analysis further supports the accuracy of the identified species, providing comparative context through sequences from similar and closely related species. The inclusion of new records and recognition of uncertain species expands the known diversity of cartilaginous fishes, not only in the Visayas region but across the Philippines.

Information gathered in this study is essential for effective conservation planning and sustainable management of marine resources. Given the impact of climate change and destructive human activities such as overfishing and coastal restructuring, it is imperative that taxonomic studies receive greater attention from the scientific community to prevent the extinction of species before they are even known, as exemplified by the critically endangered Rhinobatos whitei, which was only described in 2014.

Author contributions

Conceptualization: RCB, HM, RPB, MCM. Data curation: RCB, DFP, ESD. Formal analysis: RCB. Funding acquisition: RPB. Investigation: RCB, DFP, HM, RPB, MCM. Visualization: RCB, ESD. Project administration: RPB, MCM. Writing – original draft: RCB. Writing – review and editing: RCB, DFP, ESD, RPB, HM, MCM.

Acknowledgments

We gratefully acknowledge the funding support under the UP System Emerging Inter-Disciplinary Research Program (OVPAA-EIDR-C08-011-R) and Leverage fund from the Office of the Vice Chancellor for Research and Extension (OVCRE), University of the Philippines Visayas (2020-13-SP). The first author is grateful for the scholarship support provided by the Japanese Ministry of Education, Culture, Sports, Science and Technology (MEXT).We thank the Department of Agriculture Bureau of Fisheries and Aquatic Resources (DA-BFAR), Provincial Government Units, the Office of Provincial Agriculture (PAO) from the provinces of Aklan, Antique, Capiz, Iloilo, and Guimaras, especially Ms. Grace-an Villareal Perlas (PAO staff, Aklan), and Ms. Ramie Lyn G. Bañares (PAO staff, Capiz), the Municipal Agriculture Office (MAO) of Batan and Tangalan Aklan, especially to Ms. Glenda Sanchez (MAO, Batan), the Local Government Units (Guimbal, Tangalan, Batan, Tibiao, Pandan, San Jose, Sibunag, Cabalagnan, Batad, and Roxas) for their administrative support, generous assistance and contributions throughout the project. Contributions of our field enumerators (A. Abisan, P. Aguilos Jr., M. Asgar, M. Batay, J. Fernandez, S. Gelera, M.C. Macavinta, S. Sucgang, and J. Tubillara) are gratefully acknowledged. We are also grateful to the Philippine Genome Center-Visayas Satellite Facility (PGC-VSF), the National Institute of Molecular Biology and Biotechnology (UPV-NIMBB), and the UPV Museum of Natural Sciences (UPVMI) for allowing use of their facilities. Our gratitude and appreciation to D. Mediodia, K.D. Barnuevo, E. Obar, M.V. Aranjuez, N. Ylaron, L. Mooc, C. Garinggan, J. Ariñez, A.G. Deallo, N. Cartago, J. Velo, S. Garcia, and C. Javier for their generous help throughout the project.

References

Alava MN, Gaudiano JP, Utzurrum JT, Capuli EE, Aquino MTR, Luchvez-Maypa MMA, Santos MD (2014) Pating Ka Ba?: An identification guide to sharks, batoids and chimaeras of the Philippines. Department of Agriculture Bureau of Fisheries and Aquatic Resources, National Fisheries Research and Development Institute and the Marine Wildlife Watch of the Philippines, 200 pp.

Allen GR, Erdmann MV (2012) Reef fishes of the East Indies. Volumes I–III. Tropical Reef Research, Perth, Australia. University of Hawai’i Press, Honolulu, HI, USA, 1260 pp.

Aquino MTR, Alava MNR, Utzurrum J, Abalo DL, Gaudiano JPA, Santos MD (2023) Annotated checklist of cartilaginous fishes in Palawan waters. Philippine Journal of Fisheries 30(2): 162–198. https://doi.org/10.31398/tpjf/30.2.2022-0021

Baum JK, Myers RA, Kehler DG, Worm B, Harley SJ, Doherty PA (2003) Collapse and conservation of shark populations in the northwest Atlantic. Science 299(5605): 389–392. https://doi.org/10.1126/science.1079777

Bennett ET (1830) Class Pisces. In: Raffles SH (Ed.) Memoir of the life and public services of Sir Thomas Stamford Raffles. Volume 1. Duncan, London, 686–694.

Bleeker P (1852) Bijdrage tot de kennis der Plagiostomen van den Indischen Archipel. Verhandelingen van het Bataviaasch Genootschap van Kunsten en Wetenschappen 24: 1–92. https://doi.org/10.1163/22134379-90000832

Borsa P, Durand JD, Shen KN, Arlyza IS, Solihin DD, Berrebi P (2013) Himantura tutul sp. nov. (Myliobatoidei: Dasyatidae), a new ocellated whipray from the tropical Indo-West Pacific, described from its cytochrome-oxidase I gene sequence. Comptes Rendus Biologies 336(2): 82–92. https://doi.org/10.1016/j.crvi.2013.01.004

Borsa P, Arlyza IS, Hoareau TB, Shen KN (2018) Diagnostic description and geographic distribution of four new cryptic species of the blue-spotted maskray species complex (Myliobatoidei: Dasyatidae; Neotrygon spp.) based on DNA sequences. Journal of Oceanology and Limnology 36(3): 827–841. https://doi.org/10.1007/s00343-018-7056-2

Borsa P, Williams CT, McIvor AJ, Hoareau TB, Berumen ML (2021) Neotype designation and re-description of Forsskål’s reticulate whipray Himantura uarnak. Marine Biodiversity 51(2): 1–28. https://doi.org/10.1007/s12526-021-01180-1

Carugati L, Melis R, Cariani A, Cau A, Crobe V, Ferrari A, Follesa MC, Geraci ML, Iglésias SP, Pesci P, Tinti F, Cannas R (2022) Combined COI barcode‐based methods to avoid mislabelling of threatened species of deep‐sea skates. Animal Conservation 25(1): 38–52. https://doi.org/10.1111/acv.12716

Choi Y, Kim IS, Nakaya K (1998) A taxonomic revision of genus Carcharhinus (Pisces: Elasmobranchii) with Description of Two New Records in Korea. Animal Systematics, Evolution and Diversity 14(1): 43–49.

Compagno LJV (1999) Chimaeras. In: Carpenter KE, Niem VH (Eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes, part 1 (Elopidae to Linophrynidae). FAO, Rome, 1531–1537.

Compagno LJV (2001) Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Volume 2. Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes and Orectolobiformes). FAO, Rome, 269 pp.

Compagno LJV, Last PR (1999) Plesiobatidae. In: Carpenter KE, Niem VH (Eds) FAO Species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes, part 1 (Elopidae to Linophrynidae). FAO, Rome, 1467–1468.

Compagno LJV, Niem VH (1998a) Hemiscylliidae. In: Carpenter KE, Niem VH (Eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians, and sharks. FAO, Rome, 1249–1259.

Compagno LJV, Niem VH (1998b) Scyliorhinidae. In: Carpenter KE, Niem VH (Eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians, and sharks. FAO, Rome, 1279–1292.

Compagno LJV, Niem VH (1998c) Carcharhinidae. In: Carpenter KE, Niem VH (Eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians, and sharks. FAO, Rome, 1312–1360.

Compagno LJV, Last PR, Stevens JD, Alava MNR (2005) Checklist of Philippine Chondrichthyes. CSIRO Marine Laboratories Report 243, 103 pp.

Cortés E, Brooks EN, Shertzer KW (2015) Risk assessment of cartilaginous fish populations. ICES Journal of Marine Science 72(3): 1057–1068. https://doi.org/10.1093/icesjms/fsu157

de Carvalho MR, Soares MC (2022) Family Aetobatidae. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa: 624–626, pls. 78–79.

Dulvy NK, Fowler SL, Musick JA, Cavanagh RD, Kyne PM, Harrison LR, Pollock CM (2014) Extinction risk and conservation of the world’s sharks and rays. eLife 3: e00590. https://doi.org/10.7554/eLife.00590

Dulvy NK, Derrick D, Rigby CL, Tanay D, Utzurrum JAT (2021) Rhinobatos whitei. The IUCN Red List of Threatened Species 2021: e.T104007203A104007215. https://doi.org/10.2305/IUCN.UK.2021-2.RLTS.T104007203A104007215.en

Ebert DA (2014) Deep–sea Cartilaginous Fishes of the Indian Ocean. Volume 2. Batoids and Chimaeras. FAO Species Catalogue for Fishery Purposes. Rome, FAO, 129.

Ebert DA, Aschliman NC (2022) Family Plesiobatidae, Deepwater stingray. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa, 598, pl. 69.

Ebert DA, Wilms HA (2013) Pristiophorus lanae sp. nov., a new sawshark species from the Western North Pacific, with comments on the genus Pristiophorus Müller and Henle, 1837 (Chondrichthyes: Pristiophoridae). Zootaxa 3752(1): 86–100. https://doi.org/10.11646/zootaxa.3752.1.7

Ebert DA, White WT, Ho HC, Last PR, Nakaya K, Seret B, Straube N, Naylor GJP, de Carvalho MR (2013) An annotated checklist of the chondrichthyans of Taiwan. Zootaxa 3752(1): 279–386. https://doi.org/10.11646/zootaxa.3752.1.17

Euphrasen BA (1790) Raja (Narinari). Kongliga Vetenskaps Akademiens nya Handlingar, Stockholm, 11, 217–219.

Fahmi F, White W (2015) Atelomycterus erdmanni, a new species of catshark (Scyliorhinidae: Carcharhiniformes) from Indonesia. Journal of the Ocean Science Foundation 14: 14–27.

Ferretti F, Worm B, Britten GL, Heithaus MR, Lotze HK (2010) Patterns and ecosystem consequences of shark declines in the ocean. Ecology Letters 13(8): 1055–1071. https://doi.org/10.1111/j.1461-0248.2010.01489.x

Field IC, Meekan MG, Buckworth RC, Bradshaw CJA (2009) Susceptibility of sharks, rays and chimaeras to global extinction. Advances in Marine Biology 56: 275–363. https://doi.org/10.1016/S0065-2881(09)56004-X

Finucci B, White WT, Kemper JM, Naylor GJP (2018) Redescription of Chimaera ogilbyi (Chimaeriformes; Chimaeridae) from the Indo-Australian region. Zootaxa 4375(2): 191–210. https://doi.org/10.11646/zootaxa.4375.2.2

Forskål P (1775) Descriptiones Animalium, Avium, Amphibiorum, Piscium, Insectorum, Vermium; quae in Itinere Orientali Observavit Petrus Forskål. Post Mortem Auctoris editit Carsten Niebuhr. Adjuncta est materia Medica Kahirina. Mölleri, Hafniae, 20 + xxxiv + 164 pp. https://doi.org/10.5962/bhl.title.2154

Fricke R, Eschmeyer WN, van der Laan R (Eds) (2024). Eschmeyer’s catalog of fishes: Genera, species, references. California Academy of Sciences, San Francisco, CA, USA. [Accessed on 22 August 2024] http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp

Gonzales BG (2013) Field guide to coastal fishes of Palawan. Coral Triangle Initiative on Coral Reefs. Fisheries, and Food Security (CTI-CFF), 107 pp.

Gmelin JF (1789) Caroli a Linné [...] Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species; cum characteribus, differentiis, synonymis, locis. Editio decimo tertia, aucta, reformata, vol 1, pars III. GE Beer, Lipsiae:1033–1516.

Griffiths AM, Sims DW, Cotterell SP, El Nagar A, Ellis JR, Lynghammar A, McHugh M, Neat FC, Pade NG, Queiroz N, Serra-Pereira B, Rapp T, Wearmouth VJ, Genner MJ (2010) Molecular markers reveal spatially segregated cryptic species in a critically endangered fish, the common skate (Dipturus batis). Proceedings of the Royal Society B: Biological Sciences 277(1687): 1497–1503. https://doi.org/10.1098/rspb.2009.2111

Hata E, Motomura H (2024) Neotrygon yakkoei, a new bluespotted maskray (Dasyatidae) from Japan. Ichthyological Research: 1–11. https://doi.org/10.1007/s10228-024-00989-7

Human BA (2022) Family Hemiscylliidae, bamboo sharks. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa: 454–457, pls. 18–20.

Human BA, Ebert DA (2022) Family Scyliorhinidae, catsharks. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa: 487–488, pls. 32.

IUCN (2023) International Union for Conservation of Nature. In: The IUCN Red List of Threatened Species. Version 2022–2. [Accessed 29 July 2023] https://www.iucnredlist.org

Ivanova NV, Zemlak TS, Hanner RH, Hebert PDN (2007) Universal primer cocktails for fish DNA barcoding. Molecular Ecology Notes 7(4): 544–548. https://doi.org/10.1111/j.1471-8286.2007.01748.x

Jordan DS, Snyder JO (1900) A list of fishes collected in Japan by Keinosuke Otaki, and by the United States steamer Albatross, with descriptions of fourteen new species. Proceedings of the United States National Museum 23(1213): 335–380. https://doi.org/10.5479/si.00963801.23-1213.335

Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16(2): 111–120. https://doi.org/10.1007/BF01731581

Koeda K, Manjaji-Matsumoto BM (2017) Family Pristiophoridae. In: Motomura H, Alama UB, Muto N, Babaran RP, Ishikawa S. (Eds) Commercial and bycatch market fishes of Panay Island, Republic of the Philippines. Kagoshima University Museum, Kagoshima, University of the Philippines Visayas, Iloilo, and Research Institute for Humanity and Nature, Kyoto, 23.

Kuhl H, van Hasselt JC (1823) Uittreksel uit een’ brief van Dr. J. C. van Hasselt, aan den Heer C. J. Temminck. Algemein Konst- en Letter-bode I Deel, No. 20, 315–317.

Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35(6): 1547–1549. https://doi.org/10.1093/molbev/msy096

Last PR, Compagno LJV (1999) Dasyatidae. In: Carpenter KE, Niem VH (Eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes, part 1 (Elopidae to Linophrynidae). FAO, Rome, 1479–1505.

Last PR, White WT, Pogonoski JJ (2008) Chimaera argiloba sp. nov., a new species of chimaerid (Chimaeriformes: Chimaeridae) from Northern Australia. In: Last PR, White WT, Pogonoski JJ (Eds) Descriptions of new Australian chondrichthyans. CSIRO Marine and Atmospheric Research Paper 22: 341–348.

Last PR, White WT, Caira JN Dharmadi, Fahmi , Jensen K, Lim APK, Manjaji-Matsumoto BM, Naylor GJP, Pogonoski JJ, Stevens JD, Yearsley GK (2010) Sharks and Rays of Borneo. Collingwood: CSIRO Publishing, Australia, 306 pp.

Last PR, Corrigan S, Naylor GP (2014) Rhinobatos whitei, a new shovelnose ray (Batoidea: Rhinobatidae) from the Philippine Archipelago. Zootaxa 3872(1): 31–47. https://doi.org/10.11646/zootaxa.3872.1.3

Last PR, White WT, Naylor G (2016a) Three new stingrays (Myliobatiformes: Dasyatidae) from the Indo–west Pacific. Zootaxa 4147(4): 377–402. https://doi.org/10.11646/zootaxa.4147.4.2

Last PR, White WT, Seret B (2016b) Taxonomic status of maskrays of the Neotrygon kuhlii species complex (Myliobatoidei: Dasyatidae) with the description of three new species from the Indo–West Pacific. Zootaxa 4083(4): 533–561. https://doi.org/10.11646/zootaxa.4083.4.5

Lee HT, Liao CH, Hsu TH (2024) DNA metabarcoding unveils the hidden species composition in fish surimi: Implications for the management of unlabeled and mixed seafood products. Heliyon 10(16): 1–10. https://doi.org/10.1016/j.heliyon.2024.e36287

Leeney RH, Bagarinao-Regalado A, Verdote D, Salgado CD (2024) A rapid assessment of the status of sawfishes in the Philippines. Endangered Species Research 53: 97–113. https://doi.org/10.3354/esr01295

Lesueur CA (1818) Descriptions of several new species of North American fishes. Journal of the Academy of Natural Sciences of Philadelphia 1: 359–368. https://doi.org/10.2307/1004927

Loh KH, Lim KC, Then AYH, Adam S, Leung AJX, Hu W, Bong CW, Wang A, Sade A, Musel J, Du J (2023) Advancing DNA barcoding to elucidate elasmobranch biodiversity in Malaysian waters. Animals 13(6): 1002. https://doi.org/10.3390/ani13061002

Manjaji-Matsumoto BM (2017) Family Rhinobatidae, Family Urolophidae, Family Dasyatidae. In: Motomura H, Alama UB, Muto N, Babaran RP, Ishikawa S (Eds) Commercial and bycatch market fishes of Panay Island, Republic of the Philippines. Kagoshima University Museum, Kagoshima, University of the Philippines Visayas, Iloilo, and Research Institute for Humanity and Nature, Kyoto, Japan, 24–25.

Manjaji-Matsumoto BM, Last PR (2008) Himantura leoparda sp. nov., a new whipray (Myliobatoidei: Dasyatidae) from the Indo-Pacific. CSIRO Marine and Atmospheric Research Paper No. 022: 293–301.

Manjaji-Matsumoto BM, de Carvalho MR, Santos HRS, Gomes UL, Last PR (2022) Family Dasyatidae. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa, 597, pl. 69.

McCulloch AR (1916) Report on some fishes obtained by the F. I. S. “Endeavour” on the coasts of Queensland, New South Wales, Victoria, Tasmania, South and South-Western Australia. Part IV. Biological Results Endeavour v. 4 (pt 4): 169–199, Pls. 49–58.

Motomura H, Ishikawa S (2013) Fish collection building and procedures manual: English edition. The Kagoshima University Museum, Kagoshima and the Research Institute for Humanity and Nature, Kyoto, 70 pp.

Müller J, Henle FGJ (1838–1841) Systematische Beschreibung der Plagiostomen. Berlin (Verlag von Veit and Comp), 200 pp. https://doi.org/10.5962/bhl.title.6906

Nardo GD (1827) Prodromus observationum et disquisitionum Adriaticae ichthyologiae. Giornale di fisica, chimica e storia naturale, medicina, ed arti 10(2): 22–40.

Naylor GJP, Caira JN, Jensen K, Rosana KAM, White WT, Last PR (2012) A DNA sequence-based approach to the identification of shark and ray species and its implications for global elasmobranch diversity and parasitology. Bulletin of the American Museum of Natural History 367: 1–262. https://doi.org/10.1206/754.1

Nelson JS, Grande TC, Wilson MV (2016) Fishes of the World. Fifth Edition. John Wiley and Sons, Hoboken, NJ, USA, 752 pp.

Okonechnikov K, Golosova O, Fursov M (2012) Unipro UGENE: A unified bioinformatics toolkit. Bioinformatics (Oxford, England) 28(8): 1166–1167. https://doi.org/10.1093/bioinformatics/bts091

Randall JE, Cea A (2011) Shore fishes of Easter Island. University of Hawai’i Press, 164 pp.

Richards VP, Henning M, Witzell W, Shivji MS (2009) Species delineation and evolutionary history of the globally distributed spotted eagle ray (Aetobatus narinari). Journal of Heredity 100(3): 273–283. https://doi.org/10.1093/jhered/esp005

Rigby CL, Derrick D, Dicken M, Harry AV, Pacoureau N, Simpfendorfer C (2021) Carcharhinus plumbeus. The IUCN Red List of Threatened Species 2021: e.T3853A2874370. https://doi.org/10.2305/IUCN.UK.2021-2.RLTS.T3853A2874370.en

Santos M, Torres Jr F, Tumilba VM, Sprong F (Eds) (2017) . Sharks and Rays “Pating” at “Pagi” Philippine Status Report and National Plan of Action 2017–2022. Bureau of Fisheries and Aquatic Resources – National Fisheries Research and Development Institute – Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) GmbH on behalf of the German Federal Ministry for the Environment, Nature Conservation, Building and Nuclear Safety, Quezon City, Philippines, 176 pp.

SEAFDEC (2022) The Southeast Asian state of fisheries and aquaculture 2022. Southeast Asian Fisheries Development Center, 1–283. http://hdl.handle.net/20.500.12066/6752

Serena F (2005) Field identification guide to the sharks and rays of the Mediterranean and Black Sea. FAO, Rome, 97 pp.

Simpfendorfer CA, Hueter RE, Bergman U, Connett SMH (2002) Results of a fishery-independent survey for pelagic sharks in the western North Atlantic, 1977–1994. Fisheries Research 55(1–3): 175–192. https://doi.org/10.1016/S0165-7836(01)00288-0

Soufi-Kechaou E, Amor KOB, Souissi JB, Amor MMB, Capapé C (2018) The capture of a large predatory shark, Carcharhinus plumbeus (Chondrichthyes: Carcharhinidae), off the Tunisian coast (Central Mediterranean). Annales, Series Historia Naturalis 28: 23–28.

Springer S (1950) A revision of North American sharks allied to the genus Carcharhinus. American Museum Novitates, No. 1451. https://doi.org/10.2307/1439109

Stevens JD, Bonfil R, Dulvy NK, Walker PA (2000) The effects of fishing on sharks, rays, and chimaeras (chondrichthyans), and the implications for marine ecosystem. ICES Journal of Marine Science 57(3): 476–494. https://doi.org/10.1006/jmsc.2000.0724

Valenciennes A (1839) Carcharias (Prionodon) In: Müller J, Henle J (Eds). Systematische Beschreibung der Plagiostomen. Berlin (Verlag von Veit and Comp). p. 45, pl. 16.

Waite ER (1898) New South Wales fisheries. Trawling operations off the coast of New South Wales, between the Manning River and Jervis Bay, carried on by H. M. C. S. “Thetis”. Scientific report on the fishes. Official report, Sydney, 62 pp.

Wallace JH (1967) The batoid fishes of the east coast of southern Africa. Part II: Manta, Eagle, Duckbill, Cownose, Butterfly and Sting rays. South African Association for Marine Biological Research. Oceanographic Research. Oceanographic Research Institute Investigational Report, No.16. Ward RD, Zemlak TS, Innes BH, Last PR, Hebert PDN (2005) DNA barcoding Australia’s fish species. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 360(1462): 1847–1857. https://doi.org/10.1098/rstb.2005.1716

Ward RD, Holmes BH, White WT, Last PR (2008) DNA barcoding Australasian chondrichthyans: Results and potential uses in conservation. Marine and Freshwater Research 59(1): 57–71. https://doi.org/10.1071/MF07148

Weigmann S (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology 88(3): 837–1037. https://doi.org/10.1111/jfb.12874

Weigmann S, Gon O, Leeney RH, Barrowclift E, Berggren P, Jiddawi N, Temple AJ (2020) Revision of the sixgill sawsharks, genus Pliotrema (Chondrichthyes, Pristiophoriformes), with descriptions of two new species and a redescription of P. warreni Regan. PLoS One 15(3): e0228791. https://doi.org/10.1371/journal.pone.0228791

White WT, Last PR, Naylor GJP, Jensen K, Caira JN (2010) Clarification of Aetobatus ocellatus (Kuhl, 1823) as a valid species, and a comparison with Aetobatus narinari (Euphrasen, 1790) (Rajiformes: Myliobatidae). In: Last PR, White WT, Pogonoski JJ (Eds) Descriptions of new sharks and rays from Borneo. CSIRO Marine and Atmospheric Research Paper, 032: 141–164.

White WT, Last PR, Naylor GJP (2016) Rhinobatos manai sp. nov., a new species of guitarfish (Rhinopristiformes: Rhinobatidae) from New Ireland, Papua New Guinea. Zootaxa 4175(6): 588–600. https://doi.org/10.11646/zootaxa.4175.6.6

Winton MV, Ebert DA (2022) Family Carcharhinidae, requiem sharks. In: Heemstra PC, Heemstra E, Ebert DA, Holleman W, Randall JE (Eds) Coastal fishes of the western Indian Ocean. Volume 1. South African Institute for Aquatic Biodiversity, Makhanda, South Africa: 510–538, pls. 40–48.

Supplementary material

Supplementary material 1

List of sequences used in the study

Roxanne Cabebe-Barnuevo, Dianne Frances Penuela, Emmanuel S. Delloro Jr., Ricardo P. Babaran, Hiroyuki Motomura, Maria Celia D. Malay

Data type: docx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Download file (26.25 kb)

﻿Abstract

Keywords

﻿Introduction

﻿Materials and methods

﻿Results

﻿Family Chimaeridae Rafinesque, 1815

﻿ Chimaera phantasma Jordan et Snyder, 1900

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Hemiscylliidae Gill, 1862

﻿ Chiloscyllium plagiosum (Bennett, 1830)

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿ Chiloscyllium punctatum Müller et Henle, 1838

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Scyliorhinidae Gill, 1862

﻿ Atelomycterus marmoratus (Bennett, 1830)

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Carcharhinidae Jordan et Evermann, 1896

﻿ Carcharhinus plumbeus (Nardo, 1827)

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿ Carcharhinus sorrah (Valenciennes, 1839)

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Pristiophoridae Bleeker, 1859

﻿ Pristiophorus lanae Ebert et Wilms, 2013

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Rhinobatidae Müller et Henle, 1837

﻿ Rhinobatos whitei Last, Corrigan et Naylor, 2014

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Plesiobatidae Nishida, 1990

﻿ Plesiobatis daviesi (Wallace, 1967)

Material examined.

Morphological description.

Distribution.

DNA barcode.

Conservation status.

Remarks.

﻿Family Urolophidae Müller et Henle, 1841

﻿ Urolophus sp.

Material examined.

DNA barcode.

Remarks.

Abstract

Introduction

Materials and methods

Results

Family Chimaeridae Rafinesque, 1815

Chimaera phantasma Jordan et Snyder, 1900

Family Hemiscylliidae Gill, 1862

Chiloscyllium plagiosum (Bennett, 1830)

Chiloscyllium punctatum Müller et Henle, 1838

Family Scyliorhinidae Gill, 1862

Atelomycterus marmoratus (Bennett, 1830)

Family Carcharhinidae Jordan et Evermann, 1896

Carcharhinus plumbeus (Nardo, 1827)

Carcharhinus sorrah (Valenciennes, 1839)

Family Pristiophoridae Bleeker, 1859

Pristiophorus lanae Ebert et Wilms, 2013

Family Rhinobatidae Müller et Henle, 1837

Rhinobatos whitei Last, Corrigan et Naylor, 2014

Family Plesiobatidae Nishida, 1990

Plesiobatis daviesi (Wallace, 1967)

Family Urolophidae Müller et Henle, 1841

Urolophus sp.

Family Dasyatidae Jordan, 1888

Himantura sp.

Genus Neotrygon Castelnau, 1873

Neotrygon orientalis Last, White et Serét, 2016

Genus Taeniura Müller et Henle, 1837

Taeniura lymma (Forsskål, 1775)

Family Aetobatidae Agassiz, 1858

Aetobatus ocellatus (Kuhl in Kuhl and van Hasselt, 1823)

Discussion

Conclusions

Author contributions

Acknowledgments

References