Description of a Remarkable New Skate Species of Leucoraja Malm, 1877 (Rajiformes, Rajidae) from the Southwestern Indian Ocean: Introducing 3D Modeling as an Innovative Tool for the Visualization of Clasper Characters.

A remarkable new deep-water skate, Leucoraja longirostris n. sp., is described based on eight specimens caught during different expeditions to the southern Madagascar Ridge in the southwestern Indian Ocean. The new species differs from all congeners by its remarkably long and acutely angled snout (horizontal preorbital length 17.2-22.6% TL vs. 8.5-11.9% TL and 4.2-6.1 vs. 1.7-3.5 times orbit length, snout angle 65-85° vs. 90-150°). Furthermore, it is apparently endemic to the Madagascar Ridge, distant from the known distribution areas of all congeners. In addition to L. fullonica and L. pristispina, L. longirostris n. sp. is also the only species with plain dorsal coloration. Furthermore, the new species is the only Leucoraja species with an external clasper component dike and, besides L. wallacei, the only one with four dorsal terminal (dt) cartilages. The shape of the accessory terminal 1 (at1) cartilage with four tips is also unique within the genus. A new approach for the visualization of the clasper characters is introduced based on 3D models of all skeletal and external features. This enables a much easier and much more precise interpretation of every single clasper component, of the entire structure, and, in particular, the relationship between external features and skeletal cartilages. A new English translation of the first diagnosis of Leucoraja is provided, along with a revised generic diagnosis and a key to the species of Leucoraja in the Indian Ocean.

Revision of the Western Indian Ocean Angel Sharks, Genus Squatina (Squatiniformes, Squatinidae), with Description of a New Species and Redescription of the African Angel Shark Squatina africana Regan, 1908.

Sampling efforts on the Saya de Malha Bank (part of the Mascarene Plateau, western Indian Ocean) unveiled three unusual small juvenile angel shark specimens, that were a much paler color than the only known western Indian Ocean species, Squatina africana Regan, 1908. However, it took many years before further specimens, including adults of both sexes, and tissue samples were collected. The present manuscript contains a redescription of S. africana based on the holotype and additional material, as well as the formal description of the new species of Squatina. All specimens of the new species, hereafter referred to as Squatina leae sp. nov., were collected in the western Indian Ocean off southwestern India and on the Mascarene Plateau at depths of 100-500 m. The new species differs from S. africana in a number of characteristics including its coloration when fresh, smaller size at birth, size at maturity, and adult size, genetic composition, and distribution. Taxonomic characteristics include differences in the morphology of the pectoral skeleton and posterior nasal flap, denticle arrangement and morphology, vertebral counts, trunk width, pectoral-pelvic space, and clasper size. A key to the species of Squatina in the Indian Ocean is provided.

Morphological and Molecular Evidence Reveals the Longnose Skate Zearaja brevicaudata (Marini, 1933) to be a Senior Synonym of Dipturus lamillai Concha, Caira, Ebert & Pompert 2019.

Longnose skates have great economic importance in South American fisheries, and in order to preserve them, it is important to have a well-defined taxonomic status of their species. Dipturus lamillai was recently described for Malvinas Islands waters based on morphological and molecular comparisons with Zearaja chilensis. Although D. lamillai has been compared with several congeneric species, it was not properly compared with the morphologically similar Zearaja brevicaudata, the most abundant longnose skate in the Southwest Atlantic. Here, these species were compared by morphological and molecular analyses in order to evaluate their conspecificity. Linear morphometric variables of holotype and paratypes of D. lamillai and 69 specimens of Z. brevicaudata were compared and investigated using Principal Component Analysis. In addition, thorn patterns, denticle distributions, color, and clasper morphology were compared. No body proportions or other single character that could differentiate D. lamillai from Z. brevicaudata were found. Molecular analyses comprised of the comparison of the Cytochrome oxidase subunit I (COI) and the NADH dehydrogenase subunit 2. The results of the Maximum Likelihood (ML) carried out for each molecular marker showed that sequences from D. lamillai clustered together with those of Z. brevicaudata, and the molecular distance determined by Kimura two-parameter were lower than the expected for different species. Additionally, the Automatic Barcode Gap Discovery method and the Bayesian implementation of the Poisson tree processes were carried out with COI sequences to explore species limits, and their results were consistent with ML analyses. In summary, the results obtained showed that there are no morphological or molecular differences between these nominal species of the valid skate genus Zearaja, leading to the conclusion that they are conspecific. Therefore, we designated Z. brevicaudata as a senior synonym of D. lamillai.

Notes on batoid fishes of the Socotra Archipelago (north-western Indian Ocean), with four new records.

The species composition of batoid fishes from coastal waters of the Socotra Archipelago is reviewed, with confirmed records of the wedgefish Rhynchobatus djiddensis (Forsskål, 1775) and four new records of sharkrays, wedgefishes, and guitarfishes based on collected specimens, including one species from Abd al-Kuri Island, Rhina ancylostoma Bloch Schneider, 1801 (Rhinidae), and three species from the main island Socotra, Acroteriobatus salalah (Randall Compagno, 1995) and Rhinobatos punctifer Compagno Randall, 1987 (Rhinobatidae), and Rhynchobatus australiae Whitley, 1939 (Rhinidae). Among the new records for the Socotra Archipelago, R. australiae represents the first verified record for the Arabian region. In addition, records of four stingray species (Dasyatidae) are verified based on underwater observations accompanied with photographs. All recorded batoid fishes are commercial species caught in the local small-scale fishery. Information on the identification and distribution of each species is provided.

Revision of the sixgill sawsharks, genus Pliotrema (Chondrichthyes, Pristiophoriformes), with descriptions of two new species and a redescription of P. warreni Regan.

Recent sampling efforts in Madagascar and Zanzibar, as well as examinations of six-gilled sawsharks in several museum collections provided evidence for a complex of species within Pliotrema warreni Regan. The present manuscript contains a redescription of P. warreni involving the syntypes and additional material, as well as formal descriptions of two new species of Pliotrema Regan. All specimens of both new species were found in the western Indian Ocean. Individuals of the first new species, hereafter referred to as P. kajae sp. nov., were identified originating from Madagascar and the Mascarene Ridge. Specimens of the second new species, hereafter referred to as P. annae sp. nov., were only found off Zanzibar. Pliotrema kajae sp. nov. appears to inhabit upper insular slopes and submarine ridges at depths of 214-320 m, P. annae sp. nov. so far is only known from shallow waters (20-35 m). Both new species differ from P. warreni in a number of characteristics including the known distribution range and fresh coloration. Taxonomical differences include barbels that are situated approximately half way from rostral tip to mouth, with prebarbel length equidistant from barbel origin to symphysis of the upper jaw in P. kajae sp. nov. and P. annae sp. nov. (vs. about two thirds way from rostral tip to mouth, with prebarbel length about twice the distance from barbel origin to symphysis of upper jaw in P. warreni) and rostra that are clearly and slightly constricted between barbel origin and nostrils, respectively (vs. rostrum not constricted). Pliotrema kajae sp. nov. differs from P. annae sp. nov. in a longer snout, more numerous large lateral rostral teeth and upper jaw tooth rows, jaw teeth with (vs. without) sharp basal folds, and coloration, particularly pale to light brown (vs. medium to dark brown) dorsal coloration with (vs. without) two indistinct yellowish stripes. A revised diagnosis of Pliotrema and a key to the species are provided.

New records and range extension of the Portuguese dogfish Centroscymnus coelolepis in the south-western Atlantic Ocean, with comments on its morphology.

The Portuguese dogfish Centroscymnus coelolepis is a wide-ranging deep-water shark and a common by-catch component of the catches of several mid- to deep-water fisheries. In the present study, two new records from the south-western Atlantic Ocean are reported based on specimens caught by bottom-longline fishing vessels operating in the Argentinean-Uruguayan Common Fishing Zone. Species identification based on morphology and detailed morphometrics, as well as molecular data are presented for one of the specimens. The distribution of the species over the south-western Atlantic is discussed on the basis of the available bibliography and a thorough revision of museum collections. The records presented here expand the species' previously acknowledged distribution southwards, from around 21° S to at least 38° S, suggesting it occurs continuously along the shelf break of eastern South America. However, given the limited access to specimens of deep-water sharks in the region, the abundance and real extent of C. coelolepis distribution in the south-western Atlantic as well as its interaction with deep-water fisheries remain to be fully assessed.

A new genus and species of catshark (Carcharhiniformes: Scyliorhinidae) from eastern Indonesia.

A new genus and species of catshark is described based on a single specimen collected off Ambon in the Maluku Islands of eastern Indonesia. Akheilos suwartanai belongs to the subfamily Schroederichthyinae which differs from the other catsharks in a combination of: similar sized dorsal fins, supraorbital crests present, pseudosiphon present on claspers, broad subocular ridges under eyes, posterior nasal flaps present, tips of rostral cartilage fused into a rostral node. It represents the first record of this subfamily outside of the Americas. Akheilos differs from the other genus in the subfamily, Schroederichthys in a combination of: clasper groove not fused dorsally, ventral lobe of caudal fin produced, more intestinal valve turns, anal fin slightly larger than second dorsal fin, and in colour pattern.

Updated taxonomic list and conservation status of chondrichthyans from the exclusive economic zone of Venezuela, with first generic and specific records.

Using the last taxonomic review of chondrichthyans of the world, we selected the species distributed in the north-western Atlantic Ocean (NWA) and compared it with the available published literature related to the class Chondrichthyes in the Venezuelan exclusive economic zone. We also revised information from worldwide databases such as: FAO (NWA-31 area), GBIF, iSpecies, IUCN and OBIS, as well as available museum collection databases. The taxonomic validity was checked using the Catalogue of Fishes of the California Academy of Sciences and recent references. The past published Venezuelan lists of chondrichthyans combined included nine orders, 30 families and 108 species. The updated list with 12 orders, 36 families and 122 species increased by three new orders, six families, three shark genera, nine shark species (one replacing another species), one chimaera genus, two chimaera species, three batoid genera and six batoid species (two replacing other species). Four holotype specimens (two sharks and two rays) are deposited in Venezuelan Museums. Most of the species have an IUCN conservation status, including four species catalogued as Critically Endangered, six as Endangered and 18 species as Vulnerable. Deep-sea fisheries, scientific exploration and taxonomic/genetic revisions might add future increments to the Venezuelan chondrichthyan list.

A new microendemic species of the deep-water catshark genus Bythaelurus (Carcharhiniformes, Pentanchidae) from the northwestern Indian Ocean, with investigations of its feeding ecology, generic review and identification key.

A new deep-water catshark, Bythaelurus stewarti, is described based on 121 examined specimens caught on the Error Seamount (Mount Error Guyot) in the northwestern Indian Ocean. The new species differs from all congeners in the restricted distribution, a higher spiral valve turn count and in the morphology of the dermal denticles. It is distinguished from its morphologically and geographically closest congener, B. hispidus (Alcock), by the larger size (maximum size 44 vs. 39 cm TL, maturity size of males 35-39 vs. 21-28 cm TL), darker fresh coloration and dark grayish-brown mottling of the ventral head (vs. ventral head typically uniformly yellowish or whitish). Furthermore, it has a strongly different morphology of dermal denticles, in particular smaller and less elongate branchial, trunk and lateral caudal denticles that are set much less densely and have a surface that is very strongly and fully structured by reticulations (vs. structured by reticulations only in basal fourth). In addition, the new species differs from B. hispidus in having more slender claspers that are gradually narrowing to the bluntly pointed tip without knob-like apex (vs. claspers broader and with distinct knob-like apex), more spiral valve turns (11-12 vs. 8-10) and numerous statistical differences in morphometrics. A review of and a key to the species of Bythaelurus are given.

Bythaelurus vivaldii, a new deep-water catshark (Carcharhiniformes, Scyliorhinidae) from the northwestern Indian Ocean off Somalia.

A new very small deep-water catshark, Bythaelurus vivaldii, is described based on two female specimens caught off Somalia in the northwestern Indian Ocean during the German 'Valdivia' expedition in 1899. It is morphologically closest to the recently described B. bachi, which is the only other Bythaelurus species in the western Indian Ocean that shares a stout body of large specimens and the presence of oral papillae. It further resembles B. vivaldii in the broad mouth and broad posterior head, but differs in the presence of composite oral papillae and a higher diversity in dermal denticle morphology. Additionally, the new species differs from all congeners in the western Indian Ocean in a larger pre-second dorsal fin length, a longer head, a larger interdorsal space, a larger intergill length, a longer pectoral-fin posterior margin, a shorter caudal fin, an intermediate caudal fin preventral margin, and a larger internarial width. Furthermore, the second dorsal fin of the new species is smaller than in its congeners in the western Indian Ocean except for B. lutarius, which is easily distinguished by the slender body and virtual absence of oral papillae, as well as the aforementioned further characters. An updated key to all valid species of Bythaelurus is provided.

A new deepwater legskate, Sinobatis kotlyari n. sp. (Rajiformes, Anacanthobatidae) from the southeastern Indian Ocean on Broken Ridge.

Sinobatis kotlyari n. sp. is described, based on the nearly adult 331 mm TL holotype male from 1400 m depth in the southeastern Indian Ocean at Broken Ridge. The new species is assigned to Sinobatis due to mainly characters of its clasper and cranium fontanelle distinguishing it from congeners and other anacanthobatid skates with clasper features known. It further differs from its morphologically closest congener, S. borneensis, which is only known from the northwestern Pacific Ocean, e.g. in larger maturing size, longer snout and head, the absence of a snout filament and skin fold on tail, a longer caudal fin, and in colouration. The two congeners known from the eastern Indian Ocean, S. bulbicauda and S. cae-rulea, mature at much larger size, grow to larger maximum size and differ in numerous morphometric and meristic           characters, as well as in colouration. The type locality appears to be very remote as compared with other Indian Ocean legskates, which have primarily been found on continental and insular slopes.

Bythaelurus bachi n. sp., a new deep-water catshark (Carcharhiniformes, Scyliorhinidae) from the southwestern Indian Ocean, with a review of Bythaelurus species and a key to their identification.

A new deep-water catshark, Bythaelurus bachi, is described based on 44 specimens caught on the southern Madagascar Ridge in the southwestern Indian Ocean. The new species is the only stout-bodied Bythaelurus with oral papillae in the region and is distinguished from all congeners by the plain beige to light gray-brown coloration, high diversity in dermal denticle morphology, and presence of composite oral papillae. Despite resemblance in body shape, Bythaelurus bachi n. sp. is distinguished from its closest congener, B. naylori Ebert & Clerkin, 2015, by the presence of numerous large, partially composite papillae on the tongue and roof of the mouth (vs. papillae lacking), plain light coloration (vs. medium to dark brown ground color, light fin edges and a distinctly dark dusky-colored snout), only slightly enlarged dermal denticles on the anterior upper caudal-fin margin (vs. dermal denticles distinctly enlarged), a higher diversity in dermal denticle morphology in general, and smaller maximum size and size at maturity. The distinction of both species is also supported by molecular results. The new species differs from all other congeners in the western Indian Ocean in the stout body shape of large specimens, coloration, larger size, as well as several morphometrics, including larger claspers, longer eyes and dorsal fins, and shorter pelvic-anal and pelvic-caudal spaces. The genus is reviewed, a key to its species given.

A new skate genus Orbiraja (Rajiformes: Rajidae) from the Indo-West Pacific.

Molecular analyses and information gleaned from an examination of the newly available adult male of the North-West Pacific skate, Okamejei jensenae Last & Lim, supported earlier concerns that the species might be mis-assigned. Morphological data based on this specimen supported its placement in a new genus Orbiraja that is assigned to the recently named Rostrorajini based on molecular evidence. This subgroup of the family Rajidae also includes Malacoraja, Neoraja, Rostroraja and an unresolved 'amphi-American Assemblage' (sensu McEachran & Dunn, 1998). Orbiraja is unique within the rajids in having the combination of three, very closely spaced median thorn rows on the tail, no dark-edged ventral pores, and a clasper skeleton with a prominent accessory terminal 3 cartilage formed by a medio-distal extension of the accessory terminal 2 cartilage. Its spiracle appears to be situated posteriorly with respect to the orbit. The group contains two other nominal species, Orbiraja powelli (Alcock) and O. philipi (Lloyd), and an un-named species from Indonesia that needs further investigation. Orbiraja jensenae is rediagnosed based on characteristics of the adult male.

Sinobatis brevicauda n. sp., a new deep-water legskate (Rajiformes, Anacanthobatidae) and first generic record from the western Indian Ocean.

A new deep-water legskate, Sinobatis brevicauda, is described based on two specimens caught on the remote Saya de Malha Bank in the central western Indian Ocean. The new species is easily distinguished from all other described anacanthobatids by the short tail. It is the only Sinobatis species described from the western Indian Ocean and differs from the other anacanthobatid legskates in this area by its large size and light coloration. All other species of Sinobatis are described from the eastern Indian and, particularly, western Pacific oceans. In addition to the short tail, the new species clearly differs from its morphologically closest congener, the Australian S. bulbicauda, in a bicolored white and grayish ventral coloration with gray and white blotches (vs. uniformly pale or white and skin somewhat translucent) and a filamentous tail without flattened, bulbous tip. Furthermore, it is distinguished by several morphometric and meristic differences, e.g. a longer body (length 65% TL vs. 39-61% TL), longer head (dorsal length 34% TL vs. 21-31% TL, ventral length 41% TL vs. 23-36% TL), longer snout (preorbital length 28-29% TL vs. 14-26% TL, preoral length 30% TL vs. 16-28% TL, prenasal length 28% TL vs. 14-25% TL), and fewer diplospondylous (102-112 vs. 121-142) and total (131-141 vs. 148-168) vertebrae. S. brevicauda clearly differs from the other anacanthobatids in the western Indian Ocean, Anacanthobatis marmorata and Indobatis ori, by having a much shorter tail, strongly different coloration, much larger size, and in many morphometric and meristic differences.

Bythaelurus tenuicephalus n. sp., a new deep-water catshark (Carcharhiniformes, Scyliorhinidae) from the western Indian Ocean.

A new dwarf deep-water catshark, Bythaelurus tenuicephalus, is described based on one adult and one juvenile male specimen from off Tanzania and Mozambique in the western Indian Ocean. The new species differs from its congeners by its slender head and snout, which is only slightly bell-shaped in dorsoventral view without distinct lateral indention. All other Bythaelurus species have distinctly bell-shaped snouts with a strong lateral indention anterior to outer nostrils. Compared to its congeners in the western Indian Ocean, B. tenuicephalus n. sp. also has broader claspers in adult males (base width 2.1% TL vs. 1.5-1.8% TL). It further differs from B. clevai by attaining a smaller maximum size and having a color pattern of fewer and smaller blotches, larger oral papillae, a shorter snout, and broader claspers without knob-like apex and with a smaller envelope and a subtriangular (vs. subrectangular) exorhipidion. Compared to B. hispidus, the new species has a longer snout, a longer dorsal-caudal space, broader clasper without knob-like apex, and fewer vertebral centra. In contrast to B. lutarius, B. tenuicephalus attains a smaller maximum size and has a blotched (vs. largely plain) coloration, numerous (vs. lacking) oral papillae, shorter anterior nasal flaps, a longer caudal fin, a shorter pelvic anal space, and shorter and broader claspers.

Apristurus breviventralis, a new species of deep-water catshark (Chondrichthyes: Carcharhiniformes: Scyliorhinidae) from the Gulf of Aden.

A new deep-water catshark of the genus Apristurus Garman, 1913 is described based on nine specimens from the Gulf of Aden in the northwestern Indian Ocean. Apristurus breviventralis sp. nov. belongs to the 'brunneus group' of the genus and is characterized by having pectoral-fin tips reaching beyond the midpoint between the paired fin bases, a much shorter pectoral-pelvic space than the anal-fin base, a low and long-based anal fin, and a first dorsal fin located behind pelvic-fin insertion. The new species most closely resembles the western Atlantic species Apristurus canutus, but is distinguishable in having greater nostril length than internarial width and longer claspers in adult males. Apristurus breviventralis sp. nov. represents the sixth species of Apristurus from the western Indian Ocean and the 38th species globally. 

Rajella paucispinosa n. sp., a new deep-water skate (Elasmobranchii, Rajidae) from the western Indian Ocean off South Mozambique, and a revised generic diagnosis.

A new species of the widely in temperate and tropical latitudes distributed skate genus Rajella is described based on an almost adult male specimen from the western Indian Ocean off South Mozambique. The holotype of R. paucispinosa n. sp. was caught during cruise 17 of RV 'Vityaz' along the deep western Indian Ocean in 1988/89. It is the northernmost record of a Rajella specimen in the western Indian Ocean. The new species is the 18th valid species of the genus and the fifth species in the western Indian Ocean. It differs from its congeners in the small maximal total length of about 50 cm and only few thorns on the dorsal surface. The new species has only two thorns on each orbit, one nuchal thorn, one right scapular thorn (left one not detectable, abraded), and one median row of tail thorns. Other species of Rajella typically have half rings of thorns on orbital rims, a triangle of thorns on nape-shoulder region, and at least three rows of tail thorns. Another conspicuous feature of the new species is the almost completely white dorsal and ventral coloration. 

Carcharhinus humani sp. nov., a new whaler shark (Carcharhiniformes:Carcharhinidae) from the western Indian Ocean.

A new species of whaler shark, Carcharhinus humani sp. nov., is described based on five type specimens from the western Indian Ocean near the Socotra Islands, off Kuwait, Mozambique, and South Africa. The new species represents the fifth species of the C. dussumieri/sealei group and the third species of the C. sealei subgroup. The new species is the only species of the C. sealei subgroup known from the western Indian Ocean. Within the C. sealei subgroup, C. humani differs from C. sealei in having a sharply demarcated black apical marking on the second dorsal fin which does not extend onto body surface (vs. black marking diffuse-edged and usually extending onto upper sides of trunk), a longer horizontal prenarial length (4.1-4.7 vs. 3.4-4.2% TL), and a longer preoral length (6.8-7.6 vs. 5.7-6.5% TL); C. humani differs from C. coatesi in having a taller second dorsal fin (its height 4.0-4.5 vs. 2.9-3.6% TL), a shorter first dorsal fin (its length 13.4-14.6 vs. 14.8-17.3% TL), and more vertebrae (total centra 152-167 vs. 134-147).

Complementary redescription of Anacanthobatis ori (Wallace, 1967) and its assignment to Indobatis n. g. (Elasmobranchii, Anacanthobatidae), with comments on other legskates.

Anacanthobatis ori is one of the least known species of the family Anacanthobatidae with only four juvenile specimens reported. The species remained assigned to the genus Anacanthobatis sensu lato due to the lack of an adult male as external and skeletal clasper characters are the essential diagnostic features for the differentiation of genera and subgenera within the family Anacanthobatidae. Since an adult male of A. ori became available, along with an adult female and six further juveniles, the authors reinvestigated the species and present its so far unknown diagnostic characters of clasper morphology and skeleton and scapulocoracoid. The clasper turned out to be the most complex one of all known anacanthobatids as the external components flag, slit, pseudosiphon-like cavity, pecten, and two sentinas are not known from any other anacanthobatid species. Furthermore, a dorsal terminal 1 cartilage is present but displaced proximally of the terminal clasper skeleton, the outer edge of dorsal terminal 2 is deeply serrated, the ventral terminal has a very long, curved, strap-like process, and the proximal part of accessory terminal 1 is embedded in the cavity of the baseball-glove-like head of accessory terminal 2. Due to the strong differences in external and internal clasper characters to all other known anacanthobatid species, A. ori is placed in its own, newly erected genus, Indobatis.