On the Taxonomy of the First Record of Rare Deep-Water Rough Shark Species of Oxynotidae (Chondrichthyes: Squaliformes) in the Western Indian Ocean

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Journal of Threatened Taxa Building evidence for conservaton globally www.threatenedtaxa.org ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) Communication On the taxonomy of the first record of rare deep-water rough shark species of Oxynotidae (Chondrichthyes: Squaliformes) in the western Indian Ocean

Sarah Viana & Mark W. Lisher

26 May 2018 | Vol. 10 | No. 6 | Pages: 11732–11742 10.11609/jot.3916.10.6.11732-11742

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Threatened Taxa Taxonomy of a rough shark from the WesternJournal Indian Ocean of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742Viana & Lisher

On the taxonomy of the first record of rare

Communication deep-water rough shark species of Oxynotidae

ISSN 0974-7907 (Online) (Chondrichthyes: Squaliformes) in the western Indian Ocean ISSN 0974-7893 (Print) Sarah Viana 1 & Mark W. Lisher 2

OPEN ACCESS 1 South African Insttute for Aquatc Biodiversity, Private Bag 1015, Grahamstown, 6140, South Africa 2 The Scotsh Associaton for Marine Science, Scotsh Marine Insttute, Dunbeg, Oban, Argyll, Scotland, PA37 1QA, United Kingdom 1 [email protected] (corresponding author), 2 [email protected]

Abstract: An immature female specimen of rough shark was collected south of Reunion Island in the Madagascar Basin in 2009 aboard R/V Dr. Fridtjof Nansen, representng the frst ofcial record of the family Oxynotdae in the western Indian Ocean. The specimen is herein identfed as Oxynotus sp. due to morphological diferences with its closely similar congeners O. centrina and O. bruniensis regarding morphometrics, shape of dorsal, pectoral and caudal fns, shape of the head and colouraton, refutng the hypothesis of occurrence of these two species in the region. These results indicate that Oxynotus sp. is possibly an undescribed species. A general descripton of the external morphology, external morphometrics and photographs of Oxynotus sp. are provided. The specimen has a hepatosomatc index of 36.33% which reveals that it was possibly approaching maturaton, suggestng that a viable populaton of Oxynotus sp. exists in the western Indian Ocean. Intraspecifc variatons in O. centrina from the Mediterranean Sea and eastern Atlantc Ocean are also notced, requiring further taxonomic scrutny.

Keywords: Madagascar Basin, morphology, Oxynotus, rough sharks, taxonomy, western Indian Ocean.

DOI: htp://doi.org/10.11609/jot.3916.10.6.11732-11742 | ZooBank: urn:lsid:zoobank.org:pub:FFDA138B-7CB7-4A5B-ACE5-D8E46772CAE3

Editor: E. Vivekanandan, Central Marine Fisheries Research Insttute, Chennai, India. Date of publicaton: 26 May 2018 (online & print)

Manuscript details: Ms # 3916 | Received 26 November 2017 | Final received 03 April 2018 | Finally accepted 22 April 2018

Citaton: Viana, S. & M.W. Lisher (2018). On the taxonomy of the frst record of rare deep-water rough shark species of Oxynotdae (Chondrichthyes: Squaliformes) in the western Indian Ocean. Journal of Threatened Taxa 10(6): 11732–11742; htp://doi.org/10.11609/jot.3916.10.6.11732-11742

Copyright: © Viana & Lisher 2018. Creatve Commons Atributon 4.0 Internatonal License. JoTT allows unrestricted use of this artcle in any medium, reproducton and distributon by providing adequate credit to the authors and the source of publicaton.

Funding: None.

Competng interests: The authors declare no competng interests.

Author Details: Sarah Viana is a research associate at the South African Insttute for Aquatc Biodiversity with expertse in taxonomy, phylogeny and biogeography of Chondrichthyes, especially Squaliformes. Her currently interests are conservaton genetcs and molecular phylogeography of southern African endemic species. Mark W. Lisher worked for several years in the feld of collecton management and biodiversity data in South Africa. Currently, he is a MSc scholar at The Scotsh Associaton for Marine Science. His interests are ichthyology, biodiversity, and taxonomy of marine fshes.

Author Contributon: SV and MWL contributed equally in the conceptualizaton, collecton and analysis of data, discussion of the results and writng up of this manuscript.

Acknowledgements: The authors thank O. Gon, A. Whitield, R. Bills, B. Konqobe and N. Mazungula (SAIAB), and E. Mitchell (SAMS) for curatorial and technical assistance. Special thanks to A. Hay and M. McGrouther (AMS) for providing informaton on comparatve material, and A. Orlov (VNIRO) for helping with acquisiton of original descriptons. The frst author was supported by the South African Insttute for Aquatc Biodiversity.

11732 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

INTRODUCTION congeners by having grey or brownish grey body with broad dark brown blotches dorsal-laterally, eyes with Oxynotdae is a monotypic family of small to medium heavy supra-ocular ridge and knobs, spiracles large sized rare deep-water rough sharks with endemic regional and vertcal (Compagno 2016), and few proportonal geographical distributons on contnental and insular external measurements (e.g., dorsal fn-spines and shelves and upper contnental slopes of the Atlantc frst dorsal-fn apex, precaudal length and origin of frst and Pacifc Oceans (Ebert et al. 2013). The occurrence dorsal fn spine) (Yano & Matsuura 2002). It inhabits of Oxynotdae and records of its representatves in the muddy and coralline algal botoms of contnental and western Indian Ocean were speculatve (e.g., Ebert insular shelves and upper contnental slopes between 2013, 2015; Ebert & van Hees 2015; Compagno 2016; 40–800 m depth, and is occasionally caught as bycatch Weigmann 2016) tll recently in Fricke et al. (2018). during trawls and longlining (Ebert & Stehmann 2013; Members of Oxynotdae are characterized by having a Ragonese et al. 2013; Compagno 2016). This species body triangular in cross-secton, conspicuously deep, occurs throughout the Eastern Atlantc Ocean from arched dorsally and fatened ventrally at trunk, rough Norway to the western coast of South Africa (Bass et skin with large dermal dentcles, head small and wide, al. 1976; Cadenat & Blache 1981; Ebert & Stehmann snout blunt at tp and short with fringed anterior 2013; Ebert 2015; Compagno 2016). According to Ebert margin of nostrils, mouth circular with papillose lips and & Stehmann (2013), Compagno et al. (2015), Ebert ventrally located, eyes rounded and conspicuously large, (2015) and Compagno (2016), O. centrina or a separate located laterally, spiracles large and crescent, located similar species possibly occurs in Mozambique, or of behind eyes, dorsal fns sail-like, conspicuously upright Madagascar (Ebert & van Hees 2015). Ebert (2013) and and tall, dorsal-fn spines conical, thick and markedly Weigmann (2016) later listed O. centrina in the western small, located prior to each dorsal fn and never Indian Ocean although without scrutny, indicatng that transcending the fn apex, pectoral fns leaf-like, pointed the identfcaton of this species or any other member of distally at apex, and inconspicuous free-rear tps, lateral Oxynotdae in the region is uncertain. Fricke et al. (2018) ridges prominent at trunk, caudal fn small and broadly inaccurately listed this species as from of northern sub-triangular with inconspicuous lower caudal lobe and Madagascar, representng the frst ofcial record of evident subterminal notch. It bears teeth dissimilar in Oxynotdae in the region, but again without detailed both jaws, upper teeth smaller than lower teeth, upper taxonomic informaton. The present study thus aimed teeth thin and lanceolate, lower teeth broad and blade- to provide a general morphological descripton, external like (Compagno 1999). morphometrics, details of the collecton and biological Oxynotus Rafnesque, 1810 currently comprises data of this specimen, and to clarify the occurrence of fve valid species: O. centrina (Linnaeus, 1758), type- Oxynotdae in the western Indian Ocean. Taxonomic species, originally from the Mediterranean Sea; O. problems on species of Oxynotus are also discussed paradoxus Frade, 1929 from the northeastern Atlantc herein. Ocean, ranging from Morocco to Senegal; O. caribbeaus Cervigon, 1961 from the Caribbean; O. bruniensis (Ogilby, 1893) from the southwestern Pacifc Ocean, MATERIAL AND METHODS partcularly Australia and New Zealand; O. japonicus Yano & Murofshi, 1985 from Japanese and Taiwanese The southern Indian Ocean seamounts cruise coasts (Ho & Nakaya 2016). Litle is known about their expediton was undertaken aboard the R/V Dr. Fridtjof biological and ecological traits, including populaton Nansen along the southwestern Indian Ridge between size, distributon and life history, and studies regularly 12 November and 19 December 2009. During this cruise, focus on reproductve biology and feeding of O. centrina a single specimen of Oxynotus sp. was captured on 14 (e.g., Capapé et al. 1999; Capapé 2008; Guallart et al. November 2009 during Trawl 1, Staton 2, Madagascar 2015; Kousteni & Megalofonou 2016). Currently, these Basin, south of Reunion Island, Event 7, at a maximum species are listed as Data Defcient in the IUCN Red List depth of 600m using a pelagic Ǻkra trawl net fted with of Threatened Species (Francis 2003; Leandro 2004; a multsampler. The Ǻkra trawl net used was a Flytetrål Yano 2004; Soldo & Freitas 2009), with the excepton of 152 MSK x 3200mm, with a 20m net mouth opening O. centrina, which is considered Vulnerable (Bradaï et al. (Rogers et al. 2010). Prior to each trawl, ice-trays and 2007). labels were prepared for sortng. Once on deck, samples Oxynotus centrina is usually distnguished from its were empted into large trays containing ice. A small

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 11733 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher amount of seawater was added to the trays to prevent Skin sample measuring 1cm2 was taken from below frst the samples freezing to the ice. Specimen of Oxynotus dorsal fn (right side) for analysis of dermal dentcles. sp. was labelled by tying a label to the fsh using string, Liver weight and conditons follow Capapé et al. (1999). then either fxed in a 4% formalin soluton or frozen Hepatosomatc index (HIS) was calculated using the (Rogers et al. 2010). This specimen was later donated formula HSI (%) = liver weight (g) / dressed weight (g) and logged at the South African Insttute for Aquatc X 100. Specimen was photographed in dorsal, lateral, Biodiversity (SAIAB) where further curatorial procedures and ventral views as well as individual body parts (e.g., such as identfcaton and cataloguing took place, and fns, liver) using digital camera. Photographs of dermal put through a series of 10% and 50% ethanol solutons, dentcles were taken using stereoscope microscope before fnally being preserved in 70% ethanol soluton. atached to digital camera at the Scotsh Associaton for External measurements were obtained from a Marine Science. Map of collectng locality of examined specimen preserved in 70% ethanol using a digital calliper specimen was generated using QGIS 2.14.2 Essen (QGIS and metric tape for measurements greater than 150mm. Development Team, QGIS Geographic Informaton Measurements follow Last et al. (2007) and Ebert et al. System, Open Source Geospatal Foundaton Project; (2013), and are expressed as percentage of total length htp://qgis.osgeo.org/) and Google Earth (htp:// (% TL). Morphometric data of O. centrina recorded www.google.co.uk/intl/en_uk/earth). Insttutonal elsewhere were compiled from Yano & Murofushi abbreviatons are in accordance to Sabaj (2016). (1985), Barrull & Mate (2001), Yano & Matsuura (2002), Megalofonou & Damalas (2004), Dragicervic et al. TAXONOMY (2009), Mofah et al. (2011), Kousteni & Megalofonou (2016), and Yigin et al. (2016) for comparisons. Family Oxynotdae Gill, 1863 Terminology of general external morphology, dermal Genus Oxynotus Rafnesque, 1810 dentcles and colouraton in descripton follows Yano Oxynotus sp. & Murofushi (1985), Garrick (1960) and Compagno Images 1–6, Table 1 (2016), and for dentton follows Herman et al. (2005). Colouraton is based on specimen preserved in 70% Material examined: SAIAB 192249, juvenile female, ethanol. Maturaton was based on Capapé et al. (1999). 519mm TL, south of Reunion Island, Madagascar Basin,

Image 1. Map of southern Africa region, showing the collecton locality (red dot) of Oxynotus sp. in the Madagascar Basin, western Indian Ocean.

11734 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

Image 2. Specimen of Oxynotus sp., SAIAB 192249, juvenile female, 519mm TL in lateral view (A), and detail of head in dorsal (B), ventral (C), and lateral (D) views. Scale bars: 50mm.

26.931250S & 56.18948330E, Trawl 1, Staton 2, Event and vertcal, situated posterior to eyes. Mouth large 7, 600m depth, trawled on 14.xi.2009 aboard R/V Dr. and transverse, constricted and circular with papillose Fridtjof Nansen, coll. K.M. Kemp, P.H. Boersch-Supan, and feshy lips, located ventrally; preoral clef elongate, O. Alvheim, D. Benivary, V. Mangar, N. Mazungula, T.B. thick, distally rounded, situated medially in the upper lip. Letessier and A.D. Rogers. Upper teeth smaller and thinner than lower teeth; upper General descripton of external morphology: teeth compressed and triangular, conspicuously slender External measurements for Oxynotus sp. are provided at crown and crown vertcally elongated, unicuspid with in Table 1. Body robust and triangular in cross-secton, pointed cusp, mesial and distal cutng edges straight; conspicuously deep at trunk and abdomen, and thin at upper teeth irregularly distributed in mouth; lower precaudal peduncle; body arched dorsally from head to teeth compressed and interlocked, broadly triangular abdomen, fatened ventrally at trunk with lateral ridges at crown, crown vertcally low, unicuspid with small prominent at trunk; body wide from head to abdomen serratons in the blades, mesial and distal cutng edges and narrow at precaudal penduncle. Head small, sigmoid. Gill slits markedly tall and vertcal. Dorsal narrow anteriorly and wide posteriorly, markedly arched fns sail-like and triangular, conspicuously upright, tall anterior-dorsally. Snout blunt at tp and short with nostrils and broad at fn web; anterior and posterior margins broad and horizontal with anterior margin of nostrils of frst and second dorsal fns straight; apex and free elongated, wide and fringed distally. Eyes elliptcal rear tps of frst and second dorsal fns pointed. Dorsal- horizontally and conspicuously large, located laterally fn spines conical, thick and markedly small, located with prominent supraorbital and infraorbital ridges and prior to each dorsal fn and never transcending the inconspicuous knob. Spiracles large, subcrescent, thin fn apex; frst dorsal-fn spine directed forwardly and

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 11735 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

© Sarah Viana & Mark Lisher to caudal fn; pelvic-fn margins straight; pelvic-fn apex and free rear tps rounded. Caudal fn small and sub- triangular with inconspicuous lower caudal lobe and evident subterminal notch; caudal fork inconspicuous; dorsal caudal margin straight; preventral caudal margin somewhat convex; terminal caudal margin straight; postventral caudal margins straight; posterior, ventral and subterminal caudal tps rounded. Dermal dentcles markedly large, measuring 1mm width across tps of lateral cusps. Dentcles heavy, sparsely distributed with diferent sizes throughout, and positoned in right angle to the longitudinal body axis. Dentcles crown-like with four pointed cusps distally, including two lateral cusps and two median cusps. Median cusps comprised of one anterior cusp and one posterior cusp. Anterior median cusp much more elongate than remaining cusps. Posterior median cusp small and hook-like. Large dentcles with three thin anterior ridges, although a single anterior ridge is shown in small dentcles. Lateral ridges thin and small. Crown base of dentcles conspicuously broad. Body brownish laterally and ventrally with large black blotches dorso-laterally throughout head, trunk and tail; black botches also found ventrally at head

Image 3. Dorsal fns of Oxynotus sp., SAIAB 192249, juvenile female, posterior to mouth tll vertcal traced at inserton of 519mm TL. A: frst dorsal fn; B: second dorsal fn. Scale bars: pectoral fns. Dorsal fns brownish, dorsal-fns posterior 20mm. margin and free rear tps white. Pectoral fns brown dorsally and blackish ventrally with pectoral-fn anterior and posterior margins broadly white. Pelvic fns dark second dorsal-fn spine directed backwardly. Pectoral brown with black blotch ventrally; pelvic-fn anterior and fns leaf-like and wide, placed laterally; pectoral-fn posterior margins whitsh. Caudal fn dark brown with anterior and inner margins markedly convex; pectoral- postventral caudal margins slightly white, black caudal fn posterior margin somewhat straight and slightly strip prominent and black caudal blotch evident in the concave distally; pectoral-fn apex pointed distally; preventral caudal margin. pectoral-fn free-rear tps inconspicuous. Pelvic fns Biological traits. The specimen weighed a total sub-quadrangular and wide, located ventrally just prior of 1.944kg. The liver is markedly heavy with total

Image 4. Pectoral (A) and pelvic (B) fns in dorsal view, and caudal fn (C) of Oxynotus sp. (SAIAB 192249, juvenile female, 519mm TL). Scale bars: 20mm.

11736 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

Image 5. Dermal dentcles of Oxynotus sp. (SAIAB 192249, juvenile female, 519mm TL) taken from below frst dorsal fn. A: dorsal view; B: anterior view; C: posterior view, showing posterior median cusp; D: anterior view of a large dentcle. weight of 518g, entrely occupying the abdominal DISCUSSION cavity. It comprises two asymmetrical lobes, somewhat cylindrical, triangular in cross-secton, pointed distally The Oxynotus specimen from the western Indian and convex proximally. Liver lobes are conspicuously Ocean is herein identfed as Oxynotus sp. due to elongate with right lobe measuring 275mm and lef inherent diferentaton in external morphology and lobe 262mm in length. The hepatosomatc index (HSI) morphometrics with specimens of O. centrina and O. for the specimen is 36.33%. According to Capapé et al. bruniensis. Oxynotus sp. is distnct from O. centrina (1999) adult females show HSI ranging from 23%–42% from the Mediterranean Sea and northeastern Atlantc while Megalofanou & Damalas (2004) notced 18.8% Ocean by having body much more robust and deeper HSI for a gravid female of O. centrina, supportng that from head to tail with heights of head, trunk, tail and HSI considerably decreases in breeding females. The caudal peduncle greater in length than in O. centrina (vs. hepatosomatc index in the immature female of Oxynotus body slender and low in O. centrina), knob behind eyes sp. comprises twice that of gravid females of O. centrina. inconspicuous (vs. conspicuous in O. centrina), dorsal Capapé et al. (1999) stated that the HSI of O. centrina fns with posterior margin straight and apex directed increases as juveniles approach maturaton, taking this upwardly (vs. conspicuously concave and apex directed into account the high HSI of Oxynotus sp. indicates that posteriorly in O. centrina), pectoral fns broad with apex the specimen was possibly approaching maturaton slightly pointed (vs. thin with apex conspicuously pointed at the tme of capture. Thus, a viable populaton of and thin in O. centrina), and caudal fn with lower caudal Oxynotus sp. may exist within the Madagascar Basin, lobe inconspicuous or small (vs. lower caudal lobe although this is the only ofcial record of this species and conspicuous in O. centrina). Oxynotus sp. also difers family from the region suggestng that the populaton is by having smaller dorsal-caudal space than those of O. rather small and vulnerable. centrina, and body brownish with many black blotches

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length, height of ffh gill slit, length of frst dorsal-fn anterior and posterior margins, height of pelvic fn, dorsal-caudal and pectoral-pelvic space, height of eyes. Bass et al. (1976) stated that the interdorsal space as percentage of precaudal length in specimens from the southeastern Atlantc Ocean (e.g., Angola, Namibia and South Africa) is smaller than those from the northeastern Atlantc Ocean and Mediterranean Sea. According to Yano & Matusuura (2002), interdorsal space as percentage of precaudal length overlaps in O. centrina and the specimen of Oxynotus from Namibia (29.1%–34.6% PCL vs. 30.1% PCL). Oxynotus sp. from the western Indian Ocean shows a much smaller interdorsal space, corresponding to 25.8% PCL, which is in accordance to the results of Bass et al. (1976) and against those of Yano & Matusuura (2002). It is important to notce, however, that the measurement provided in Yano & Matusuura (2002) for the Namibian specimen was calculated from the illustraton taken from Bass et al. (1976), which might have interfered in their results. When the interdorsal space is calculated as percentage of total length, Oxynotus sp. (20.7% TL) and O. caribbaeus (19.2% TL) taken from Yano & Matsuura (2002) overlap with those of O. centrina (18.0%–21.0% TL). Interdorsal space in Oxynotus sp. clearly difers from those of O. bruniensis (12.7% TL), O. japonicus (16.7% TL) and O. paradoxus (21.9% TL) taken from Yano & Matsuura (2002). Besides interdorsal space, Oxynotus sp. is separated from O. bruniensis sensu Ogilby (1893), Cadenat & Blache (1981), Last & Stevens (2009) and Ebert (2013) by having body dark brown with black blotches present from head to tail (vs. body sandy light brown with no evident black blotches in O. bruniensis), supraorbital and infraorbital ridges conspicuous (vs. inconspicuous in O. bruniensis), head markedly convex anterior-dorsally (vs. © Sarah Viana & Mark Lisher markedly concave anterior-dorsally in O. bruniensis), Image 6. Liver of Oxynotus sp. (SAIAB 192249, juvenile female, spiracles vertcal (vs. spiracles rounded in O. bruniensis), 519mm TL) in dorsal (A) and ventral (B) views, showing asymmetrical lobes. Scale bar: 30mm. and dorsal fn with apex pointed (vs. apex rounded in O. bruniensis). It is further distnct from O. bruniensis by having interdorsal space 1.67 tmes second dorsal- throughout the body dorsal, lateral and ventrally (vs. fn base length (vs. interdorsal space 0.93 tmes second body greyish-white with brown or reddish-brown dorsal-fn base length in O. bruniensis, taken from Yano blotches throughout the body in O. centrina). Oxynotus & Matsuura 2002). These fndings support that O. sp. has head, preorbital, pre-branchial, prespiracular bruniensis recognized in the Eastern Indian Ocean in and prepelvic lengths greater than in O. centrina. The Ebert (2013) and Weigmann (2016) has no occurrence in former species also shows pre-vent length, pelvic-caudal the western Indian Ocean. space, trunk width, interorbital space, postventral Oxynotus sp. may represent a species distnct from the caudal margin length much greater than in O. centrina. Mediterranean O. centrina and possibly an undescribed Other diferences in external measurements between species. The present study clarifes that this species occurs Oxynotus sp. and O. centrina are notced for preoral in the Madagascar Basin from of south Reunion Island

11738 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

Table 1. External measurements for Oxynotus sp. (SAIAB 192249) expressed as percentage of the total length (% TL). TL is given in millimeter. Morphometrics of O. centrina are summarized as mean values for comparisons.

Total length (mm) 519 440–545 555 210–545 690 800 – 533–790 651

mm %TL Mean %TL

Precaudal length 417.00 80.35 80.24 80 78.8 81 79 78.49 77.68 80.18

Pre-frst dorsal length 131.20 25.28 29.24 19.25 15.7 24 22 22.38 21.85 23.81 Pre-second dorsal 325.00 62.62 65.57 58.15 57.6 62 60 59.05 64.82 length Head length 107.75 20.76 15 15 17.56 18.85 18.13

Prebranchial length 91.96 17.72 15.69 15.4 15.03

Prespiracular length 60.02 11.56 8.96 9.65 8.6 9.48

Preorbital length 37.17 7.16 3.66 6 3.3 4.75

Pre-pectoral length 103.58 19.96 19.64 18.55 19.7 15 16 17.21 17.25 18.43

Pre-pelvic length 330.00 63.58 62.57 61.55 61.7 59.55 62.28

Pre-vent length 372.00 71.68 66.46 65.5

Interdorsal space 107.68 20.75 19.36 19.7 21 18 18.93 20.43

Dorsal-caudal space 45.80 8.82 10.42 10 9.15

Pectoral-pelvic space 190.00 36.61 37.2 39.68

Pelvic-caudal space 52.63 10.14 9.19 9.4

Head width 94.28 18.17

Trunk width 98.03 18.89 15.69 16

Head height 98.95 19.07 8 9 13.48 13.67

Trunk height 120.42 23.20 17.37 18 18 20 16.35 16.08 16.9

Tail height 44.68 8.61 7.73 Caudal peduncle 25.62 4.94 4 4 4.08 4.15 height Eye length 20.36 3.92 4.52 4.05 4.5 4 3 4.1 3.69

Eye height 7.18 1.38 2.05 1.5 1.6

Interorbital space 60.34 11.63 9.75 9.4 7.5

Spiracle length 16.83 3.24 2.92 3 3.1

Nostril width 17.64 3.40 2.8

Internarial space 7.91 1.52 1.13 1.8 1.2 2.85

Mouth width 32.76 6.31 5.78 4.4 5.6 7.5

Prenarial length 13.35 2.57 1.56 2.8 1.5 1.68

Preoral length 30.60 5.90 4.79 4.7 4.8 5.25

First gill slit height 8.27 1.59 1.25 1.5 1.4 1.8

Fifh gill slit height 11.94 2.30 1.46 1.5 1.6 1.85

Intergill length 19.61 3.78 4.55 4.43 First dorsal fn anterior 112.13 21.61 19 21 20.38 18.89 margin length First dorsal fn height 66.29 12.77 14.46 12.35 14.5 14 13 13.18 12.29 First dorsal fn inner 18.82 3.63 4.15 margin length First dorsal fn posterior margin 76.50 14.74 15.4 15.33 length

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 11739 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

Oxynotus sp. Oxynotus centrina Barrull Yano & Yano & Megalofonou Dragičević Mofah Kousteni & Yigin & SAIAB 192249 Murofushi Matsuura & Damalas et al. et al. Megalofonou et al. Mate (1985) (2002) (2004) (2009) (2011) (2016) (2016) (2001) n=1 n=5 n=2 n=8 n=1 n=1 n=1 n=4 n=1 First dorsal fn base 96.00 18.50 19.9 24.4 12 18 17.3 12.6 length First dorsal fn length 114.48 22.06 24.05 27.3 First dorsal-fn spine 6.70 1.29 0.9 length Second dorsal fn 87.43 16.85 15 16 17.1 15.05 anterior margin length Second dorsal fn 57.57 11.09 11.02 10.45 11.1 16 10 9.4 10.14 height Second dorsal fn inner 19.10 3.68 3.8 margin length Second dorsal fn posterior margin 55.80 10.75 10.8 10 12 10.05 8.6 length Second dorsal fn base 64.48 12.42 11.95 11.05 12 9 12 12.15 9.98 length Second dorsal fn 81.38 15.68 14.85 15.9 length Second dorsal-fn spine 3.98 0.77 0.9 length Pectoral-fn anterior 88.33 17.02 17.79 15.75 17.8 19 16 16.88 17.2 margin length Pectoral-fn inner 17.79 3.43 3.5 margin length Pectoral-fn posterior 77.38 14.91 14.2 11.6 11.95 margin length Pelvic-fn anterior 51.15 9.86 9.79 6.4 9.8 10 10 8.78 8.29 margin length Pelvic fn height 49.55 9.55 9 8 7.83 6.91 Pelvic-fn posterior 54.88 10.57 11 13 8.38 9.52 margin length Dorsal caudal margin 112.44 21.66 21.09 20.55 21.5 17 18 21.43 18.74 length Terminal caudal margin 31.05 5.98 6 6 6.3 5.22 length Preventral caudal 64.07 12.34 11.7 11 11 12.8 11.52 margin length

(not northern Madagascar as seen in Fricke et al. 2018) (2016) due to overlapping of external morphological and speculatve records elsewhere (e.g., Mozambique characters and interdorsal space. Vasil’eva et al. (2001), and Madagascar) requires future scientfc expeditons however, considers O. shubnikovi as valid following in the region. To elucidate the taxonomic status of this earlier assumptons of Bass et al. (1976), Myagkov species an ongoing taxonomic investgaton on species (1986), and Compagno et al. (1991). of the genus is being undertaken (Viana & Lisher in Preliminary comparatve morphological observatons prep.) by integratng molecular analysis of the preserved in O. centrina reveal strong intraspecifc variatons specimen and comparatve morphological examinaton between specimens from the eastern and western of material from the Eastern Atlantc Ocean, partcularly Mediterranean Sea as well as northeastern Atlantc specimens from South African and Namibian waters, Ocean regarding shape of dorsal and pectoral fns, body as well as congeners described elsewhere. Elucidaton colouraton and body depth. External measurements of the taxonomic status of other available nominal also vary for trunk height, interorbital and internarial species is also essental such as O. shubnikovi Myagkovi, spaces, mouth width, preoral length and heights of 1986 from of Kunene River in Namibia. This species frst and ffh gill slits, length of frst dorsal-fn base, and is considered junior synonym of O. centrina in Yano & length of second dorsal-fn posterior margin (see Table Matsuura (2002), Compagno (2016), and Weigmann 1). These variatons possibly represent the existence

11740 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher of another nominal species distnct from O. centrina RMNH.PISC. 17805, 20 miles from Barra Nova; RMNH. in this region as well. Oxynotus salviani Risso, 1827 PISC. 34809, 20 miles from Barra Nova. described from Nice, France has been in synonymy with O. centrina without detailed taxonomic examinaton in Kref & Tortonese (1973), demanding verifcaton. REFERENCES Divergences on the morphometric methodology applied in earlier accounts on species of Oxynotus (e.g., Yano Barrull, J. & I. Mate (2001). First confrmed record of angular roughshark Oxynotus centrina (Linnaeus, 1758) predaton on shark & Murofushi 1985; Myagkov 1986; Yano & Matsuura egg case of small-spoted catshark Scyliorhinus canicular (Linnaeus, 2002; Kousteni & Megalofonou 2016) and the present 1758) in Mediterranean waters. Annales Series Historia Naturalis study exist, and may have interfered in the assumptons 11: 23–28. Bass, A.J., J.D. D’Aubrey & N. Kistnasamy (1976). Sharks of the pointed out in the present study, indicatng that a east coast of southern Africa. VI. The families Oxynotdae, standardizaton of external measurements for Oxynotus Squalidae, Dalatidae and Echinorhinidae. Investgatonal Report, species is essental for future comparatve purposes. Oceanographic Research Insttute, Durban 45(1): 1–103. Bradai, M.N., F. Serena, I. Bianchi & D.A. Ebert (2007). Oxynotus The diferences and intraspecifc variatons observed in centrina. The IUCN Red List of Threatened Species 2007: the external morphology among species of the genus, e.T63141A12622296. Downloaded on 7 June 2017; htp://doi. org/10.2305/IUCN.UK.2007.RLTS.T63141A12622296.en however, denote that the taxonomy of the group is Cadenat, J. & J. Blache (1981). Requins de Méditerranée et d’Atlantque more complex than previously notced. Re-evaluaton of (plus partculierment de La Côte Ocidentale d’Afrique). Colleton these morphological aspects combined with molecular Fauna Tropicale XXI. ORSTOM, Paris, 330pp. Capapé, C. (2008). Diet of the Angular Rough Shark Oxynotus centrina analysis of species of Oxynotus, especially Oxynotus (Chondrichthyes: Oxynotdae) of the Languedocian Coast (Southern sp. and O. centrina, is urgently required in order to France, North-Western Mediterranean). Viet et Milieu - Life and elucidate if Oxynotus sp. is a valid species and if O. Environment 58(1): 57–61. Capapé, C., A.A. Seck & J.P. Quignard (1999). Observatons on the centrina comprises a species with broad geographical reproductve biology of the Angular Rough Shark, Oxynotus centrina distributon and strong intraspecifc variatons or if it has (Oxynotdae). Cybium 23(3): 259–271. been misidentfed with other regional congeners (e.g., Compagno, L.J.V. (1999). Checklist of living elasmobranchs, pp. 471– 498. In: Hamlet, W.C. (ed.). Sharks, Skates, and Rays. The biology of O. paradoxus). elasmobranch fshes. John Hopkins University Press, Baltmore and Oxynotus sp. is a rare deep-water species, possibly London, 528pp. comprising a small and vulnerable populaton with Compagno, L.J.V. (2016). Sharks, pp. 1123–1336. In: Carpenter, K.E. & N. De Angelis (eds.). FAO Species Identfcaton Guide for Fisheries scatered endemic distributon in the western Indian Purpose. The Living Marine Resources for Eastern Central Atlantc. Ocean. Fishery trends are currently shifing from Volume 2: Bivalves, gastropods, hagfshes, sharks, batoid fshes, and targetng coastal habitats to abyssal basins where fsh chimaeras. FAO Species Identfcaton Guide for Fishery Purposes, Rome, pp. 665-1509. stocks are stll widely available for exploitaton, including Compagno, L.J.V., D. A. Ebert & P.D. Cowley (1991). Distributon of the Madagascar Basin where this species occurs. Thus, ofshore demersal cartlaginous fsh (Class Chondrichthyes) of the West coast of Southern Africa, with notes on their systematcs. it is assumed that Oxynotus sp. may occasionally be South African Journal of Marine Science 11 (1): 43–139; htp://doi. caught as by-catch during botom trawls by commercial org/10.2989/025776191784287664 vessels in this region, as it is notced for its congeners Compagno, L.J.V., M. Dando & S. Fowler (2005). Sharks of the World. Princeton University Press, New Jersey, 368pp. elsewhere, including O. centrina, O. bruniensis, and O. Dragičević, B., J. Dulčić & C. Capapé (2009). Capture of a rare shark, japonicus (Francis 2003; Yano 2004; Bradaï et al. 2007). Oxynotus centrina (Chondrichthyes: Oxynotdae) in the eastern Moreover, the high HSI of this species provided in this Adriatc Sea. Journal of Applied Ichthyology 25(1): 56–59; htp://doi. org/10.1111/j.1439-0426.2009.01265.x study indicates that the populaton of Oxynotus sp. is Ebert, D.A. (2013). Deep-sea cartlaginous fshes of the Indian Ocean. approaching the breeding stage of the life cycle, making Volume 1: Sharks. FAO Species Catalogue for Fisheries Purposes No. this species even more vulnerable to fshing threats. It 8, Rome, 256pp. Ebert, D.A. (2015). Deep–Sea Cartlaginous Fishes of the Southeastern is of urgent conservaton concern to consider Oxynotus Atlantc Ocean. FAO Species Catalogue for Fishery Purposes, No. 9, sp. as a species vulnerable to overfshing, even though Rome, 251pp. fsheries have no apparent economic interest in rough Ebert, D.A. & K.E. Van Hees (2015). Beyond Jaws: rediscovering the ‘lost sharks’ of southern Africa. African Journal of Marine Science sharks. 37(3): 141–156; htp://doi.org/10.2989/1814232X.2015.1048730 Ebert, D.A., S. Fowler & L.J.V. Compagno (2013). Sharks of the World: A Fully Illustrated Guide. Wild Nature Press, London, 528pp. Comparatve material Ebert, D.A. & M.F.W. Stehmann (2013). Sharks, batoids and chimaeras Oxynotus centrina (fve specimens): AMS I.1752, from the North Atlantc. FAO Species Catalogue for Fisheries adult female, 635mm TL, Mediterranean Sea (stufed Purposes, No.7, Rome, 523pp. Francis, M.P. (2003). Oxynotus bruniensis. The IUCN Red List of specimen, destroyed); CAS 234693, 195mm TL, neonate Threatened Species 2003: e.T41840A10578144. Downloaded male, Guiné-Bissau; RMNH.PISC. 4248, Cantraine; on 7 June 2017; htp://doi.org/10.2305/IUCN.UK.2003.RLTS.

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742 11741 Taxonomy of a rough shark from the Western Indian Ocean Viana & Lisher

T41840A10578144.en Myagkov, N.A. (1986). Oxynotdae, pp. 168–173. In: Gubanov, E.P., Fricke, R., J. Mahafna, F. Behivoke, H. Jaonalison, M. Léopold & D. V.V. Kondyurin & N.A. Myagkov, (eds.). Sharks of the World Ocean: Ponton (2018). Annotated checklist of the fshes of Madagascar, Identfcaton handbook. [=Akuly Mirovogo okeana: Spravochnik- southWestern Indian Ocean, with 158 new records. Fishtaxa 3(1): opredelitel’]. Agropromizdat, Moscow, 272pp. 1–432. Ogilby, J.D. (1893). Descripton of a New Shark from the Tasmanian Garrick, J.A.F. (1960). Studies on New Zealand Elasmobranchii PartXI. Coast. Records of the Australian Museum 2(5): 62–63; htp://doi.or Squaloids of the genera Deania, Etmopterus, Oxynotus and Dalatas g/10.3853/j.0067-1975.2.1893.1194 in New Zealand waters. Transactons of the Royal Society of New Ragonese, S., S. Vitale, M. Dimech & S. Mazzola (2013). Abundances Zealand 88(3): 489–517. of Demersal Sharks and Chimaera from 1994-2009 Scientfc Surveys Guallart, J., P. Garcia-Salinas, A.E. Ahuir-Baraja, M. Guimerans, J.R. in the Central Mediterranean Sea. PLoS ONE 8(9): e74865; htp:// Ellis & M. Roche (2015). Angular roughshark Oxynotus centrina doi.org/10.1371/journal.pone.0074865 (Squaliformes: Oxynotdae) in captvity feeding exclusively on Rogers, A.D., O. Alvheim, E. Bemanaja, D. Benivary, P.H. Boersch- elasmobranch eggs: an overlooked feeding niche or a mater of Supan, T. Bornman, R. Cedras, N. Du Plessis, S. Gotheil, A. Hoines, individual taste? Journal of Fish Biology 87(4): 1072–1079; htp:// K. Kemp, J. Kristansen, T. Letessier, V. Mangar, N. Mazungula, T. doi.org/10.1111/jf.12761 Mørk, P. Pinet, J. Read & T. Sonnekus (2010). Cruise report “Dr. Herman, J., M. Hovestadt-Euler & D.C. Hovestadt (2005). Contributons Fridtjof Nansen”. Southern Indian Ocean Seamounts. IUCN/ GEF/ to the odontological study of living Chondrichthyes. 2. The genus UNDP/ ZSL/ ASCLME/ NERC/ EAF Nansen Project/ ECOMAR/ ACEP Oxynotus Rafnesque, 1810. Bulletn de L’Insttut Royal Des Sciences 2009 Cruise 410, 182pp. Naturelles de Belgique, Biologie 75: 5–20. Sabaj, M.H. (2016). Standard symbolic codes for insttutonal resource Ho, H-C. & K. Nakaya (2016). New record of the Japanese Roughshark, collectons in herpetology and ichthyology: an Online Reference. Oxynotus japonicus Yano & Murofushi, 1985 (Elasmobranchii: In: American Society of Ichthyologists and Herpetologists, Squaliformes: Oxynotdae) in Taiwan. Acta Ichthyologica et Washington, DC., Version 6.5 (16 August 2016), htp://www.asih. Piscatoria 46(4): 357–360; htp://doi.org/10.3750/AIP2016.46.4.08 org. Downloaded on 2 September 2016. Kref, G. & E. Tortonese (1973). Oxynotdae, pp. 35–36. In: Hureau, Soldo, A. & M. Freitas (2009). Oxynotus paradoxus. The IUCN Red List J.C. & T. Monod (eds.). Check-list of the Fishes of the North-eastern of Threatened Species 2009: e.T161361A5406422. Downloaded Atlantc and of the Mediterranean. CLOFNAM Unesco, Paris, 683pp. on 7 June 2017; htp://doi.org/10.2305/IUCN.UK.2009-2.RLTS. Kousteni, V. & P. Megalofonou (2016). Observatons on the biological T161361A5406422.en traits of the rare shark Oxynotus centrina (Chondrichthyes: Vasil’eva, E.D., N.N. Sazonov & I.A. Verigina. (2001). Types of Oxynotdae) in the Hellenic Seas. Journal of Fish Biology 89(3): Vertebrates in the Zoological Museum of Moscow University. 1880–1888; htp://doi.org/ 10.1111/jf.13077 Moscow University Publisher, Moscow, 250pp. Last, P.R. & J.D. Stevens (2009). Sharks and Rays of Australia, 2nd Yano, K. (2004). Oxynotus japonicus. The IUCN Red List of Threatened Editon. Harvard University Press, Cambridge, 644pp. Species 2004: e.T44206A10870794. Downloaded on 7 June 2017; Last, P.R., W.T. White, J.J. Pogonoski, D.C. Gledhill, G.K. Yearsleay htp://doi.org/10.2305/IUCN.UK.2004.RLTS.T44206A10870794.en & R.D. Ward (2007). Part 1 – Applicaton of a rapid taxonomic Yano, K. & M. Murofushi (1985). A New Prickly Dogfsh, Oxynotus approach to the genus Squalus, pp. 1–10. In: Last, P.R., W.T. White japonicus, from Japan. Japanese Journal of Ichthyology 32(2): 129– & J.J. Pogonoski (eds.). Descriptons of new dogfshes of the genus 135. Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Yano, K. & K. Matsuura (2002). A Review of the Genus Oxynotus Research Paper 014, Hobart, 130pp. (Squaliformes, Oxynotdae). Bulletn of the Natonal Science Leandro, L. (2004). Oxynotus caribbaeus. The IUCN Red List of Museum, Tokyo, Series A 28(2): 109–117. Threatened Species 2004: e.T44568A10907800. Downloaded Yigin, C.C., A. Ismen & U. Onal (2016). Occurrence of a rare shark, on 7 June 2017; htp://doi.org/10.2305/IUCN.UK.2004.RLTS. Oxynotus centrina (Chondrichthyes: Oxynotdae), from Saros Bay, T44568A10907800.en North Aegean Sea, Turkey. Journal of the Black Sea/Mediterranean Megalofonou, P. & D. Damalas (2004). 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e27001; htp://doi.org/10.1371/journal.pone.0027001 Threatened Taxa

11742 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 May 2018 | 10(6): 11732–11742

ISSN 0974-7907 (Online); ISSN 0974-7893 (Print)

May 2018 | Vol. 10 | No. 6 | Pages: 11703–11830 Date of Publicaton: 26 May 2018 (Online & Print) www.threatenedtaxa.org DOI: 10.11609/jot.2018.10.6.11703-11830

Communicatons Early stages and larval host plants of some northeastern Indian buterfies -- Tarun Karmakar, R. Nitn, Vivek Sarkar, Sarika Baidya, Subhajit Mazumder, Home range and spatal organizaton by the Hoary Fox Lycalopex vetulus V.K. Chandrasekharan, Rudraprasad Das, G.S. Girish Kumar, Swapnil (Mammalia: Carnivora: Canidae): response to social disrupton of two Lokhande, Joyce Veino, Lightson Veino, Rakoveine Veino, Zeeshan Mirza, neighboring pairs Rajesh V. Sanap, Bimal Sarkar & Krushnamegh Kunte, Pp. 11780–11799 -- Julio C. Dalponte, Herson S. Lima, Stuart Klorfne & Nelton C. da Luz, Pp. 11703–11709 Inventory of teloganodid mayfies (Ephemeroptera: Teloganodidae) from southern India with records of endemic taxa People’s attude towards wild elephants, forest conservaton and Human- -- C. Selvakumar, K.G. Sivaramakrishnan, T. Kubendran & Kailash Chandra, Elephant confict in Nilambur, southern Western Ghats of Kerala, India Pp. 11800–11805 -- C.K. Rohini, T. Aravindan, K.S. Anoop Das & P.A. Vinayan, Pp. 11710–11716

Analysis of regurgitated pellets of Spoted Owlet Athene brama Notes (Temminck, 1821) (Aves: Strigiformes: Strigidae) from Punjab, India -- Renuka Malhotra & Neena Singla, Pp. 11717–11724 Durga Das’s Leaf-nosed Bat Hipposideros durgadasi Khajuria, 1970 (Mammalia: Chiroptera: Hipposideridae): a new distributon record in Species diversity and abundance of birds on Bharathiar University Campus, northern India hidden in the Natonal Zoological Collectons Tamil Nadu, India -- M. Kamalakannan, Tauseef Hamid Dar & C. Venkatraman, Pp. 11806– -- L. Arul Pragasan & M. Madesh, Pp. 11725–11731 11811

On the taxonomy of the frst record of rare deep-water rough shark species A new range record of noctuid moth Owadaglaea elongata (Lepidoptera: of Oxynotdae (Chondrichthyes: Squaliformes) in the western Indian Ocean Noctuidae: Xyleninae) from India -- Sarah Viana & Mark W. Lisher, Pp. 11732–11742 -- P.R. Shashank & Balázs Benedek, Pp. 11812–11814

Forest evergreenness and tree endemism in the central Western Ghats, Natural history of Large Cabbage White Pieris brassicae nepalensis southern India Gray, 1846 (Lepidoptera: Pieridae) on Nasturtum, Tropaeolum majus -- Divakar K. Mesta & Ganesh R. Hegde, Pp. 11743–11752 (Tropaeolaceae) in Utarakhand, India -- Bhawana Kapkot Negi & Ravindra K. Joshi, Pp. 11815–11817 Distributon of Rhododendron falconeri Hook. F. (Ericales: Ericaceae) in Yuksam-Dzongri trekking corridor of Khangchendzonga Natonal Park, An account of the occurrence of Wedge Sea Hare Dolabella auricularia Sikkim, India (Lightoot, 1786) (Gastropoda: Aplysiidae) from Andaman Islands, India -- Aseesh Pandey & Hemant K. Badola, Pp. 11753–11759 -- Vikas Pandey, Ganesh Thiruchitrambalam, M. Savurirajan, Raj Kiran Lakra, Jawed Equbal, Kunal Satyam, P. Shanmukha Sainath & Rokkarukala Samson, Pp. 11818–11821 Peer Commentary New pteridophytc records from Mizoram, northeastern India The characteristcs, representatveness, functon and conservaton -- Sachin Sharma, Amit Kumar, Bhupendra Singh Kholia & Surendra Singh importance of tropical dry evergreen forest on India’s Coromandel Coast Bargali, Pp. 11822–11826 -- Mark Everard, Pp. 11760–11769 Clarke’s Morning Glory Ipomoea clarkei Hook.f. (Convolvulaceae): additon to the fora of Eastern Ghats Short Communicatons -- L. Rasingam, J. Swamy & M. Sankara Rao, Pp. 11827–11829

Mugger Crocodile Crocodylus palustris Lesson, 1831 (Reptlia: Crocodilia: Crocodylidae) in river Saberi of Godavari system in southern Odisha, India: Miscellaneous conservaton implicatons -- Subrat Debata, Swetashree Purohit, Anirban Mahata, Sudheer Kumar Jena Natonal Biodiversity Authority & Sharat Kumar Palita, Pp. 11770–11774 Member A new record of the lesser-known buterfy Small Woodbrown Lethe nicetella de Nicéville, 1887 (Lepidoptera: Nymphalidae: Satyrinae) from Khangchendzonga Natonal Park, Sikkim, India -- Sailendra Dewan, Bhoj Kumar Acharya & Sudeep Ghatani, Pp. 11775–11779

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