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Host relationships and geographic distribution of species of Acanthobothrium Blanchard, 1848 (Onchoproteocephalidea, Onchobothriidae) in elasmobranchs: a metadata analysis
expand article infoFrancisco Zaragoza-Tapia, Griselda Pulido-Flores§, Scott L. Gardner§, Scott Monks§
‡ Universidad Autónoma del Estado de Hidalgo, Hidalgo, Mexico
§ University of Nebraska, Lincoln, United States of America
Open Access

Abstract

Species of Acanthobothrium have been documented as parasites of the spiral intestine of elasmobranchs. Results of a metadata analysis indicate that 114 species of elasmobranchs have been reported as hosts of 200 species of Acanthobothrium. The metadata analysis revealed that 3.7% of species of sharks and 14.9% of species of rays that have been reported as hosts to date; some species are parasitized by more than one species of Acanthobothrium. This work provides a Category designation, as proposed by Ghoshroy and Caira (2001), for each species of Acanthobothrium. These Category designations are a tool to facilitate comparisons among members of Acanthobothrium for descriptions of new species in the future.

Keywords

Biodiversity, Elasmobranchii, Eucestoda, geographic distribution, rays, sharks

Introduction

According to Last et al. (2016b), there are 34 families comprised of 516 valid species of sharks and 26 families that include 633 valid species of rays. Since that publication, six new species of sharks and rays were described by: Yokota and Carvalho (2017) (two species of rays), Vaz and Carvalho (2018) (one species of shark), Rutledge (2019) (one species of ray), Grace et al. (2019) (one species of shark) and Concha et al. (2019) (one species of ray). This brought the current number of recognized species to 517 species of sharks and 637 species of rays.

Elasmobranchs (sharks, skates and rays) are host to a great variety of parasites in nature, particularly helminths. Acanthobothrium Blanchard, 1848 (Onchoproteocephalidea) is the most diverse genus that has been reported as parasite of the spiral intestine of elasmobranchs (Caira and Jensen 2017). At the present time, 201 species of Acanthobothrium are considered to be valid (Maleki et al. 2013; Caira and Jensen 2017; Rodríguez-Ibarra et al. 2018; Franzese and Ivanov 2018; Maleki et al. 2019; Zaragoza-Tapia et al. 2019, 2020). The genus consists of species that exclusively parasitize elasmobranchs as adults and, in many cases, individual species are thought to parasitize only a single species of elasmobranch (Caira 2011; Caira and Jensen 2017). Therefore, the genus Acanthobothrium is an excellent model for future studies of host-parasite co-speciation.

The main goal of this work is to provide a revised checklist based on a metadata analysis of the host relationships of members of Acanthobothrium and their geographic distribution based on records that have been generated from different parts of the world. The checklist focuses on the 201 valid species of Acanthobothrium and reports correlated with the genera and species of elasmobranchs, and includes the geographical distribution of each.

The number of species of Acanthobothrium continues to grow and there are still regions of the world without a single report of this genus (see Figure 1). For some time, the process of distinguishing new species of Acanthobothrium from existing species has become more and more unwieldy because of the large number of species. As an identification tool, Ghoshroy and Caira (2001) developed a categorical method for identifying species for initial comparisons. Therefore, in order to provide an update to this method, categorical designations are provided in the present checklist for each species of Acanthobothrium in the manner proposed by Ghoshroy and Caira (2001). The categories are based on and obtained from the combination of four quantitative characters: total length of the worm; the number of proglottids comprising the strobila; the number of testes per proglottid; and symmetry of the ovarian lobes. This categorical designation allows parasitologists working with this genus to postulate a group of similar species, those of the same category designation, for comparison of a new species or to aid in the preliminary identification of known species. As an additional aid, in the checklist the accession number, if known, of type specimens of each species is provided.

Materials and methods

The checklist, updated until March 2020, was based on bibliographical information from two sources of information: 1. a compilation of the records of species of Acanthobothrium as originally described, complemented by information gathered from Global Cestode Database (Caira et al. 2019) and from recent compilation studies (e.g., Ghoshroy and Caira 2001; Campbell and Beveridge 2002; Fyler and Caira 2006; Caira and Jensen 2017); and 2. information for the distribution and taxonomy of elasmobranchs that integrated a bibliographical search using different databases of literature published to date (e.g., Del Moral-Flores et al. 2015; Last et al. 2016b; Merlo-Serna and García-Prieto 2016; Alves et al. 2017) and data from FishBase (Froese and Pauly 2019).

In the checklist, the species of Acanthobothrium are arranged in alphabetical order. The scientific names and geographic distribution of elasmobranchs have been updated based on Last et al. (2016a, 2016b), Amaral et al. (2018) and Froese and Pauly (2019). The regional classification scheme of the geographic distribution of the hosts is according to Last et al. (2016b) with additional information from Froese and Pauly (2019). The following abbreviations are used for biogeographic regions (see Figure 1):

Figure 1. 

Type localities of species of Acanthobothrium reported worldwide and the biogeographic regions (Last et al. 2016b) of the geographic distribution of their hosts (see Table 1).

ARC Arctic Ocean;

ECA Eastern Central Atlantic;

ECP Eastern Central Pacific;

EIO Eastern Indian Ocean;

ENA Eastern North Atlantic;

ENP Eastern North Pacific;

ESA Eastern South Atlantic;

ESP Eastern South Pacific;

MED Mediterranean Sea;

NIO Northern Indian Ocean;

SOC Southern (Antarctic) Ocean;

WCA Western Central Atlantic;

WCP Western Central Pacific;

WIO Western Indian Ocean;

WNA Western North Atlantic;

WNP Western North Pacific;

WSA Western South Atlantic;

WSP Western South Pacific.

Information for each species of Acanthobothrium presented herein includes the name of the species, authority (original description referenced in the literature cited), abbreviation of the name of the collection where specimens are deposited and the accession numbers of the specimens, followed by the status of the specimens (holotype, paratype, neotype, syntype or voucher). The acronym “NR” was used for data that are not reported in the original source. Localities (type or/and additional localities) were given and referenced in the literature cited. A Category designation was supplied for all species using the categorical method proposed by Ghoshroy and Caira (2001).

The categorical method was developed as a system of grouping species of Acanthobothrium based on the combination of four qualitative characters: the total length of worms- ≤ 15 mm = S (short) or > 15 mm = L (long); the number of proglottids comprising the strobila- ≤ 50 = F (few) or > 50 = M (many); the number of testes per proglottid- ≤ 80 = F (few) or > 80 = M (many); and symmetry of the ovarian lobes- symmetrical = S or asymmetrical = A. Of the possible combinations the following 10 categories currently are recognized and coded as follows: 1 = SFFS; 2 = SFFA; 3 = LMMA; 4 = LMMS; 5 = LMFS; 6 = LMFA; 7 = LFFA; 8 = SMFS; 9 = LFFS; 10 = SMMS. This method limited the number of necessary comparisons required in the description between known species with new species assigned to the same Category. For this work, the categories and characteristics were used as in Ghoshroy and Caira (2001) and Fyler and Caira (2006) but the character values are as given in the original descriptions or as supplemented by the most recent taxonomic publications. In the Category designation, the type species is identified by number for this classification; the symbol “–” was used for the additional reports of species with additional hosts and/or localities.

For specimens deposited in a formal collection, acronyms are as follows:

AMS Australian Museum, Sydney;

CH-MHNJP Colecciones Helmintológicas del Museo de Historia Natural “Javier Prado” y del Instituto de Medicina Tropical “Daniel. A. Carrión”, Universidad Mayor de San Marcos, Perú;

CHE Colección de Helmintos, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo, Pachuca, México;

CHIOC Coleção Helmintológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil;

CNHE Colección Nacional de Helmintos del Instituto de Biología, Universidad Nacional Autónoma de México, México;

DMNZ Dominion Musem (=National Museum), New Zealand;

DZAUW Department of Zoology, Andhra University, Waltair, India;

DZCJ Department of Zoology, Bipin Bihari, P. G. College, Jhansi, India;

HWML University of Nebraska State Museum, Harold W. Manter Laboratory, Division of Parasitology, Lincoln, Nebraska, United States;

IPCAS Institute of Parasitology, Academy of Sciences of the Czech Republic, České Budějovice, Czech Republic;

IPMB Institut Penyelidikan Marin Borneo (Borneo Marine Research Institute), Universiti Malaysia Sabah, Kota Kinabalu, Sabah, Malaysia;

LRP Lawrence R. Penner Parasitology Collection, Helminthological Collection, University of Connecticut, Storrs, Connecticut, United States;

MACN-Pa Museo Argentino de Ciencias Naturales, Colección Parasitológica, Buenos Aires, Argentina;

MEPN Museum of the Escuela Politecnica Nacional, Quito, Ecuador;

MHNLS Museo de Historia Natural La Salle, Caracas, Venezuela;

MHNP Museo de Historia Natural, Lima, Peru;

MLP Museo de Ciencias Naturales de La Plata, Departamento de Zoología Invertebrados (Parasitología), Argentina;

MNHG Museum of Natural History, Geneva, Switzerland;

MNHN Muséum National d’Histoire Naturelle, Paris;

MNHNC Museo Nacional de Historia Natural de Chile;

MPM Meguro Parasitology Museum, Tokyo, Japan;

MZUM (P) Muzium Zoologi, Universiti Malaya, Kuala Lumpur, Malaysia;

MZUSP Museu de Zoologia da Universidade de São Paulo, Brazil;

NHMUK The Natural History Museum, London;

NMNS National Museum of Natural Science, Taichung, Taiwan;

PRLXU Parasitology Research Laboratory, Xiamen University, China;

QM Queensland Museum, Brisbane, Queensland, Australia;

SAM AHC South Australian Museum, Adelaide, Australia;

SBC Sarawak Biodiversity Center, Kuching, Sarawak, Malaysia;

SPUK School of Parasitology, Department of Zoology, University of Karachi, Pakistan;

SYSU School of Life Sciences, Sun Yat-sen University;

UAA Department of Zoology, University of Allahabad, Allahabad, India;

USNPC United States National Parasite Collection, Beltsville, Maryland, United States;

ZCUOK Zoological Collection, University of Kurdistan, Sanandaj, Iran;

ZIMC Collection of the Zoological Survey of India, Indian Museum, Calcutta and the Collection of the Department of Zoology, the University of Allahabad, India;

ZMB Natural History Museum Berlin, Germany;

ZUTC Collection of the Zoological Museum, University of Tehran, Tehran, Iran.

Results

The information obtained from the metadata analysis (Table 1) is comprised of 336 reports of the 201 valid species of Acanthobothrium. The list includes the type host of each species, type locality, and additional hosts and/or localities. Five of the elasmobranchs that were reported as hosts of Acanthobothrium were only identified to genus and four others are reported as “cf.” (= similar to) (see Table 1).

Species of Acanthobothrium reported from the different species of elasmobranchs of the world. Abbreviations: Gd = Geographical distribution; Ht = Holotype; Nt = Neotype; Pt = Paratype; Va = Voucher; Loc = Locality; Sou = Source; Cd = Category designation; * = Additional host; † = Additional locality; ‡ = Category designation obtained from Ghoshroy and Caira (2001); § = Category designation obtained from Fyler and Caira (2006); ¶ = Category designation obtained in this study from original descriptions; ** = Host identification requiring confirmation.

Species of Acanthobothrium Ht Nt, Pt or Va Species of Host Gd Loc Sou Cd
A. adlardi Campbell & Beveridge, 2002 SAM AHC 28210 SAM AHC 22723, 22724 Pristiophorus cirratus (Latham, 1794) EIO, WSP Port Stanvac, South Australia Campbell and Beveridge (2002)
A. aetiobatidis (Shipley, 1900) Southwell, 1925 NR NR Aetobatus narinari** (Euphrasen, 1790) WSA, WCA, WNA, ECA Lifu, Loyalty Islands Shipley (1900),Southwell (1925), Baer and Euzet (1962),Goldstein (1967)
A. amazonensis Mayes, Brooks & Thorson, 1978 USNPC 74806 USNPC 74807; HWML 20562 Potamotrygon circularis German, 1913 WSA Itacuari River, Brazil Mayes et al. (1978) 5‡
A. americanum Campbell, 1969 USNPC 71355 USNPC 71356 Hypanus americanus (Hildebrand & Schroeder, 1928) WSA, WCA, WNA Chesapeake Bay, Virginia, USA Campbell (1969) 6‡
A. americanum NR NR Hypanus americanus WSA, WCA, WNA Isla Margarita, Venezuela Mayes and Brooks (1981)
A. angelae Campbell & Beveridge, 2002 SAM AHC 22661 SAM AHC 22709, 22712 Hypnos monopterygius (Shaw, 1795) EIO, WSP Yarraville Shoals, South Australia Campbell and Beveridge (2002)
A. annapinkiensis Carvajal & Goldstein, 1971 MNHNC 20.003 NR Zearaja chilensis (Guichenot, 1848) ESP, WSA, Anna Pink Bay, Chile Carvajal-G. and Goldstein (1971) 2‡
A. arlenae Campbell & Beveridge, 2002 SAM AHC 28225 SAM AHC 28226 Aetobatus narinari** WSA, WCA, WNA, ECA Fog Bay, Timor Sea, North Australia Campbell and Beveridge (2002)
A. asnihae Fyler & Caira, 2006 MZUM (P) 142 USNPC 96413; LRP 3809-3812, LRP 3814 (including cross sections and SEM specimens); MZUM (P) 143–144; IPMB 77.14.04 Urogymnus polylepis (Bleeker, 1852) NIO, WCP Off Kampung Abai, Kinabatangan River, Sabah, Malaysia Fyler and Caira (2006)
A. asrinae Maleki, Malek & Palm, 2015 ZUTC 1325 ZUTC 1326; ZMB E.7569; SEM voucher ZUTC 1327 Rhynchobatus cf. djiddensis** (Forsskå, 1775) WIO, NIO Persian Gulf, Iran Maleki et al. (2015) 1
A. atahualpai Marques, Brooks & Barringa, 1997 MEPN 3029 MNHG 22098; CNHE 3029 Gymnura afuerae (Hildebrand, 1946) ECP, ESP Puerto Bolivar, Provincia de El Oro, Ecuador Marques et al. (1997a) 1‡
A. australis Robinson, 1965 AMS AMS Squalus megalops (Macleay, 1881) ENA, MED, ECA, ESA, WIO, EIO, WSP Eden, New South Wales, Australia Robinson (1965)
A. australis NR SAM AHC 22696 Squalus megalops ENA, MED, ECA, ESA, WIO, EIO, WSP Beachport, South Australia Campbell and Beveridge (2002)
A. bajaensis Appy & Dailey, 1973 USNPC 72567 USNPC 72568 Heterodontus francisci (Girard, 1855) ECP, ESP San Quintin Bay, Baja California, Mexico Appy and Dailey (1973) 4‡
A. bajaensis NR NR Heterodontus francisci ECP, ESP Newport Bay, California, USA Appy and Dailey (1973)
A. bartonae Campbell & Beveridge, 2002 SAM AHC 28235 NR Rhynchobatus djiddensis** WIO, NIO Broome, Western Australia Campbell and Beveridge (2002)
A. batailloni Euzet, 1955 NR NR Myliobatis aquila (Linnaeus, 1758) ENA, MED, ECA, ESA, WIO Mediterranean Sea, Gulfe du Lion Euzet (1955) 7(2)‡
A. batailloni*† NR MNHNC 20015 Myliobatis chilensis** Philippi, 1892 ESP Antofagasta, Chile Carvajal-G. and Jeges-G. (1980)
A. batailloni*† NR NR Myliobatis chilensis** ESP Coquimbo, Chile Carvajal-G. and Jeges-G. (1980)
A. batailloni*† NR NR Myliobatis chilensis** ESP Trujillo, Peru Escalante-A. (1986)
A. benedenii (Lönnberg, 1889) NR NR Raja clavata Linnaeus, 1758 ENA, MED, ECA, ESA, WIO Mediterranean Sea Lönnberg (1889) 2
A. benedenii*† NR NR Pteroplatytrygon violacea** (Bonaparte, 1832) ENP, ECP, ESP, WSA, WCA, WNA, ENA, MED, ECA, ESA, WIO, NIO, EIO, WSP, WCP, WNP Naples, Italy Baer (1948)
A. benedenii*† NR NR Torpedo marmorata** Risso, 1810 ENA, MED, ECA, ESA Casablanca, Marruecos Euzet (1952), Euzet (1959)
A. bengalense Baer & Euzet, 1962 NR NR Pastinachus sephen (Forsskål, 1775) NIO Nagapattinam, India Baer and Euzet (1962)
A. blairi Campbell & Beveridge, 2002 SAM AHC 28211 SAM AHC 28212 Dipturus whitleyi (Iredale, 1938) EIO, WSP Stanley, Tasmania Campbell and Beveridge (2002)
A. blairi NR NR Dipturus whitleyi EIO, WSP Spencer Gulf, South Australia Campbell and Beveridge (2002)
A. bobconniorum Fyler & Caira, 2010 QM G232499 QM G232500–G232501; USNPC 104278; LRP 7583–7585; cross sections of one paratype worm and voucher LRP 7586, 7588, 7589, SEM LRP 7587–7590 Rhynchobatus laevis** (Bloch & Schneider, 1801) NIO, WNP Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia Fyler and Caira (2010) 2
A. brachyacanthum Riser, 1955 USNPC 37418 NR Raja stellulata (Gilbert, 1915) ENP, ECP Monterey Bay, California, USA Riser (1955) 2‡
A. brachyacanthum* NR NR Beringraja binoculata** (Gilbert, 1855) ENP, ECP Monterey Bay, California, USA Riser (1955)
A. brayi Campbell & Beveridge, 2002 SAM AHC 22670 SAM AHC 22730 Sutorectus tentaculatus (Peters, 1864) EIO, WSP Eastern Shoal, South Australia Campbell and Beveridge (2002)
A. brevissime Linton, 1909 USNPC 9008 NR Hypanus say (Lesueur, 1817) WSA, WCA, WNA Dry Tortugas, Florida, USA Linton (1908), Goldstein (1964) 2‡
A. brevissime*† NR NR Raja eglanteria Bosc, 1800 WCA, WNA Gulf of Mexico, Chesapeake Bay, Virginia, USA Campbell (1969)
A. brevissime*† NR USNPC 71349, 71350 Hypanus americanus WSA, WCA, WNA Gulf of Mexico, Chesapeake Bay, Virginia, USA Campbell (1969)
A. brevissime*† NR CH-MHNJP 727 Myliobatis peruvianus** Garman, 1913 ESP Lima, Peru Tantaleán-Vidaurre (1991)
A. brevissime USNPC 9008 USNPC 60178 (neotype) Hypanus say WSA, WCA, WNA Gulf of Mexico, Chesapeake Bay, Virginia, USA Campbell (1969), Vardo-Zalik and Campbell (2011)
A. bullardi Ghoshroy & Caira, 2001 CNHE 4045 CNHE 4046–4047; LRP 2060–2062; USNPC 90466–90468 Hypanus dipterurus (Jordan & Gilbert, 1880) ECP Bahía de Los Angeles, Gulf of California, Mexico Ghoshroy and Caira (2001) 2‡
A. bullardi NR NR Hypanus dipterurus ECP Puertecitos, Gulf of California, Mexico Ghoshroy and Caira (2001)
A. bullardi NR NR Hypanus dipterurus ECP Santa Rosalia, Gulf of California, Mexico Ghoshroy and Caira (2001)
A. cairae Vardo-Zalik & Campbell, 2011 USNPC 103801 USNPC 103802–103814 Bathytoshia centroura (Mitchill, 1815) WSA, WCA, WNA Narragansett Bay off Sakonnet Point, Rhode Island, USA Vardo-Zalik and Campbell (2011) 3
A. campbelli Marques, Brooks & Monks, 1995 MNHG 20014 MNHG 20015–20016; HWML 38546; CNHE 3033 Urotrygon chilensis (Günther, 1872) ECP, ESP Costa de Pajaros, Puntarenas, Costa Rica Marques et al. (1995) 2‡
A. campbelli*† NR MEPN 3033 Hypanus longus (Garman, 1880) ECP Puerto Huatulco, Provincia de El Oro, Ecuador Marques et al. (1997a)
A. cannoni Campbell & Beveridge, 2002 SAM AHC 28236 SAM AHC 28237 Himantura uarnak (Gmelin, 1789) WIO, NIO, EIO, WCP Fog Bay, Timor Sea, North Australia Campbell and Beveridge (2002)
A. cartagenensis Brooks & Mayes, 1980 USNPC 75159 NR Urobatis jamaicensis (Cuvier, 1816) WCA, WNA Cartagena, Colombia Brooks and Mayes (1980) 1
A. cartagenensis NR CNHE 9706; HWML 101020; CHE P00061 Urobatis jamaicensis WCA, WNA Ría Lagartos, Yucatán, Quintana Roo Monks et al. (2015)
A. cartagenensis NR CNHE 9706; HWML 101020; CHE P00061 Urobatis jamaicensis WCA, WNA Isla Contoy, Quintana Roo Monks et al. (2015)
A. cartagenensis NR CNHE 9706; HWML 101020; CHE P00061 Urobatis jamaicensis WCA, WNA Isla Cozumel, El Paso de los Cedros, Quintana Roo Monks et al. (2015)
A. cartagenensis NR CNHE 9706; HWML 101020; CHE P00061 Urobatis jamaicensis WCA, WNA Xcalak, Quintana Roo Monks et al. (2015)
A. cestraciontis (Yamaguti, 1934) NR NR Heterodontus japonicus Miklouho-Maclay & Macleay, 1884 WNP, WCP Pacific Ocean, Japan Yamaguti (1934)
A. cestraciontis NR NR Sphyraena japonica** (Bloch &Schneider, 1801) ? Pacific Ocean, Japan Goldstein (1967)
A. chabahariense Maleki, Malek & Rastgoo, 2018 ZCUOK 100 ZCUOK 101– 112 and (SME specimen) ZCUOK 113 Pastinachus cf. sephen** NIO Chabahar coasts, the coast of the Gulf of Oman, Iran Maleki et al. (2018) 1
A. chengi Cornford, 1974 USNPC 72958 USNPC 72959 Bathytoshia lata (Garman, 1880) ECP, ENA, MED, ECA, WIO, NIO, EIO, WSP, WCP, WNP Oahu, Hawaii Cornford (1974)
A. chilensis Rego, Vincente & Herrera, 1968 CHIOC 30.308 a-c NR Sarda chiliensis** (Cuvier, 1832) ? Paita, Piúra, Peru Rêgo et al. (1968) 3‡
A. chisholmae Campbell & Beveridge, 2002 SAM AHC 28223 SAM AHC 28224 Pastinachus sephen** NIO Nickol Bay, Western Australia Campbell and Beveridge (2002)
A. cimari Marques, Brooks & Monks, 1995 MNHG 20017 MNHG 20018–20020; HWML 38547 Hypanus longus ECP Punta Morales, Puntarenas Province, Costa Rica Marques et al. (1995) 2‡
A. clarkeae Campbell & Beveridge, 2002 SAM AHC 28349 SAM AHC 28350 Urolophus paucimaculatus Dixon, 1969 EIO, WSP Queenscliff, Victoria, Australia Campbell and Beveridge (2002)
A. clarkeae*† NR SAM AHC 28243, 28244 Urolophus cruciatus (Lacepède, 1804) EIO, WSP Devonport, Tasmania Campbell and Beveridge (2002)
A. clarkeae*† NR SAM AHC 28208 Urolophus expansus McCulloch, 1916 EIO Beachport, South Australia Campbell and Beveridge (2002)
A. cleofanus Monks, Brooks & Lonce de Leon, 1996 CNHE 2670 CNHE 2671; MNHG 38576; HWML 38576. Hypanus longus ECP Chamela Bay, Jalisco, Mexico (Monks et al. 1996) 3‡
A. colombianum Brooks & Mayes, 1980 USNPC 75160 USNPC 75161 Aetobatus narinari WSA, WCA, WNA, ECA Cartagena, Colombia Brooks and Mayes (1980) 9‡
A. confusum Baer & Euzet, 1962 NR NR Neotrygon kuhlii** (Müller & Henle, 1841) WSP Indian Ocean, Sri Lanka Baer and Euzet (1962)
A. coquimbensis Carvajal & Jeges, 1980 MNHNC 20016 NR Myliobatis chilensis ESP Antofagasta, Chile Carvajal-G. and Jeges-G. (1980) 2‡
A. coquimbensis NR NR Myliobatis chilensis ESP Coquimbo, Chile Carvajal-G. and Jeges-G. (1980)
A. coronatum (Rudolphi, 1819), Blanchard, 1848 NR NR Dipturus batis (Linnaeus, 1758) ENA Mediterranean Sea, Italy Rudolphi (1819), Baer (1948)
A. coronatum* NR NR Scyliorhinus stellaris (Linnaeus, 1758) ENA, MED, ECA Mediterranean Sea, Italy Rudolphi (1819), Baer (1948)
A. coronatum* NR NR Torpedo marmorata ENA, MED, ECA, ESA Mediterranean Sea, Italy Rudolphi (1819), Baer (1948)
A. coronatum* NR NR Torpedo torpedo (Linnaeus, 1758) ENA, MED, ECA Mediterranean Sea, Italy Rudolphi (1819), Baer (1948)
A. coronatum* NR NR Dasyatis pastinaca (Linnaeus, 1758) ENA, MED, ECA Mediterranean Sea, Italy Rudolphi (1819), Baer (1948)
A. coronatum*† NR NR Hemitrygon akajei** (Müller & Henle, 1841) WNP Nakatsu, West Japan Yoshida (1917)
A. coronatum*† NR NR Aetobatus narinari** WSA, WCA, WNA, ECA Batavia, Java, Indonesia MacCallum (1921)
A. coronatum*† NR NR Scyliorhinus stellaris ENA, MED, ECA Sète, France Euzet (1959)
A. coronatum*† NR NR Scyliorhinus stellaris ENA, MED, ECA Concarneau, France Euzet (1959)
A. coronatum*† NR NR Mustelus mustelus (Linnaeus, 1758) ENA, MED, ECA, ESA Naples, Italy Euzet (1959)
A. coronatum*† NR NR Scyliorhinus stellaris ENA, MED, ECA Cardigan Bay, UK Rees and Williams (1965)
A. coronatum* NR NR Carcharodon carcharias (Linnaeus, 1758) MED Mediterranean Sea Goldstein (1967)
A. coronatum*† NR MNHG 40003, 40009 Scyliorhinus canicula (Linnaeus, 1758) ENA, MED, ECA Naples, Italy Euzet (1959), Vardo-Zalik and Campbell (2011)
A. costarricense Marques, Brooks & Monks, 1995 MNHG 20008 MNHG 20009–20010; HWML 38544; CNHE 3034 Hypanus longus ECP Punta Morales, Puntarenas Province, Costa Rica Marques et al. (1995) 2‡
A. costarricense NR MEPN 3034 Hypanus longus ECP Puerto Huatulco, Provincia de El Oro, Ecuador Marques et al. (1997a)
A. crassicolle Wedl, 1855 NR MNHG 40014 88/77 Dasyatis pastinaca ENA, MED, ECA Arcacho, Gironde, France Dollfus (1926), Baer (1948), Goldstein (1967)
A. cribbi Campbell & Beveridge, 2002 SAM AHC 28251 SAM AHC 28252 Gymnura australis (Ramsay & Ogilby, 1886) EIO, WSP, WCP Gulf of Carpentaria, Northern Territory, Australia Campbell and Beveridge (2002)
A. dasi Ghoshroy & Caira, 2001 CNHE 4043 CNHE 4044; HWML 15549–15551; LRP 2051–2054; USNPC 90463–90465 Hypanus dipterurus ECP Puertecitos, Gulf of California, Mexico Ghoshroy and Caira (2001) 2‡
A. dasybati Yamaguti, 1934 NR NR Hemitrygon akajei WNP Tarumi, Kobe, Japan Yamaguti (1934)
A. dasybati*† NR NR Okamejei kenojei** (Müller & Henle, 1841) WNP Maisaka, Japan Yamaguti (1952)
A. dasybati*† NR NR Urolophus sp.** (U. fuscus?) ? Hamazima, Mie, Japan Yamaguti (1952)
A. dighaensis Srivastava & Capoor, 1980 UAA NR Pateobatis uarnacoides (Bleeker, 1852) NIO, WCP Digha, Orissa, India Srivastav and Capoor (1980)
A. dollyae Caira & Burge, 2001 CNHE 4169 CNHE 4170; LRP 2097–2101; USNPC 90837–90839 Diplobatis ommata (Jordan and Gilbert, 1890) ECP Bahía de Los Angeles, Gulf of California, Mexico Caira and Burge (2001) 1
A. dollyae NR NR Diplobatis ommata ECP Isla San Esteban, Gulf of California, Mexico Caira and Burge (2001)
A. dollyae NR NR Diplobatis ommata ECP Punta Arena, Gulf of California, Mexico Caira and Burge (2001)
A. dujardini van Beneden, 1850 NR NR Raja clavata ENA, MED, ECA, ESA, WIO English Channel,Belgium van Beneden (1850), Goldstein (1967)
A. dujardini NR NR Raja clavata ENA, MED, ECA, ESA, WIO English Channel, Belgium Williams (1969)
A. dujardini*† NR NR Raja brachyura** Lafont, 1871 ENA, MED, ECA Roscoff, France Euzet (1959)
A. dujardini*† NR NR Raja montagui** Fowler, 1910 ENA, MED British Isles Williams (1960)
A. dysbiotos (MacCallum, 1921) Williams, 1969 NR NR Aetobatus narinari** WSA, WCA, WNA, ECA Batavia, Java, Indonesia MacCallum (1921), Williams (1969)
A. edmondsi Campbell & Beveridge, 2002 SAM AHC 28205 SAM AHC 28206, 22704 Parascyllium ferrugineum McCulloch, 1911 EIO, WSP Port Stanvac, South Australia Campbell and Beveridge (2002)
A. edmondsi NR NR Parascyllium ferrugineum EIO, WSP Holdfast Bay, South Australia Campbell and Beveridge (2002)
A. edmondsi NR NR Parascyllium ferrugineum EIO, WSP Esperance, Western Australia Campbell and Beveridge (2002)
A. edwardsi Williams, 1969 NR NR Leucoraja fullonica (Linnaeus, 1758) ENA, MED, ARC West coast of Britain, United Kingdom Williams (1969)
A. electricolum Brooks & Mayes, 1978 USNPC 74728 USNPC 74729 Narcine brasiliensis (Olfers, 1831) WSA Caribbean Sea, near Cartagena, Colombia Brooks and Mayes (1978) 9‡
A. elongatum Subhapradha, 1955 NR NR Rhynchobatus djiddensis WIO, NIO Madras Coast, India Subhapradha (1955) 4
A. etini Fyler & Caira, 2006 MZUM (P) 145 USNPC 96414–96415; LRP 3815-3824 (including cross sections and SEM specimens); MZUM (P) 146; IPMB 77.14.05 Urogymnus polylepis NIO, WCP Off Kampung Abai, Kinabatangan River, Sabah, Malaysia Fyler and Caira (2006)
A. filicolle (Zschokke, 1888) Yamaguti, 1959 NR NR Torpedo marmorata ENA, MED, ECA, ESA Mediterranean Sea Zschokke (1888), Yamaguti (1959b) 1(8)
A. filicolle* NR NR Torpedo torpedo ENA, MED, ECA Mediterranean Sea Williams (1969)
A. floridensis Goldstein, 1964 USNPC 60025 NR Raja eglanteria WCA, WNA Gulf of Mexico and Coast of Massachusetts Goldstein (1964) 8(10)‡
A. floridensis*† NR USNPC 103848–103850 Raja texana Chandler, 1921 WCA Gulf of Mexico Vardo-Zalik and Campbell (2011)
A. floridensis NR NR Raja eglanteria WCA, WNA Gulf of Mexico, Chesapeake Bay, Virginia, USA, USA Campbell (1969)
A. fogeli Goldstein, 1964 USNPC 60024 NR Gymnura micrura (Bloch & Schneider, 1801) WSA, WCA, WNA, ECA Northeastern Gulf of Mexico, Florida Goldstein (1964) 1‡
A. fogeli NR NR Gymnura micrura WSA, WCA, WNA, ECA Isla Margarita, Venezuela Mayes and Brooks (1981)
A. foulki Reyda & Caira, 2006 MZUM (P) 168(h) USNPC 97463–97464; LRP 3850–3853 (including cross sections and SEM specimens); MZUM (P) 169(p)–171(p); IPMB 77.08.14 Pateobatis uarnacoides NIO, WCP Off Kampung Tetabuan, Sabah, Malaysia Reyda and Caira (2006) 1
A. franus Marques, Centritto & Stewart, 1997 CNHE 3139 USNPC 87374; CHIOC 33754a, b; CNHE 3140 Narcine entemedor Jordan & Starks, 1895 ECP Cuajiniquil Beach, Gulf of Santa Helena, Guanacaste, Costa Rica Marques et al. (1997b) 5(8)‡
A. fylerae Maleki, Malek & Palm, 2015 ZUTC 1319 ZUTC 1320–1323; ZMB E.7568; SEM voucher ZUTC 1324 Rhynchobatus cf. djiddensis** WIO, NIO Gulf of Oman, Iran Maleki et al. (2015) 1
A. gasseri Campbell & Beveridge, 2002 SAM AHC 28217 SAM AHC 28218 Pastinachus sephen** NIO Nickol Bay, Western Australia Campbell and Beveridge (2002)
A. gibsoni Campbell & Beveridge, 2002 SAM AHC 28239 NR Rhynchobatus djiddensis** WIO, NIO Fog Bay, Timor Sea, North Australia Campbell and Beveridge (2002)
A. giganticum Sanaka, Lakshmi & Hanumantharao, 1993 NR NR Gymnura micrura** WSA, WCA, WNA, ECA Waltair coast, India Sanaka et al. (1993)
A. gloveri Campbell & Beveridge, 2002 SAM AHC 22600 SAM AHC 22715 Trygonorrhina fasciata Müller & Henle, 1841 WSP Goolwa, South Australia Campbell and Beveridge (2002)
A. gnomus Reyda & Caira, 2006 MZUM (P) 172(h) USNPC 97465–97466; LRP 3854–3859 (includes cross sections and SEM specimens); MZUM (P) 173(p)–175(p); IPMB 77.08.15 Pateobatis uarnacoides NIO, WCP Off Kampung Tetabuan, Sabah, Malaysia Reyda and Caira (2006) 1
A. goldsteini Appy & Dailey, 1973 USNPC 72569 USNPC 72570 Platyrhinoidis triseriata (Jordan & Gilbert, 1880) ENP, ECP Seal Beach, California, USA Appy and Dailey (1973) 5(9)‡
A. gonzalesmugaburoi Severino & Sarmiento, 1979 CH-MHNJP 340 CH-MHNJP 341, 341a, 341b Myliobatis peruvianus ESP Callao, Lima, Peru Severino and Sarmiento (1979) 7(6)
A. gracile Yamaguti, 1952 NR NR Narke japonica (Temminck & Schlegel, 1850) WNP Tokushima, Japan Yamaguti (1952)
A. grandiceps Yamaguti, 1952 MPM 22638 NR Telatrygon zugei (Müller & Henle, 1841) WCP, WNP East China Sea, Japan Yamaguti (1952), Yang et al. (2016)
A. grandiceps* NR NR Hemitrygon akajei WNP East China Sea, Japan Yamaguti (1952)
A. guanghaiense Yang, Sun, Zhi, Iwaki, Reyda & Yang, 2016 MPM 21229 MPM 21230; SYSU 20140818-1-4 Hemitrygon akajei WNP Off Guanghai Port, Taishan, Guangdong Province, China Yang et al. (2016) 2
A. halehae Maleki, Malek & Palm, 2019 ZCUOK 127 ZCUOK 128–130; ZUTC Platy. 1342–1343, 1 SEM voucher ZUTC Platy. 1344 Gymnura cf. poecilura** (Shaw, 1804) NIO, EIO, WCP, WNP Chabahar coast, Gulf of Oman, Iran Maleki et al. (2019) 1
A. hanumantharaoi Rao, 1977 NR NR Aetomylaeus nichofii (Bloch & Schneider, 1801) NIO, EIO, WCP, WNP Waltair coast, Benegal Bay, India Rao (1977)
A. herdmani Southwell, 1912 NR NR Neotrygon kuhlii** WSP Ceylon Pearl Bank, Sri Lanka Southwell (1912), Southwell (1925), Southwell (1930) 3
A. heterodonti Drummond, 1937 NR NR Heterodontus portusjacksoni (Meyer, 1793) EIO, WSP Lady Julia Perey Island, Victoria, Australia Drummond (1937)
A. heterodonti NR SAM AHC 22595, 22597, 15744 Heterodontus portusjacksoni EIO, WSP Derwent Estuary, Hobart, Tasmania Campbell and Beveridge (2002)
A. heterodonti NR NR Heterodontus portusjacksoni EIO, WSP Bunbury, Western Australia Campbell and Beveridge (2002)
A. himanturi Brooks, 1977 USNPC 73963 USNPC 73964; HWML 20260 Styracura schmardae (Werner, 1904) WCA Caribbean Sea, La Cienaga, Magdalena, Colombia Brooks (1977) 1‡
A. hispidum Riser, 1955 USNPC 37416 NR Tetronarce californica (Ayres, 1855) ENP, ECP, WNP Monterey Bay, California, USA Riser (1955) 5‡
A. holorhini Alexander, 1953 USNPC 47853 USNPC 47854 Myliobatis californicus Grill, 1865 ENP, ECP Long Beach Harbor, California, USA Alexander (1953) 3‡
A. holorhini*† NR CHIMTDC 542 Myliobatis chilensis ESP Callao, Peru Rodriguez and Tantaleán-Vidaurre (1980)
A. hypanus Zaragoza-Tapia, Pulido-Flores & Monks, 2020 CNHE 11255 CNHE 11256; HWML 216261 Hypanus longus ECP La Puntilla, Mazatlán, Sinaloa, Mexico Zaragoza-Tapia et al. (2020) 2
A. hypermekkolpos Fyler & Caira, 2010 QM G232506 QM G232507; USNPC 104280; LRP 7591, hologenophores LRP 7592–7593 Rhynchobatus laevis** NIO, WNP Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia Fyler and Caira (2010) 1
A. icelandicum Manger, 1972 NR NR Dipturus batis ENA Faxa Bay, Western coasts Iceland Manger (1972)
A. ijimai Yoshida, 1917 NR MPM 22639 Hemitrygon akajei WNP Tokyo, Japan Yoshida (1917), Williams (1969),Yang et al. (2016)
A. ijimai NR NR Hemitrygon akajei WNP East China Sea, Japan Yamaguti (1952)
A. inbiorium Marques, Centritto & Stewart, 1997 CNHE 3137 USNPC 87373; CHIOC 33753a, b; CNHE 3138 Narcine entemedor ECP Cuajiniquil Beach, Gulf of Santa Helena, Guanacaste, Costa Rica Marques et al. (1997b) 5‡
A. incognita (MacCallum, 1921) Wardle & McLeod, 1952 NR NR Dasyatis pastinaca ENA, MED, ECA New York Aquarium MacCallum (1921), Southwell (1925), Williams (1969)
A. indicum (Subhapradha, 1955) NR NR Narcine brasiliensis** WSA Madras Coast, India Subhapradha (1955), Williams (1969)
A. intermedium Perrenoud, 1931 NR NR Dasyatis pastinaca** ENA, MED, ECA Tauranga, New Zealand Perrenoud (1931)
A. jalalii Maleki, Malek & Palm, 2013 ZUTC 1291 ZUTC 1292–1295), SEM voucher ZUTC 1296); IPCAS C–639); ZMB E.7559 Pastinachus cf. sephen** NIO Gulf of Oman, Iran Maleki et al. (2013) 1
A. jamesi Maleki, Malek & Palm, 2015 ZUTC 1328 ZMB E.7570; SEM voucher ZUTC 1329. Rhynchobatus cf. djiddensis** WIO, NIO Persian Gulf, Iran Maleki et al. (2015) 1
A. janineae Maleki, Malek & Palm, 2015 ZUTC 1311 ZUTC 1312–1316; ZMB E.7566; SEM vouchers ZUTC 1317–1318 Rhynchobatus cf. djiddensis** WIO, NIO Gulf of Oman, Iran Maleki et al. (2015) 1
A. jeanneae Fyler & Caira, 2010 QM G232502 QM G232503–G232505; USNPC 104279; LRP 7573–7575, cross sections of one paratype worm and voucher LRP 7580–7582, SEM LRP 7576–7578, hologenophore LRP 7579 Rhynchobatus laevis** NIO, WNP Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia Fyler and Caira (2010) 1
A. jonesi Campbell & Beveridge, 2002 SAM AHC 28227 SAM AHC 28228 Dasyatis sp.** ? Cape Ford, North Australia Campbell and Beveridge (2002)
A. karachiense Bilqees, 1980 NR SPUK 2000 (syntype) Mustelus manazo Bleeker, 1855 NIO, WCP, WNP Karachi Coast, Pakistan Bilqees (1980)
A. kurdistanense Maleki, Malek & Palm, 2019 ZCUOK 122 ZCUOK 123–127; ZUTC Platy. 1336–1340, 1 SEM voucher ZUTC Platy. 1341 Gymnura cf. poecilura** NIO, EIO, WCP, WNP Chabahar coast, Gulf of Oman, Iran Maleki et al. (2019) 1
A. larsoni Reyda & Caira, 2006 MZUM (P) 176(h) USNPC 97467–97468; LRP 3860–3865 (including cross sections and SEM specimens); MZUM (P) 177(p)–180(p); IPMB 77.08.16 Pateobatis uarnacoides NIO, WCP Off Kampung Tetabuan, Sabah, Malaysia Reyda and Caira (2006) 1
A. lasti Campbell & Beveridge, 2002 SAM AHC 28247 SAM AHC 28248 Rhynchobatus djiddensis** WIO, NIO Broome, Western Australia Campbell and Beveridge (2002)
A. latum Yamaguti, 1952 MPM 22637 NR Hemitrygon akajei WNP Sea of Ariake, Kyusyu, Japan Yamaguti (1952), Yang et al. (2016)
A. laurenbrownae Campbell & Beveridge, 2002 SAM AHC 28215 SAM AHC 28216 Pastinachus sephen NIO Nickol Bay, Western Australia Campbell and Beveridge (2002)
A. lentiginosum Vardo-Zalik & Campbell, 2011 USNPC 103815 USNPC 103816–103819 Pseudobatos lentiginosus (Garman, 1880) WCA, WNA Gulf of Mexico Vardo-Zalik and Campbell (2011) 1
A. lepidum Reyda & Caira, 2006 MZUM (P) 181(h) USNPC 97469; LRP 3866–3868 (including cross sections and SEM specimens); MZUM (P) 182(p)–183(p); IPMB 77.08.17 Pateobatis uarnacoides NIO, WCP Off Kampung Tetabuan, Sabah, Malaysia Reyda and Caira (2006) 1
A. lilium Baer & Euzet, 1962 NR NR Dasyatis sp.** ? Ceylon Pearl Bank, Sri Lanka Baer and Euzet (1962)
A. lineatum Campbell, 1969 USNPC 71353 USNPC 71354 Hypanus americanus WSA, WCA, WNA Chesapeake Bay, Virginia, USA Campbell (1969) 1‡
A. lintoni Goldstein, Henson & Schlicht, 1968 USNPC 62938 USNPC 62939 Narcine brasiliensis** WSA Gulf of Mexico, Texas, USA Goldstein et al. (1969) 1(8,9,5)‡
A. lintoni NR USNPC 74851 Narcine brasiliensis** WSA Gulf of Mexico, Florida, USA Goldstein et al. (1969)
A. longipedunculata Meheswari, Sanaka, Lakshmi & Rao, 1985 NR NR Himantura uarnak WIO, NIO, EIO, WCP Waltair coast, India Maheswari et al. (1985)
A. lusarmientoi Severino & Verano, 1980 CH-MHNJP 342 CH-MHNJP 343, 343a Sympterygia brevicaudata (Cope, 1877) ECP, ESP Callao, Lima, Peru Severino and Verano (1980) 7
A. macracanthum Southwell, 1925 NR NR Urogymnus sp.** ? Madras Coast, India Southwell (1925)
A. macrocephalum Wang & Yang, 2001 MPM 21231 NR Hemitrygon akajei WNP Xiamen, Fujiari, China Wang and Yang (2001), Yang et al. (2016)
A. macrocephalum MPM 21231 MPM 21232; SYSU 20140620-1-7 Hemitrygon akajei WNP Off Guanghai Port, Guangdong, China Yang et al. (2016)
A. macrocephalum NR NR Hemitrygon akajei WNP Sanya Fishing Port, Sanya, Hainan, China Yang et al. (2016)
A. maculatum Riser, 1955 USNPC 37417 NR Myliobatis californicus ENP, ECP Monterey Bay, California, USA Riser (1955) 6(3)‡
A. magnum Euzet, 1959 NR NR Pteroplatytrygon violacea ENP, ECP, ESP, WSA, WCA, WNA, ENA, MED, ECA, ESA, WIO, NIO, EIO, WSP, WCP, WNP Mediterranean Sea, France Euzet (1959)
A. makranense Maleki, Malek & Palm, 2019 ZCUOK 130 ZCUOK 131–135; ZUTC Platy. 1345–1350, 1 SEM voucher ZCUOK 139, 1 SEM voucher ZUTC Platy. 1350 Gymnura cf. poecilura** NIO, EIO, WCP, WNP Chabahar coast, Gulf of Oman, Iran Maleki et al. (2019) 1
A. manteri Hassan, 1983 IHAHE S1051/A IHAHE S1051/B Pastinachus sephen** NIO Mediterranean Sea, Egypt Hassan (1983)
A. margieae Fyler, 2011 NMNS 6356–001 NMNS 6356–002, 6356–003, 6356–004, 6356–005, 6356–006, 6356–007; LRP 7468–7477; USNPC 103274 Orectolobus japonicus Regan, 1906 WNP, WCP Off Penghu Island, East China Sea, Magong, Taiwan Fyler (2011) 8
A. marplatensis Ivanov & Campbell, 1998 MLP 4025 MLP 4026; USNMPC 87475; NHMUK 1998.2.10.1-2 Atlantoraja castelnaui (Miranda Ribeiro, 1907) WSA Mar del Plata, Buenos Aires, Argentina Ivanov and Campbell (1998) 1‡
A. marquesi Rodríguez-Ibarra, Pulido-Flores, Violante-González & Monks, 2018 CNHE 10554 CNHE 10555, 10556; HWML 139377–139384; CHE P00061–P00063 Aetobatus cf. narinari** WSA, WCA, WNA, ECA Laguna de Términos, Ciudad del Carmen, Campeche, Mexico Rodríguez-Ibarra et al. (2018) 3
A. marquesi NR NR Aetobatus cf. narinari** WSA, WCA, WNA, ECA Champotón, Campeche, Mexico Rodríguez-Ibarra et al. (2018)
A. martini Campbell & Beveridge, 2002 SAM AHC 28213 SAM AHC 28214 Myliobatis tenuicaudatus Hector, 1877 EIO, WSP Bunbury, Western Australia Campbell and Beveridge (2002)
A. maryanskii Caira & Burge, 2001 CNHE 4171 CNHE 4172; LRP 2012, 2013; USNPC 90840, 90841 Diplobatis ommata ECP Loreto, Gulfo of California, Mexico Caira and Burge (2001) 5
A. marymichaelorum Twohig, Caira & Fyler, 2008 MZUM(P) 699(H) MZUM(P) 700(P)–702(P); SBC P–00028; USNPC 100700; LRP 4162–4164 (whole mount), 4167–4168 (cross sections) Brevitrygon walga (Müller & Henle, 1841) NIO Off Sematan, Sarawak, Malaysia Twohig et al. (2008) 1
A. marymichaelorum NR NR Brevitrygon walga NIO Off Mukah, Sarawak, Malaysia. Twohig et al. (2008)
A. masnihae Fyler & Caira, 2006 MZUM (P) 147 USNPC 96416–96417; LRP 3825-3835 (including cross sections and SEM specimens); MZUM (P) 148; IPMB 77.14.06 Urogymnus polylepis NIO, WCP Kampung Abai, Kinabatangan River, Sabah, Malaysia Fyler and Caira (2006)
A. mathiasi Euzet, 1959 NR NR Mustelus mustelus ENA, MED, ECA, ESA Sète, France Euzet (1959)
A. mathiasi* NR NR Mustelus canis (Mitchill, 1815) WNA, WCA, WSA Sète, France Euzet (1959)
A. matttaylori Fyler & Caira, 2010 QM G232508 Hologenophore USNPC 104281 Rhynchobatus laevis** NIO, WNP Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia Fyler and Caira (2010) 4
A. micracantha Yamaguti, 1952 NR MPM 22635, 22636 Hemitrygon akajei WNP Nagasaki, East China Sea, Japan Yamaguti (1952), Yang et al. (2016)
A. micracantha* NR NR Gymnura micrura** WSA, WCA, WNA, ECA Nagasaki, East China Sea, Japan Yamaguti (1952)
A. micracantha* NR NR Telatrygon zugei WCP, WNP Nagasaki, East China Sea, Japan Yamaguti (1952)
A. microcephalum Alexander, 1953 USNPC 47852 NR Myliobatis californicus ENP, ECP Long Beach Harbor, California, USA Alexander (1953) 4‡
A. minus Tazerouti, Kechemir-Issad & Euzet, 2009 MNHN HEL 76, Th 180 MNHN HEL 77, Th 181, HEL 78, Th 182, HEL 79, Th 183; NHMUK 2009.2.10.1-2 Raja asterias Delaroche, 1809 ENA, MED Cap Djinet, Algérie Tazerouti et al. (2009) 2
A. minus NR NR Raja asterias ENA, MED Zemmouri El Bahri, Algérie Tazerouti et al. (2009)
A. minus NR NR Raja asterias ENA, MED Bouharoun, Algérie Tazerouti et al. (2009)
A. minusculus Marques, Brooks & Barriga, 1997 MEPN 3030 MNHG 22099; HWML 39178, CNHE 3030 Urobatis tumbesensis (Chirichigno F. & McEachran, 1979) ECP Puerto Hualtaco, Provincia de El Oro, Ecuador Marques et al. (1997a) 1‡
A. monksi Marques, Brooks & Barriga, 1997 MEPN 3031 MNHG 22100; HWML 39179; CNHE 3031 Aetobatus narinari** WSA, WCA, WNA, ECA Puerto Jelí, Provincia de El Oro, Ecuador Marques et al. (1997a) 1‡
A. mooreae Campbell & Beveridge, 2002 SAM AHC 28209 SAM AHC 22665, 22718, 28265 Trygonorrhina fasciata WSP Northhaven, South Australia Campbell and Beveridge (2002)
A. mujibi Bilqees, 1980 NR SPUK 2001 (syntype) Mustelus manazo NIO, WCP, WNP Karachi Coast, Pakistan Bilqees (1980)
A. musculosum (Baer, 1948) Yamaguti, 1959 NR NR Pteroplatytrygon violacea ENP, ECP, ESP, WSA, WCA, WNA, ENA, MED, ECA, ESA, WIO, NIO, EIO, WSP, WCP, WNP New Zealand Baer (1948), Euzet (1959), Yamaguti (1959a),Williams (1969)
A. myliomaculata Srivastav, Shweta & Noopur, 1995 DZCJ NR Aetomylaeus maculatus (Gray, 1834) NIO, WCP, WNP Madras Coast, India Srivastav et al. (1995)
A. nanogravidum Zschoche, Caira & Fyler, 2011 QM G232166 QM G232167–G23217, cross sections QM G232171, G23217; USNPC 104103); LRP 7480–7483, cross sections LRP 7486–7491, SEM LRP 7484–7485), egg mounts LRP 7492–7493 Pastinachus ater (Macleay, 1883) WIO, NIO, EIO, WSP, WCP Gulf of Carpentaria off Weipa, Queensland, Australia. Zschoche et al. (2011) 1
A. nicoyaense Brooks & McCorquodale, 1995 USNPC 84477 USNPC 84388; MNHG 18255 Aetobatus narinari** WSA, WCA, WNA, ECA Punta Morales, Golfo de Nicoya, Costa Rica Brooks and McCorquodale (1995) 1‡
A. ningdense Yang, Sun, Zhi, Iwaki, Reyda & Yang, 2016 MPM 21226 MPM 21227, 21228; SYSU 20121113-1-3, 20141002-1-27 Hemitrygon akajei WNP Fuhai aquatic market, Ningde, Fujian Province, China Yang et al. (2016) 4
A. ningdense NR NR Hemitrygon akajei WNP Off Wanjichi aquatic wholesale market, Taizhou, Zhejiang Province, China Yang et al. (2016)
A. ningdense NR NR Hemitrygon akajei WNP 8th Seafood Market, Xiamen, Fujian Province, China Yang et al. (2016)
A. ningdense NR NR Hemitrygon akajei WNP Guanghai Port, Taishan, Guangdong Province, China Yang et al. (2016)
A. ningdense NR NR Hemitrygon akajei WNP Sanya Fishing Port, Sanya, Hainan Province, China Yang et al. (2016)
A. obuncus Marques, Brooks & Barriga, 1997 MEPN 3032 MNHG 22101; HWML 39180; CNHE 3032, 3167 Hypanus longus ECP Puerto Hualtaco, Provincia de El Oro, Ecuador Marques et al. (1997a) 6‡
A. ocallaghani Campbell & Beveridge, 2002 SAM AHC 28202 SAM AHC 28203 Aptychotrema vincentiana (Haacke, 1885 EIO Musgrave Shoal, South Australia Campbell and Beveridge (2002)
A. oceanharvestae Fyler, Caira & Jensen, 2009 QM 231345 QM G231346–G231347; USNPC 101957–101958; LRP 4317–4318; cross sections QM 231349, QM G231348; SEM LRP 4319–4320, 4327–4328, hologenophores LRP 4321, LRP 4322 Urogymnus acanthobothrium Last, White & Kyne, 2016 WSP, WCP Arafura Sea, east of Wessel Islands, Northern Territory, Australia. Fyler et al. (2009), Caira and Jensen (2017) 1
A. odonoghuei Campbell & Beveridge, 2002 SAM AHC 22699 SAM AHC 22699 Urolophus expansus EIO Holdfast Bay, South Australia Campbell and Beveridge (2002)
A. odonoghuei*† NR NR Urolophus lobatus McKay, 1966 EIO Esperance, Western Australia Campbell and Beveridge (2002)
A. olseni Dailey & Mudry, 1968 USNPC 71216 NR Pseudobatos productus (Ayres, 1854) ENP, ECP Newport Beach, California, USA Dailey and Mudry (1968) 2‡
A. olseni*† NR NR Pseudobatos planiceps (Garman, 1880) ECP, ESP Lima, Chorrillos, Peru Iannacone et al. (2011)
A. olseni*† NR NR Urobatis halleri (Cooper, 1863) ENP, ECP Anaheim Bay, California, USA Appy and Dailey (1973)
A. olseni*† NR NR Urobatis halleri ENP, ECP Puerto Peñasco, Sonora, Mexico Friggens and Brown (2005)
A. omanense Maleki, Malek & Palm, 2019 ZCUOK 117 ZCUOK 118–122; ZUTC Platy. 1330–1334, 1 SEM voucher ZUTC Platy. 1335 Gymnura cf. poecilura** NIO, EIO, WCP, WNP Chabahar coast, Gulf of Oman, Iran Maleki et al. (2019) 1
A. omanense* NR NR Gymnura cf. poecilura NIO, EIO, WCP, WNP Bandar Abbas, Persian Gulf, Iran Maleki et al. (2019)
A. parviuncinatum Young, 1954 USNPC 49095 NR Urobatis halleri ENP, ECP San Diego Bays, California, USA Young (1954) 8‡
A. parviuncinatum* NR NR Gymnura marmorata (Cooper, 1864) ECP San Diego Bays, California, USA Young (1954)
A. parviuncinatum NR NR Urobatis halleri ENP, ECP Puerto Peñasco, Sonora, Mexico Friggens and Brown (2005)
A. parvum Manger, 1972 NR NR Dipturus batis ENA Faxa Bay, Western coasts Iceland Manger (1972)
A. paulum Linton, 1890 NR USNPC 07683, 35882, 71351, 71352. Bathytoshia centroura WSA, WCA, WNA Woods Hole, Massachusetts, USA Linton (1890), Vardo-Zalik and Campbell (2011) 1(8,9,5)‡
A. paulum*† NR NR Raja eglanteria WCA, WNA Chesapeake Bay, Virginia, USA Campbell (1969)
A. paulum*† NR NR Hypanus americanus WSA, WCA, WNA Chesapeake Bay, Virginia, USA Campbell (1969)
A. pearsoni Williams, 1962 NR NR Orectolobus maculatus (Bonnaterre, 1788) EIO, WSP Hastings Point NSW, Australia Williams (1962),Campbell and Beveridge (2002)
A. persicum Maleki, Malek & Palm, 2019 ZCUOK 135 ZCUOK 136–137; ZUTC Platy. 1351–1352, 1 SEM voucher ZCUOK 142, 1 SEM voucher ZUTC Platy. 1353 Gymnura cf. poecilura** NIO, EIO, WCP, WNP Bandar Abbas, Persian Gulf, Iran Maleki et al. (2019) 1
A. peruviense Reyda, 2008 USNPC 99945 USNPC 99946; LRP 4108–4111 (including whole mounts and SEM specimens); MZUSP 6393a–6393b; MHNP 2335 Potamotrygon motoro (Müller & Henle, 1841) WSA, WCA Madre de Dios River at Boca Manu, Madre de Dios Department, Peru Reyda (2008) 1(8)
A. pichelinae Campbell & Beveridge, 2002 SAM AHC 28229 SAM AHC 28230 Myliobatis tenuicaudatus EIO, WSP Devonport, Tasmania Campbell and Beveridge (2002)
A. pichelinae NR NR Myliobatis tenuicaudatus EIO, WSP Bunbury, Western Australia Campbell and Beveridge (2002)
A. pintanensis Wang, 1984 NR NR Neotrygon kuhlii** WSP Fujian Province, China Wang (1984) 4
A. polytesticularis Wang & Yang, 2001 PRLXU NR Squalus sp.** ? Xiamen, Fujiari, China Wang and Yang (2001)
A. ponticum Léon-Borcea, 1934 NR NR Raja clavata ENA, MED, ECA, ESA, WIO Agigéa, Black Sea Léon-Borcéa (1934)
A. ponticum* NR NR Dasyatis pastinaca ENA, MED, ECA Agigéa, Black Sea Léon-Borcéa (1935)
A. popi Fyler, Caira & Jensen, 2009 QM G231350 QM G231351–G231352; USNPC 101959–101960; LRP 4323–4324; cross sections QM G231353; SEM LRP 4329–4330, 4325–4326, hologenophores LRP 4331, 4332 Urogymnus acanthobothrium WSP, WCP Arafura Sea, east of Wessel Islands, Northern Territory, Australia. Fyler et al. (2009), Caira and Jensen (2017) 2
A. ppdeleoni Zaragoza-Tapia, Pulido-Flores & Monks, 2020 CNHE 11253 CNHE 11254; HWML 216260 Hypanus dipterurus ECP Bahía de Chamela, Jalisco, Mexico Zaragoza-Tapia et al. (2020) 2
A. psammobati Carvajal & Goldstein, 1969 USNPC 71357 USNPC 71358 Psammobatis scobina (Philippi, 1857) ESP South Pacific Ocean, between Papudo and Talcahuano, Chile Carvajal-G. and Goldstein (1969) 5‡
A. psammobati*† NR CH-MHNJP 342a, 342b Sympterygia brevicaudata ECP, ESP Callao, Lima, Peru Tantaleán-Vidaurre (1991)
A. puertecitense Caira & Zahner, 2001 CNHE 4175 CNHE 4176; USNPC 90843; LRP 2105–2106 Heterodontus francisci ECP, ESP Puertecitos, Gulf of California, Mexico Caira and Zahner (2001) 4
A. puntarenasense Marques, Brooks & Monks, 1995 MNHG 20005 MNHG 20006–20007; HWML 38543, CNHE 4176. Hypanus longus ECP Punta Morales, Puntarenas Province, Costa Rica Marques et al. (1995) 2‡
A. quadripartitum Williams, 1968 NR NR Leucoraja naevus (Müller & Henle, 1841) ENA, MED, ECA North Sea, off Aberdeen Williams (1968)
A. quinonesi Mayes, Brooks & Thorson, 1978 USNPC 74804 USNPC 74805; HWML 74806 Potamotrygon magdalenae (Duméril, 1865) WCA Magdalena River, Cienaga Jobo, vicinity of San Cristobal, Bolivar, Colombia Mayes et al. (1978) 5‡
A. quinonesi*† NR NR Potamotrygon yepezi Castex & Castello, 1970 WCA Lake Maracaibo area near El Congo and Represa de Tule, Rio Cachiri, Zulia, Venezuela Brooks et al. (1981)
A. rajaebatis (Rudolphi, 1810) Euzet, 1959 NR NR Dipturus batis** ENA Mediterranean Sea Rudolphi (1810)
A. rajaebatis*† NR NR Dipturus oxyrinchus (Linnaeus, 1758) ENA, MED, ECA Sète, France Euzet (1959)
A. rajaebatis*† NR NR Rostroraja alba (Lacepède, 1803) ENA, MED, ECA, ESA, WIO Sète, France Euzet (1959)
A. rajaebatis*† NR NR Rostroraja alba ENA, MED, ECA, ESA, WIO Lacépède, France Euzet (1959)
A. rajaebatis NR NR Dipturus batis** ENA Sète, France Euzet (1959)
A. rajaebatis NR NR Dipturus batis** ENA Roscoff, France Euzet (1959)
A. rajivi Ghoshroy & Caira, 2001 CNHE 4038 CNHE 4039; HWML 15552; LRP 2055–2056; USNPC 90461 Hypanus dipterurus ECP Puertecitos, Gulf of California, Mexico Ghoshroy and Caira (2001) 2‡
A. ramiroi Ivanov, 2005 MACN-Pa 412/1-4 USNPC 92521 Potamotrygon motoro WSA, WCA Río Colastiné, Santa Fé, Argentina Ivanov (2005) 4
A. ramiroi NR NR Potamotrygon motoro WSA, WCA Río Coronda, Santa Fé, Argentina Ivanov (2005)
A. regoi Brooks, Mayes & Thorson, 1981 USNPC 75709 USNPC 75710; HWML 21012, 21013 Potamotrygon hystrix (Müller & Henle, 1841) WSA Orinoco River Delta, Orinoco River near Los Castillos, Venezuela Brooks et al. (1981) 5‡
A. regoi*† NR NR Potamotrygon falkneri Castex & Maciel, 1963 WSA Paraná River, Brazil Lacerda et al. (2008)
A. regoi*† NR NR Potamotrygon motoro WSA, WCA Paraná River, Brazil Lacerda et al. (2008)
A. rhinobati Alexander, 1953 USNPC 47858 USNPC 47859 Pseudobatos productus ENP, ECP Santa Monica Harbor, California, USA Alexander (1953) 9(5)‡
A. rhinobati NR NR Pseudobatos productus ENP, ECP Ocean Park Pier, California, USA Alexander (1953)
A. robertsoni Campbell & Beveridge, 2002 SAM AHC 28197 SAM AHC 22590, 22591, 22592, 22667, 22714 Trygonorrhina fasciata WSP Middleton, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR SAM AHC 28257 Pristiophorus cirratus EIO, WSP Port Stanvac, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR NR Aptychotrema vincentiana EIO North Haven, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR NR Aptychotrema vincentiana EIO Goolwa, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR NR Dentiraja cerva (Whitley, 1939) EIO, WSP Port Stanvac, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR NR Dentiraja cerva EIO, WSP Holdfast Bay, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR SAM AHC 28260 Urolophus bucculentus Macleay, 1884 EIO, WSP Rapid Head, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR SAM AHC 22699 Urolophus expansus EIO Holdfast Bay, South Australia Campbell and Beveridge (2002)
A. robertsoni*† NR SAM AHC 28256 Urolophus lobatus EIO Esperance, Western Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP Outer Harbour, South Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP North Haven, South Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP Port Stanvac, South Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP Goolwa, South Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP Port Vincent, South Australia Campbell and Beveridge (2002)
A. robertsoni NR NR Trygonorrhina fasciata WSP Queenscliff, Victoria, Australia Campbell and Beveridge (2002)
A. robustum Alexander, 1953 USNPC 47856 USNPC 47857 Pseudobatos productus ENP, ECP Long Beach Harbor, California, USA Alexander (1953) 4‡
A. robustum*† NR NR Pseudobatos planiceps ECP, ESP Trujillo, Peru Escalante-A. (1986)
A. rodmani Fyler, Caira & Jensen, 2009 QM G231354 QM G231355–G231357; USNPC 101961–101963; LRP 4333–4335; cross sections QM G231359 G231358); cross sections LRP 4564–4569, 4563), longitudinal sections 4560–4562, 4559, SEM LRP 4336–4339, hologenophores LRP 4340, 4341 Urogymnus acanthobothrium WSP, WCP Arafura Sea, east of Wessel Islands, Northern Territory, Australia. Fyler et al. (2009), Caira and Jensen (2017) 6
A. rohdei Campbell & Beveridge, 2002 SAM AHC 28233 SAM AHC 28234 Urolophus lobatus EIO Esperance, Western Australia Campbell and Beveridge (2002)
A. romanowi Fyler, Caira & Jensen, 2009 QM G231360 QM G231361–231363; USNPC 101964–101966; LRP 4342–4344; cross sections QM G231365, G231364); cross sections LRP 4351–4356, SEM LRP 4345–4348, hologenophores LRP 4350, 4349. Urogymnus acanthobothrium WSP, WCP Arafura Sea, east of Wessel Islands, Northern Territory, Australia. Fyler et al. (2009), Caira and Jensen (2017) 1
A. rotundum Subhapradha, 1955 NR NR Rhynchobatus djiddensis WIO, NIO Madras Coast, India Subhapradha (1955) 4
A. royi Caira & Burge, 2001 CNHE 4173 CNHE 4174; LRP 2104; USNPC 90842 Diplobatis ommata ECP Punta Arena, Gulf of California, Mexico Caira and Burge (2001) 1
A. royi NR NR Diplobatis ommata ECP Loreto, Gulfo of California, Mexico Caira and Burge (2001)
A. rubrum Bilqees, 1980 NR SPUK 2002 (syntype) Mustelus manazo NIO, WCP, WNP Karachi Coast, Pakistan Bilqees (1980)
A. saliki Fyler & Caira, 2006 MZUM (P) 149 USNPC 96418–96419; LRP 3836-3843 (including cross sections and SEM specimens); MZUM (P) 150; IPMB 77.14.07 Urogymnus polylepis NIO, WCP Off Kampung Abai, Kinabatangan River, Sabah, Malaysia Fyler and Caira (2006)
A. santarosaliense Caira & Zahner, 2001 CNHE 4177 CNHE 4178; USNPC 90844; LRP 2107 Heterodontus mexicanus Taylor & Castro-Aguirre, 1972 ECP, ESP Santa Rosalia, Gulf of California, Mexico Caira and Zahner (2001) 3
A. satyanarayanaraoi Sanaka, Vijaya Lakshmi & Hanumantha Rao, 1993 DZAUW NR Glaucostegus granulatus (Cuvier, 1829) NIO Waltair coast, India Sanaka et al. (1993)
A. schalli Vardo-Zalik & Campbell, 2011 USNPC 103820 USNPC 103821–103826 Mustelus canis WNA, WCA, WSA Gulf of Mexico Vardo-Zalik and Campbell (2011) 1
A. schalli* NR NR Mustelus norrisi Springer, 1939 WNA, WCA, WSA Gulf of Mexico Vardo-Zalik and Campbell (2011)
A. semnovesiculum Verma, 1928 ZIMC NR Pastinachus sephen NIO Allahabad (Ganges and Jumna), India Verma (1928)
A. semnovesiculum NR NR Pastinachus sephen** NIO Fog Bay, Timor Sea, North Australia Campbell and Beveridge (2002)
A. semnovesiculum NR NR Pastinachus sephen** NIO Nickol Bay, Western Australia Campbell and Beveridge (2002)
A. septentrionale Baer & Euzet, 1962 NR NR Dipturus batis ENA Atlantic, Nort Sea Baer and Euzet (1962), Baer (1948), Euzet (1959)
A. septentrionale* NR NR Dipturus oxyrinchus ENA, MED, ECA Atlantic, Nort Sea Williams (1969)
A. sinaloaensis Zaragoza-Tapia, Pulido-Flores & Monks, 2020 CNHE 11257 CNHE 11258; HWML 216262 Hypanus longus ECP La Puntilla, Mazatlán, Sinaloa, Mexico Zaragoza-Tapia et al. (2020) 2
A. soberoni Ghoshroy & Caira, 2001 CNHE 4040 CNHE 4041–4042; HWML 15548; LRP 2057–2059; USNPC 90462 Hypanus dipterurus ECP Puertecitos, Gulf of California, Mexico Ghoshroy and Caira (2001) 6‡
A. soberoni NR NR Hypanus dipterurus ECP Bahía de Los Angeles, Gulf of California, Mexico Ghoshroy and Caira (2001)
A. soniae Zaragoza-Tapia, Pulido-Flores, Violante-Gonzalez & Monks, 2019 CNHE 11136 CNHE 11137; HWML 139978; CHE P00081 Narcine entemedor ECP Bahía de Acapulco, Playa Las Hamacas, Guerrero, Mexico Zaragoza-Tapia et al. (2019) 2
A. southwelli Subhapradha, 1955 NR NR Rhinobatos schlegelii** Müller & Henle, 1841 WNP Madras Coast, India Subhapradha (1955)
A. sphaera Maleki, Malek & Palm, 2013 ZUTC 1298 ZUTC 1299–1307), SEM vouchers ZUTC 1308–1309; IPCAS C–641; ZMB E7560 Pastinachus cf. sephen** NIO Persian Gulf, Iran Maleki et al. (2013) 2
A. stefaniae Franzese & Ivanov, 2018 MACN-Pa 624 MACN-Pa 625/1–6, 626/1–3, 627/1, 628/1–2; IPCAS C-786; LRP 9403–9410 Discopyge tschudii Heckel, 1846 ESP, WSA Coastal waters off Mar Chiquita City, Buenos Aires Province Franzese and Ivanov (2018) 1
A. stefaniae NR NR Discopyge tschudii ESP, WSA Coastal waters off Villa Gesell, Argentina Franzese and Ivanov (2018)
A. stefaniae NR NR Discopyge tschudii ESP, WSA Off San Clemente del Tuyú, Argentina Franzese and Ivanov (2018)
A. stefaniae NR NR Discopyge tschudii ESP, WSA Off Camarones, Argentina Franzese and Ivanov (2018)
A. stevensi Campbell & Beveridge, 2002 SAM AHC 28198 SAM AHC 28199 Trygonorrhina fasciata WSP Marion Bay, South Australia Campbell and Beveridge (2002)
A. stevensi NR NR Trygonorrhina fasciata WSP Goolwa, South Australia Campbell and Beveridge (2002)
A. stevensi NR NR Trygonorrhina fasciata WSP Coorong, Australia Campbell and Beveridge (2002)
A. tasajerasi Brooks, 1977 USNPC 73961 USNPC 73962; HWML 20261 Styracura schmardae WCA Caribbean Sea, La Cienaga, Magdalena, Colombia Brooks (1977) 2‡
A. tasajerasi*† NR NR Hypanus guttatus (Bloch & Schneider, 1801) WSA, WCA Lake Maracaibo, Venezuela Mayes and Brooks (1981)
A. terezae Rego & Dias, 1976 CHIOC 31.215c CHIO 10.847, 10.994, 31.412a-b, 31.215a-b Potamotrygon motoro WSA, WCA Rio Salobra, Mato Grosso, Brazil Rêgo and Luna Dias (1976) 4‡
A. tetabuanense Reyda & Caira, 2006 MZUM (P) 184(h) USNPC 97470–97471; LRP 3869–3873 (including cross sections and SEM specimens); MZUM (P) 185(p)–186(p); IPMB 77.08.18 Pateobatis uarnacoides NIO, WCP Off Kampung Tetabuan, Sabah, Malaysia Reyda and Caira (2006) 2
A. thomasae Campbell & Beveridge, 2002 SAM AHC 28201 SAM AHC 22676 Aptychotrema vincentiana EIO Musgrave Shoal, South Australia Campbell and Beveridge (2002)
A. thomasae NR NR Aptychotrema vincentiana EIO Cowell, Australia Campbell and Beveridge (2002)
A. tortum (Linton, 1916) Baer & Euzet, 1962 NR NR Aetobatus narinari WSA, WCA, WNA, ECA Woods Hole, Massachusetts, USA Linton (1916) 3‡
A. tortum NR NR Aetobatus narinari WSA, WCA, WNA, ECA Caimare Chico, Gulf of Venezuela Mayes and Brooks (1981)
A. tortum NR USNPC 70494 Aetobatus narinari WSA, WCA, WNA, ECA Cape Haze Marine Laboratory, Sarasota, Florida. Campbell (1970)
A. triacis Yamaguti, 1952 NR NR Triakis scyllium Müller & Henle, 1839 WNP Hamazima, Mie, Japan Yamaguti (1952) 4
A. tripartitum Williams, 1969 NR NR Raja microocellata Montagu, 1818 ENA, ECA English Channel, Plymouth Williams (1969)
A. ulmeri Vardo-Zalik & Campbell, 2011 USNPC 103830 USNPC 103831–103837, 103839, 103842, 103846 Raja texana WCA Gulf of Mexico Vardo-Zalik and Campbell (2011) 1
A. unilateralis Alexander, 1953 USNPC 47855 NR Myliobatis californicus ENP, ECP Long Beach Harbor, California, USA Alexander (1953) 7(2)‡
A. urogymni (Hornell, 1912) Southwell, 1925 NR NR Urogymnus asperrimus (Bloch & Schneider, 1801) ECA, WIO, NIO, EIO, WSP, WCP, WNP Gulf of Mannar, India Hornell (1912), Southwell (1925)
A. urolophi Schmidt, 1973 USNPC 72284 USNPC 72284 Trygonoptera testacea Müller & Henle, 1841 WSP Glenelg Beach near Adelaide, South Australia Schmidt (1973)
A. urolophi*† NR NR Urolophus paucimaculatus EIO, WSP Devonport, Tasmania Campbell and Beveridge (2002)
A. urotrygoni Brooks & Mayes, 1980 USNPC 75162 USNPC 75163; HWML 20917 Urobatis venezuelae Schultz, 1949 WCA Cartagena, Colombia Brooks and Mayes (1980) 2‡
A. urotrygoni*† NR NR Hypanus guttatus WSA, WCA Lake Maracaibo, Venezuela Mayes and Brooks (1981)
A. urotrygoni*† NR NR Hypanus guttatus WSA, WCA Isla Margarita, Venezuela Mayes and Brooks (1981)
A. vargasi Marques, Brooks & Monks, 1995 MNHG 20011 MNHG 20012–20013; HWML 38545 Hypanus longus ECP Punta Morales, Puntarenas Province, Costa Rica Marques et al. (1995) 2‡
A. vidali Zaragoza-Tapia, Pulido-Flores, Violante-Gonzalez & Monks, 2019 CNHE 11134 CNHE 11135; HWML 139979- 139981; CHE P00082 Narcine entemedor ECP Bahía de Acapulco, Playa Las Hamacas, Guerrero, Mexico Zaragoza-Tapia et al. (2019) 6
A. walkeri Campbell & Beveridge, 2002 SAM AHC 28219 SAM AHC 28220 Pastinachus sephen** NIO Nickol Bay, Western Australia Campbell and Beveridge (2002)
A. waltairensis Uma Maheswari, Sanaka, Vijaya Lakshmi & Hanumantha Rao, 1987 NR NR Himantura uarnak WIO, NIO, EIO, WCP Waltair coast, India Maheswari et al. (1987)
A. wedli Robinson, 1959 NR DMNZ 194b,c,d, 195–197 (syntype) Zearaja nasuta (Müller & Henle, 1841) WSP Petone Beach, New Zealand Robinson (1959)
A. wedli NR NR Zearaja nasuta WSP PortobeIlo, Otago Harbour, New Zealand Robinson (1959)
A. wedli NR NR Zearaja nasuta WSP South Island, off Lyttelton, New Zealand Campbell and Beveridge (2002)
A. westi Vardo-Zalik & Campbell, 2011 USNPC 103841 USNPC 103838, 103840, 103843–103845, 103847 Raja texana WCA Gulf of Mexico Vardo-Zalik and Campbell (2011) 1
A. woodsholei Baer, 1948 NR MNHG 40028 (syntype) Bathytoshia centroura WSA, WCA, WNA Woods Hole, Massachusetts, USA Baer (1948), Vardo-Zalik and Campbell (2011) 2(7)‡
A. woodsholei NR NR Bathytoshia centroura WSA, WCA, WNA Western North Atlantic Goldstein (1964)
A. xiamenensis Yang & Lin, 1994 NR NR Rhynchobatus djiddensis** WIO, NIO Xiamen, South Fujian, China Yang (1994)
A. zainali Fyler & Caira, 2006 MZUM (P) 151 USNPC 96420–96422; LRP 3844-3849 (including cross sections and SEM specimens); MZUM (P) 152–153; IPMB 77.14.08 Urogymnus polylepis NIO, WCP Off Kampung Abai, Kinabatangan River, Sabah, Malaysia Fyler and Caira (2006)
A. zapterycum Ostrowski de Nuñez, 1971 MACN-Pa 214/1 NR Zapteryx brevirostris (Müller & Henle, 1841) WSA Mar del Plata, Buenos Aires, Argentina Ostrowski de Núñez (1971) 2‡
A. zapterycum NR MACN-Pa 214/1-2, 214/4–5, 629/1, 630/1–3, 631/1–4, 632/1–4; IPCAS C-787; LRP 9411–9417 Zapteryx brevirostris WSA Coastal waters off Villa Gessel, Argentina Franzese and Ivanov (2018)
A. zapterycum NR NR Zapteryx brevirostris WSA La Lucila del Mar, Argentina Franzese and Ivanov (2018)
A. zapterycum NR NR Zapteryx brevirostris WSA Puerto Quequén, Argentina Franzese and Ivanov (2018)
A. zapterycum NR NR Zapteryx brevirostris WSA Puerto Pirámides, Argentina Franzese and Ivanov (2018)
A. zimmeri Fyler, Caira & Jensen, 2009 QM G231366 QM G231367–G231369; USNPC 101967–101969; LRP 4357–5358; cross sections QM G231371, G231370); cross sections LRP 4364–4366, SEM LRP 4359–4361, hologenophores LRP 4363, 4362 Urogymnus acanthobothrium WSP, WCP Arafura Sea, east of Wessel Islands, Northern Territory, Australia. Fyler et al. (2009), Caira and Jensen (2017) 1
A. zschokkei Baer, 1948 MHNG 88/39 NR Torpille (common name)** ? Naples, Italy Baer (1948)
A. zschokkei*† NR NR Torpedo marmorata ENA, MED, ECA, ESA Adriatic Sea, Mediterranean Sea Goldstein (1967)
A. zschokkei*† NR NR Torpedo torpedo ENA, MED, ECA Sète, France Euzet (1959)
A. zschokkei*† NR NR Torpedo torpedo ENA, MED, ECA Adriatic Sea, Mediterranean Sea Goldstein (1967)

The type localities where species of Acanthobothrium have been reported is shown in Figure 1. The currently known diversity of sharks comprises 517 species (34 families); of these, 19 species of sharks (eight families) have been reported to be parasitized by species of Acanthobothrium (Fig. 2). Eighteen of the 201 valid species have been described from sharks. The families of sharks that have the highest number of reports are Orectolobidae (three different species of Acanthobothrium), Heterodontidae (five species) and Triakidae (six species) (Fig. 2B). In contrast, currently known diversity of rays comprises 637 species (26 families); of these, 95 species (18 families) have been reported to be parasitized by species of Acanthobothrium (Fig. 3). Of the 201 valid species of Acanthobothrium, 182 have been described from rays. The families of rays that have the highest number of reports are Rajidae (20 species of Acanthobothrium) and Dasyatidae (70 species) (Fig. 3B).

Figure 2. 

Families of sharks: A number of species of sharks per family B number of species of sharks parasitized by species of Acanthobothrium. Note: The first number within parentheses corresponds to the number of species of shark that have been reported as hosts of Acanthobothrium and the second is the number of species that have been described from that Family C percentage of species of shark reported to be parasitized within the total number of families of sharks- note: Red color = parasitized; Blue color = not parasitized.

Species of Acanthobothrium are not evenly grouped in the different categories. In Category 1 there are 55 species, 44 in Category 2, 19 in Category 3, 37 in Category 4, 17 in Category 5, 14 in Category 6, four in Category 7, four in Category 8, and three in Category 9. Although there is a Category 10, species in that category also are in grouped with those in Category 8 because their characteristics are thought to fall into both categories (Table 1). The categories of four species of Acanthobothrium were classified as unknown (“?”) because the original descriptions do not have sufficient information for assignment in one of the ten categories (Table 1).

Figure 3. 

Families of rays: A number of species of rays per family B number of species of rays parasitized by species of Acanthobothrium. Note: The first number within parentheses corresponds to the number of species of ray that have been reported as hosts of Acanthobothrium and the second is the number of species that have been described from that Family C percentage of species of rays reported to be parasitized within the total number of families of rays- note: Red color = parasitized; Blue color = not parasitized.

Discussion

Currently, 517 species of sharks have been described worldwide with 3.7% (19 of the 517 species) have been reported as hosts for species of Acanthobothrium (Fig. 2C). In contrast, 637 species of rays have been described with 14.9% (95 of the 637 species) have been reported as hosts for species of Acanthobothrium (Fig. 3C). Estimates of cestode diversity in elasmobranchs discussed by Caira (2011) assumes that the fauna of cestodes of a species of elasmobranchs does not vary substantially across in its distribution. Knowledge of life cycles are essential in understanding the distribution of species of Acanthobothrium; however, for this study it is assumed that the distribution of adults of these parasites normally is limited to that of its normal definitive host. Thus, it is hypothesized that the limits of the distribution of the host limits the species of its parasites to the same biogeographic regions proposed for the distribution of elasmobranchs by Last et al. (2016b). It is recognized that an infected elasmobranch could move outside of the region where it has been designated, but until an extension to its distribution has been reported, it must be assumed that the normal distribution for each species of parasite also is that same designated region. The information in the table will be subject to future research, not forgetting that there is a lack of knowledge of the life cycle of the species of Acanthobothrium; a partial life cycle of a single species has been reported (Holland and Wilson 2009). Publication of molecular sequences for more species will provide new discoveries in this subject.

Figure 4. 

Number of species of Acanthobothrium reported from elasmobranchs in each biogeographic region (Last et al. 2016b).

The information in the Figures 1 and 4 indicates that there is an absence of reports from several regions of the world, such as ECA, ESA, WIO, ARC, and SOC. According to the percentages of species of elasmobranchs that have been reported as hosts of species of Acanthobothrium, we can infer that there are still many new species of Acanthobothrium to be discovered. In the GenBank database records, molecular sequences of only 16 of the 201 species of Acanthobothrium have been reported. However, more molecular information about species of Acanthobothrium is required for future analyzes, both for identification and life cycle studies; these would provide more solid information for delimiting distributions.

In Table 1, Acanthobothrium chilensis Rêgo, Vicente & Herrera, 1968, was included for reference, although it was described from a fish, Sarda chiliensis (Cuvier, 1832) (Perciformes: Scombridae) (see Rêgo et al. 1968). Extensive recent studies of this species of fish (Chero et al. 2016; Luque et al. 2016) failed to report A. chilensis; there is only the report by Rêgo et al. (1968). The report of the host for this species of Acanthobothrium likely is an accidental infection and not a normal host.

According to Fyler et al. (2009) and Franzese and Ivanov (2018), species of Acanthobothrium appear to exhibit oioxenous specificity for their elasmobranch hosts. In the present metadata analysis, for species exclusively in elasmobranchs, 83% of the species of Acanthobothrium show remarkable host specificity for their definitive host, and thus, should be considered to be an oioxenous species. In contrast, 34 of the 200 species (17%) of Acanthobothrium have been reported in more than one species of elasmobranch (Table 1). However, with the metadata analysis of the distribution of the hosts and the reports of the species of Acanthobothrium, 45 of the type specimens of Acanthobothrium require confirmation of the host (Table 1) because some appear to be problematic identifications and other hosts were reported as “cf.” or only as an unidentified member of a particular genus In addition, there are reports of species of Acanthobothrium that suggest misidentification of the parasites; these should reevaluated in future studies. To mention obvious cases, A. batailloni has been reported from the Mediterranean Sea and from the Pacific coast of Peru and Chile and A. brevissime has been reported from the Gulf of Mexico and the Pacific coast of Peru.

The categorical method developed by Ghoshroy and Caira (2001) was proposed in order to delimit the number of taxonomic comparisons when describing new species. Using the method of Ghoshroy and Caira (2001), which focused only on species from the Americas, Fyler and Caira (2006) later applied the same methodology to biodiversity data for species from other regions; those works are augmented by this study. Of the 201 known species of Acanthobothrium, 13 have been classified in more than one category (see category designations in Table 1) because some characteristics of those species overlap with those of more than one category (see descriptions found in Zschokke 1888; Linton 1890; Baer 1948; Alexander 1953; Euzet 1955; Riser 1955; Yamaguti 1959; Goldstein 1964; Williams 1969; Goldstein et al. 1969; Appy and Dailey 1973; Severino and Sarmiento 1979; Marques et al. 1997; Reyda 2008). This does not decrease the usefulness of the categorical method as a tool for the initial stages in identification.

Having more information, such as molecular sequences, could solve some problems in identification, such as the two cases mentioned above. A species of Acanthobothrium that has been assigned to more than one category suggests that the categories still need some refining, or it is an example of cryptic species that cannot be distinguished without molecular information. However, molecular information cannot replace morphological descriptions of species. One reason is the lack of material for sequencing of the vast majority of already-known species. Morphology also augments molecular data in studies of the phylogeny of platyhelminths (Zamparo et al. 2001; Littlewood 2008). A complete phylogenetic hypothesis based on total evidence (morphological and molecular data) such as that of Littlewood (2008) for any major group of cestodes is still distant. Until that time, a categorical method provides the easiest and most direct method for selection of a group of species similar to a new species of Acanthobothrium. This updated database includes the category designation for each species described to date will be an important tool for the future taxonomic studies.

Acknowledgements

The authors would like to thank to Luis García-Prieto (CNHE) for providing important bibliographic references and the Consejo Nacional de Ciencia y Tecnología (CONACYT) for a doctoral scholarship (no. 432427) to FZ-T.

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