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Corresponding author: Scott Monks ( monks.scott@gmail.com ) Academic editor: Boyko Georgiev
© 2020 Francisco Zaragoza-Tapia, Griselda Pulido-Flores, Scott L. Gardner, Scott Monks.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Zaragoza-Tapia F, Pulido-Flores G, Gardner SL, Monks S (2020) Host relationships and geographic distribution of species of Acanthobothrium Blanchard, 1848 (Onchoproteocephalidea, Onchobothriidae) in elasmobranchs: a metadata analysis. ZooKeys 940: 1-49. https://doi.org/10.3897/zookeys.940.46352
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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
Biodiversity, Elasmobranchii, Eucestoda, geographic distribution, rays, sharks
According to
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 (
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
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 (
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
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
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
For specimens deposited in a formal collection, acronyms are as follows:
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;
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;
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;
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;
MPM Meguro Parasitology Museum, Tokyo, Japan;
MZUM (P) Muzium Zoologi, Universiti Malaya, Kuala Lumpur, Malaysia;
NMNS National Museum of Natural Science, Taichung, Taiwan;
SBC Sarawak Biodiversity Center, Kuching, Sarawak, Malaysia;
SYSU School of Life Sciences, Sun Yat-sen University;
UAA Department of Zoology, University of Allahabad, Allahabad, India;
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;
The information obtained from the metadata analysis (Table
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
Species of Acanthobothrium | Ht | Nt, Pt or Va | Species of Host | Gd | Loc | Sou | Cd |
---|---|---|---|---|---|---|---|
A. adlardi Campbell & Beveridge, 2002 |
|
|
Pristiophorus cirratus (Latham, 1794) | EIO, WSP | Port Stanvac, South Australia |
|
4§ |
A. aetiobatidis (Shipley, 1900) Southwell, 1925 | NR | NR | Aetobatus narinari** (Euphrasen, 1790) | WSA, WCA, WNA, ECA | Lifu, Loyalty Islands |
|
6§ |
A. amazonensis Mayes, Brooks & Thorson, 1978 |
|
|
Potamotrygon circularis German, 1913 | WSA | Itacuari River, Brazil |
|
5‡ |
A. americanum Campbell, 1969 |
|
|
Hypanus americanus (Hildebrand & Schroeder, 1928) | WSA, WCA, WNA | Chesapeake Bay, Virginia, USA |
|
6‡ |
A. americanum† | NR | NR | Hypanus americanus | WSA, WCA, WNA | Isla Margarita, Venezuela |
|
– |
A. angelae Campbell & Beveridge, 2002 |
|
|
Hypnos monopterygius (Shaw, 1795) | EIO, WSP | Yarraville Shoals, South Australia |
|
5§ |
A. annapinkiensis Carvajal & Goldstein, 1971 |
|
NR | Zearaja chilensis (Guichenot, 1848) | ESP, WSA, | Anna Pink Bay, Chile |
|
2‡ |
A. arlenae Campbell & Beveridge, 2002 |
|
|
Aetobatus narinari** | WSA, WCA, WNA, ECA | Fog Bay, Timor Sea, North Australia |
|
6§ |
A. asnihae Fyler & Caira, 2006 | MZUM (P) 142 |
|
Urogymnus polylepis (Bleeker, 1852) | NIO, WCP | Off Kampung Abai, Kinabatangan River, Sabah, Malaysia |
|
1§ |
A. asrinae Maleki, Malek & Palm, 2015 |
|
|
Rhynchobatus cf. djiddensis** (Forsskå, 1775) | WIO, NIO | Persian Gulf, Iran |
|
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 |
|
1‡ |
A. australis Robinson, 1965 |
|
|
Squalus megalops (Macleay, 1881) | ENA, MED, ECA, ESA, WIO, EIO, WSP | Eden, New South Wales, Australia |
|
3§ |
A. australis† | NR |
|
Squalus megalops | ENA, MED, ECA, ESA, WIO, EIO, WSP | Beachport, South Australia |
|
– |
A. bajaensis Appy & Dailey, 1973 |
|
|
Heterodontus francisci (Girard, 1855) | ECP, ESP | San Quintin Bay, Baja California, Mexico |
|
4‡ |
A. bajaensis† | NR | NR | Heterodontus francisci | ECP, ESP | Newport Bay, California, USA |
|
– |
A. bartonae Campbell & Beveridge, 2002 |
|
NR | Rhynchobatus djiddensis** | WIO, NIO | Broome, Western Australia |
|
1§ |
A. batailloni Euzet, 1955 | NR | NR | Myliobatis aquila (Linnaeus, 1758) | ENA, MED, ECA, ESA, WIO | Mediterranean Sea, Gulfe du Lion |
|
7(2)‡ |
A. batailloni*† | NR |
|
Myliobatis chilensis** Philippi, 1892 | ESP | Antofagasta, Chile |
|
– |
A. batailloni*† | NR | NR | Myliobatis chilensis** | ESP | Coquimbo, Chile |
|
– |
A. batailloni*† | NR | NR | Myliobatis chilensis** | ESP | Trujillo, Peru |
|
– |
A. benedenii (Lönnberg, 1889) | NR | NR | Raja clavata Linnaeus, 1758 | ENA, MED, ECA, ESA, WIO | Mediterranean Sea |
|
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 |
|
– |
A. benedenii*† | NR | NR | Torpedo marmorata** Risso, 1810 | ENA, MED, ECA, ESA | Casablanca, Marruecos |
|
– |
A. bengalense Baer & Euzet, 1962 | NR | NR | Pastinachus sephen (Forsskål, 1775) | NIO | Nagapattinam, India |
|
4§ |
A. blairi Campbell & Beveridge, 2002 |
|
|
Dipturus whitleyi (Iredale, 1938) | EIO, WSP | Stanley, Tasmania |
|
3§ |
A. blairi† | NR | NR | Dipturus whitleyi | EIO, WSP | Spencer Gulf, South Australia |
|
– |
A. bobconniorum Fyler & Caira, 2010 |
|
|
Rhynchobatus laevis** (Bloch & Schneider, 1801) | NIO, WNP | Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia |
|
2¶ |
A. brachyacanthum Riser, 1955 |
|
NR | Raja stellulata (Gilbert, 1915) | ENP, ECP | Monterey Bay, California, USA |
|
2‡ |
A. brachyacanthum* | NR | NR | Beringraja binoculata** (Gilbert, 1855) | ENP, ECP | Monterey Bay, California, USA |
|
– |
A. brayi Campbell & Beveridge, 2002 |
|
|
Sutorectus tentaculatus (Peters, 1864) | EIO, WSP | Eastern Shoal, South Australia |
|
2§ |
A. brevissime Linton, 1909 |
|
NR | Hypanus say (Lesueur, 1817) | WSA, WCA, WNA | Dry Tortugas, Florida, USA |
|
2‡ |
A. brevissime*† | NR | NR | Raja eglanteria Bosc, 1800 | WCA, WNA | Gulf of Mexico, Chesapeake Bay, Virginia, USA |
|
– |
A. brevissime*† | NR |
|
Hypanus americanus | WSA, WCA, WNA | Gulf of Mexico, Chesapeake Bay, Virginia, USA |
|
– |
A. brevissime*† | NR | CH-MHNJP 727 | Myliobatis peruvianus** Garman, 1913 | ESP | Lima, Peru |
|
– |
A. brevissime† |
|
|
Hypanus say | WSA, WCA, WNA | Gulf of Mexico, Chesapeake Bay, Virginia, USA |
|
– |
A. bullardi Ghoshroy & Caira, 2001 | CNHE 4045 |
CNHE 4046–4047; LRP 2060–2062; |
Hypanus dipterurus (Jordan & Gilbert, 1880) | ECP | Bahía de Los Angeles, Gulf of California, Mexico |
|
2‡ |
A. bullardi† | NR | NR | Hypanus dipterurus | ECP | Puertecitos, Gulf of California, Mexico |
|
– |
A. bullardi† | NR | NR | Hypanus dipterurus | ECP | Santa Rosalia, Gulf of California, Mexico |
|
– |
A. cairae Vardo-Zalik & Campbell, 2011 |
|
|
Bathytoshia centroura (Mitchill, 1815) | WSA, WCA, WNA | Narragansett Bay off Sakonnet Point, Rhode Island, USA |
|
3¶ |
A. campbelli Marques, Brooks & Monks, 1995 | MNHG 20014 |
MNHG 20015–20016; |
Urotrygon chilensis (Günther, 1872) | ECP, ESP | Costa de Pajaros, Puntarenas, Costa Rica |
|
2‡ |
A. campbelli*† | NR | MEPN 3033 | Hypanus longus (Garman, 1880) | ECP | Puerto Huatulco, Provincia de El Oro, Ecuador |
|
– |
A. cannoni Campbell & Beveridge, 2002 |
|
|
Himantura uarnak (Gmelin, 1789) | WIO, NIO, EIO, WCP | Fog Bay, Timor Sea, North Australia |
|
4§ |
A. cartagenensis Brooks & Mayes, 1980 |
|
NR | Urobatis jamaicensis (Cuvier, 1816) | WCA, WNA | Cartagena, Colombia |
|
1¶ |
A. cartagenensis† | NR |
CNHE 9706; |
Urobatis jamaicensis | WCA, WNA | Ría Lagartos, Yucatán, Quintana Roo |
|
– |
A. cartagenensis† | NR |
CNHE 9706; |
Urobatis jamaicensis | WCA, WNA | Isla Contoy, Quintana Roo |
|
– |
A. cartagenensis† | NR |
CNHE 9706; |
Urobatis jamaicensis | WCA, WNA | Isla Cozumel, El Paso de los Cedros, Quintana Roo |
|
– |
A. cartagenensis† | NR |
CNHE 9706; |
Urobatis jamaicensis | WCA, WNA | Xcalak, Quintana Roo |
|
– |
A. cestraciontis (Yamaguti, 1934) | NR | NR | Heterodontus japonicus Miklouho-Maclay & Macleay, 1884 | WNP, WCP | Pacific Ocean, Japan |
|
4§ |
A. cestraciontis† | NR | NR | Sphyraena japonica** (Bloch &Schneider, 1801) | ? | Pacific Ocean, Japan |
|
– |
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 |
|
1¶ |
A. chengi Cornford, 1974 |
|
|
Bathytoshia lata (Garman, 1880) | ECP, ENA, MED, ECA, WIO, NIO, EIO, WSP, WCP, WNP | Oahu, Hawaii |
|
3§ |
A. chilensis Rego, Vincente & Herrera, 1968 |
|
NR | Sarda chiliensis** (Cuvier, 1832) | ? | Paita, Piúra, Peru |
|
3‡ |
A. chisholmae Campbell & Beveridge, 2002 |
|
|
Pastinachus sephen** | NIO | Nickol Bay, Western Australia |
|
2§ |
A. cimari Marques, Brooks & Monks, 1995 | MNHG 20017 |
MNHG 20018–20020; |
Hypanus longus | ECP | Punta Morales, Puntarenas Province, Costa Rica |
|
2‡ |
A. clarkeae Campbell & Beveridge, 2002 |
|
|
Urolophus paucimaculatus Dixon, 1969 | EIO, WSP | Queenscliff, Victoria, Australia |
|
1§ |
A. clarkeae*† | NR |
|
Urolophus cruciatus (Lacepède, 1804) | EIO, WSP | Devonport, Tasmania |
|
– |
A. clarkeae*† | NR |
|
Urolophus expansus McCulloch, 1916 | EIO | Beachport, South Australia |
|
– |
A. cleofanus Monks, Brooks & Lonce de Leon, 1996 | CNHE 2670 |
CNHE 2671; MNHG 38576; |
Hypanus longus | ECP | Chamela Bay, Jalisco, Mexico | ( |
3‡ |
A. colombianum Brooks & Mayes, 1980 |
|
|
Aetobatus narinari | WSA, WCA, WNA, ECA | Cartagena, Colombia |
|
9‡ |
A. confusum Baer & Euzet, 1962 | NR | NR | Neotrygon kuhlii** (Müller & Henle, 1841) | WSP | Indian Ocean, Sri Lanka |
|
5§ |
A. coquimbensis Carvajal & Jeges, 1980 |
|
NR | Myliobatis chilensis | ESP | Antofagasta, Chile |
|
2‡ |
A. coquimbensis† | NR | NR | Myliobatis chilensis | ESP | Coquimbo, Chile |
|
– |
A. coronatum (Rudolphi, 1819), Blanchard, 1848 | NR | NR | Dipturus batis (Linnaeus, 1758) | ENA | Mediterranean Sea, Italy |
|
4§ |
A. coronatum* | NR | NR | Scyliorhinus stellaris (Linnaeus, 1758) | ENA, MED, ECA | Mediterranean Sea, Italy |
|
– |
A. coronatum* | NR | NR | Torpedo marmorata | ENA, MED, ECA, ESA | Mediterranean Sea, Italy |
|
– |
A. coronatum* | NR | NR | Torpedo torpedo (Linnaeus, 1758) | ENA, MED, ECA | Mediterranean Sea, Italy |
|
– |
A. coronatum* | NR | NR | Dasyatis pastinaca (Linnaeus, 1758) | ENA, MED, ECA | Mediterranean Sea, Italy |
|
– |
A. coronatum*† | NR | NR | Hemitrygon akajei** (Müller & Henle, 1841) | WNP | Nakatsu, West Japan |
|
– |
A. coronatum*† | NR | NR | Aetobatus narinari** | WSA, WCA, WNA, ECA | Batavia, Java, Indonesia |
|
– |
A. coronatum*† | NR | NR | Scyliorhinus stellaris | ENA, MED, ECA | Sète, France |
|
– |
A. coronatum*† | NR | NR | Scyliorhinus stellaris | ENA, MED, ECA | Concarneau, France |
|
– |
A. coronatum*† | NR | NR | Mustelus mustelus (Linnaeus, 1758) | ENA, MED, ECA, ESA | Naples, Italy |
|
– |
A. coronatum*† | NR | NR | Scyliorhinus stellaris | ENA, MED, ECA | Cardigan Bay, UK |
|
– |
A. coronatum* | NR | NR | Carcharodon carcharias (Linnaeus, 1758) | MED | Mediterranean Sea |
|
– |
A. coronatum*† | NR | MNHG 40003, 40009 | Scyliorhinus canicula (Linnaeus, 1758) | ENA, MED, ECA | Naples, Italy |
|
– |
A. costarricense Marques, Brooks & Monks, 1995 | MNHG 20008 |
MNHG 20009–20010; |
Hypanus longus | ECP | Punta Morales, Puntarenas Province, Costa Rica |
|
2‡ |
A. costarricense† | NR | MEPN 3034 | Hypanus longus | ECP | Puerto Huatulco, Provincia de El Oro, Ecuador |
|
– |
A. crassicolle Wedl, 1855 | NR | MNHG 40014 88/77 | Dasyatis pastinaca | ENA, MED, ECA | Arcacho, Gironde, France |
|
3§ |
A. cribbi Campbell & Beveridge, 2002 |
|
|
Gymnura australis (Ramsay & Ogilby, 1886) | EIO, WSP, WCP | Gulf of Carpentaria, Northern Territory, Australia |
|
4§ |
A. dasi Ghoshroy & Caira, 2001 | CNHE 4043 |
CNHE 4044; |
Hypanus dipterurus | ECP | Puertecitos, Gulf of California, Mexico |
|
2‡ |
A. dasybati Yamaguti, 1934 | NR | NR | Hemitrygon akajei | WNP | Tarumi, Kobe, Japan |
|
4§ |
A. dasybati*† | NR | NR | Okamejei kenojei** (Müller & Henle, 1841) | WNP | Maisaka, Japan |
|
– |
A. dasybati*† | NR | NR | Urolophus sp.** (U. fuscus?) | ? | Hamazima, Mie, Japan |
|
– |
A. dighaensis Srivastava & Capoor, 1980 | UAA | NR | Pateobatis uarnacoides (Bleeker, 1852) | NIO, WCP | Digha, Orissa, India |
|
4§ |
A. dollyae Caira & Burge, 2001 | CNHE 4169 |
CNHE 4170; LRP 2097–2101; |
Diplobatis ommata (Jordan and Gilbert, 1890) | ECP | Bahía de Los Angeles, Gulf of California, Mexico |
|
1¶ |
A. dollyae† | NR | NR | Diplobatis ommata | ECP | Isla San Esteban, Gulf of California, Mexico |
|
– |
A. dollyae† | NR | NR | Diplobatis ommata | ECP | Punta Arena, Gulf of California, Mexico |
|
– |
A. dujardini van Beneden, 1850 | NR | NR | Raja clavata | ENA, MED, ECA, ESA, WIO | English Channel,Belgium |
|
2§ |
A. dujardini | NR | NR | Raja clavata | ENA, MED, ECA, ESA, WIO | English Channel, Belgium |
|
– |
A. dujardini*† | NR | NR | Raja brachyura** Lafont, 1871 | ENA, MED, ECA | Roscoff, France |
|
– |
A. dujardini*† | NR | NR | Raja montagui** Fowler, 1910 | ENA, MED | British Isles |
|
– |
A. dysbiotos (MacCallum, 1921) Williams, 1969 | NR | NR | Aetobatus narinari** | WSA, WCA, WNA, ECA | Batavia, Java, Indonesia |
|
4§ |
A. edmondsi Campbell & Beveridge, 2002 |
|
|
Parascyllium ferrugineum McCulloch, 1911 | EIO, WSP | Port Stanvac, South Australia |
|
5§ |
A. edmondsi† | NR | NR | Parascyllium ferrugineum | EIO, WSP | Holdfast Bay, South Australia |
|
– |
A. edmondsi† | NR | NR | Parascyllium ferrugineum | EIO, WSP | Esperance, Western Australia |
|
– |
A. edwardsi Williams, 1969 | NR | NR | Leucoraja fullonica (Linnaeus, 1758) | ENA, MED, ARC | West coast of Britain, United Kingdom |
|
2§ |
A. electricolum Brooks & Mayes, 1978 |
|
|
Narcine brasiliensis (Olfers, 1831) | WSA | Caribbean Sea, near Cartagena, Colombia |
|
9‡ |
A. elongatum Subhapradha, 1955 | NR | NR | Rhynchobatus djiddensis | WIO, NIO | Madras Coast, India |
|
4¶ |
A. etini Fyler & Caira, 2006 | MZUM (P) 145 |
|
Urogymnus polylepis | NIO, WCP | Off Kampung Abai, Kinabatangan River, Sabah, Malaysia |
|
8§ |
A. filicolle (Zschokke, 1888) Yamaguti, 1959 | NR | NR | Torpedo marmorata | ENA, MED, ECA, ESA | Mediterranean Sea |
|
1(8)¶ |
A. filicolle* | NR | NR | Torpedo torpedo | ENA, MED, ECA | Mediterranean Sea |
|
– |
A. floridensis Goldstein, 1964 |
|
NR | Raja eglanteria | WCA, WNA | Gulf of Mexico and Coast of Massachusetts |
|
8(10)‡ |
A. floridensis*† | NR |
|
Raja texana Chandler, 1921 | WCA | Gulf of Mexico |
|
– |
A. floridensis† | NR | NR | Raja eglanteria | WCA, WNA | Gulf of Mexico, Chesapeake Bay, Virginia, USA, USA |
|
– |
A. fogeli Goldstein, 1964 |
|
NR | Gymnura micrura (Bloch & Schneider, 1801) | WSA, WCA, WNA, ECA | Northeastern Gulf of Mexico, Florida |
|
1‡ |
A. fogeli† | NR | NR | Gymnura micrura | WSA, WCA, WNA, ECA | Isla Margarita, Venezuela |
|
– |
A. foulki Reyda & Caira, 2006 | MZUM (P) 168(h) |
|
Pateobatis uarnacoides | NIO, WCP | Off Kampung Tetabuan, Sabah, Malaysia |
|
1¶ |
A. franus Marques, Centritto & Stewart, 1997 | CNHE 3139 |
|
Narcine entemedor Jordan & Starks, 1895 | ECP | Cuajiniquil Beach, Gulf of Santa Helena, Guanacaste, Costa Rica |
|
5(8)‡ |
A. fylerae Maleki, Malek & Palm, 2015 |
|
|
Rhynchobatus cf. djiddensis** | WIO, NIO | Gulf of Oman, Iran |
|
1¶ |
A. gasseri Campbell & Beveridge, 2002 |
|
|
Pastinachus sephen** | NIO | Nickol Bay, Western Australia |
|
3§ |
A. gibsoni Campbell & Beveridge, 2002 |
|
NR | Rhynchobatus djiddensis** | WIO, NIO | Fog Bay, Timor Sea, North Australia |
|
3§ |
A. giganticum Sanaka, Lakshmi & Hanumantharao, 1993 | NR | NR | Gymnura micrura** | WSA, WCA, WNA, ECA | Waltair coast, India |
|
5§ |
A. gloveri Campbell & Beveridge, 2002 |
|
|
Trygonorrhina fasciata Müller & Henle, 1841 | WSP | Goolwa, South Australia |
|
2§ |
A. gnomus Reyda & Caira, 2006 | MZUM (P) 172(h) |
|
Pateobatis uarnacoides | NIO, WCP | Off Kampung Tetabuan, Sabah, Malaysia |
|
1¶ |
A. goldsteini Appy & Dailey, 1973 |
|
|
Platyrhinoidis triseriata (Jordan & Gilbert, 1880) | ENP, ECP | Seal Beach, California, USA |
|
5(9)‡ |
A. gonzalesmugaburoi Severino & Sarmiento, 1979 | CH-MHNJP 340 | CH-MHNJP 341, 341a, 341b | Myliobatis peruvianus | ESP | Callao, Lima, Peru |
|
7(6)¶ |
A. gracile Yamaguti, 1952 | NR | NR | Narke japonica (Temminck & Schlegel, 1850) | WNP | Tokushima, Japan |
|
3§ |
A. grandiceps Yamaguti, 1952 | MPM 22638 | NR | Telatrygon zugei (Müller & Henle, 1841) | WCP, WNP | East China Sea, Japan |
|
4§ |
A. grandiceps* | NR | NR | Hemitrygon akajei | WNP | East China Sea, Japan |
|
– |
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 |
|
2¶ |
A. halehae Maleki, Malek & Palm, 2019 | ZCUOK 127 |
ZCUOK 128–130; |
Gymnura cf. poecilura** (Shaw, 1804) | NIO, EIO, WCP, WNP | Chabahar coast, Gulf of Oman, Iran |
|
1¶ |
A. hanumantharaoi Rao, 1977 | NR | NR | Aetomylaeus nichofii (Bloch & Schneider, 1801) | NIO, EIO, WCP, WNP | Waltair coast, Benegal Bay, India |
|
4§ |
A. herdmani Southwell, 1912 | NR | NR | Neotrygon kuhlii** | WSP | Ceylon Pearl Bank, Sri Lanka |
|
3¶ |
A. heterodonti Drummond, 1937 | NR | NR | Heterodontus portusjacksoni (Meyer, 1793) | EIO, WSP | Lady Julia Perey Island, Victoria, Australia |
|
4§ |
A. heterodonti† | NR |
|
Heterodontus portusjacksoni | EIO, WSP | Derwent Estuary, Hobart, Tasmania |
|
– |
A. heterodonti† | NR | NR | Heterodontus portusjacksoni | EIO, WSP | Bunbury, Western Australia |
|
– |
A. himanturi Brooks, 1977 |
|
|
Styracura schmardae (Werner, 1904) | WCA | Caribbean Sea, La Cienaga, Magdalena, Colombia |
|
1‡ |
A. hispidum Riser, 1955 |
|
NR | Tetronarce californica (Ayres, 1855) | ENP, ECP, WNP | Monterey Bay, California, USA |
|
5‡ |
A. holorhini Alexander, 1953 |
|
|
Myliobatis californicus Grill, 1865 | ENP, ECP | Long Beach Harbor, California, USA |
|
3‡ |
A. holorhini*† | NR | CHIMTDC 542 | Myliobatis chilensis | ESP | Callao, Peru |
|
– |
A. hypanus Zaragoza-Tapia, Pulido-Flores & Monks, 2020 | CNHE 11255 |
CNHE 11256; |
Hypanus longus | ECP | La Puntilla, Mazatlán, Sinaloa, Mexico |
|
2¶ |
A. hypermekkolpos Fyler & Caira, 2010 |
|
|
Rhynchobatus laevis** | NIO, WNP | Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia |
|
1¶ |
A. icelandicum Manger, 1972 | NR | NR | Dipturus batis | ENA | Faxa Bay, Western coasts Iceland |
|
3§ |
A. ijimai Yoshida, 1917 | NR | MPM 22639 | Hemitrygon akajei | WNP | Tokyo, Japan |
|
4§ |
A. ijimai† | NR | NR | Hemitrygon akajei | WNP | East China Sea, Japan |
|
– |
A. inbiorium Marques, Centritto & Stewart, 1997 | CNHE 3137 |
|
Narcine entemedor | ECP | Cuajiniquil Beach, Gulf of Santa Helena, Guanacaste, Costa Rica |
|
5‡ |
A. incognita (MacCallum, 1921) Wardle & McLeod, 1952 | NR | NR | Dasyatis pastinaca | ENA, MED, ECA | New York Aquarium |
|
?¶ |
A. indicum (Subhapradha, 1955) | NR | NR | Narcine brasiliensis** | WSA | Madras Coast, India |
|
5§ |
A. intermedium Perrenoud, 1931 | NR | NR | Dasyatis pastinaca** | ENA, MED, ECA | Tauranga, New Zealand |
|
4§ |
A. jalalii Maleki, Malek & Palm, 2013 |
|
|
Pastinachus cf. sephen** | NIO | Gulf of Oman, Iran |
|
1¶ |
A. jamesi Maleki, Malek & Palm, 2015 |
|
ZMB E.7570; SEM voucher |
Rhynchobatus cf. djiddensis** | WIO, NIO | Persian Gulf, Iran |
|
1¶ |
A. janineae Maleki, Malek & Palm, 2015 |
|
|
Rhynchobatus cf. djiddensis** | WIO, NIO | Gulf of Oman, Iran |
|
1¶ |
A. jeanneae Fyler & Caira, 2010 |
|
|
Rhynchobatus laevis** | NIO, WNP | Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia |
|
1¶ |
A. jonesi Campbell & Beveridge, 2002 |
|
|
Dasyatis sp.** | ? | Cape Ford, North Australia |
|
6§ |
A. karachiense Bilqees, 1980 | NR |
|
Mustelus manazo Bleeker, 1855 | NIO, WCP, WNP | Karachi Coast, Pakistan |
|
4§ |
A. kurdistanense Maleki, Malek & Palm, 2019 | ZCUOK 122 |
ZCUOK 123–127; |
Gymnura cf. poecilura** | NIO, EIO, WCP, WNP | Chabahar coast, Gulf of Oman, Iran |
|
1¶ |
A. larsoni Reyda & Caira, 2006 | MZUM (P) 176(h) |
|
Pateobatis uarnacoides | NIO, WCP | Off Kampung Tetabuan, Sabah, Malaysia |
|
1¶ |
A. lasti Campbell & Beveridge, 2002 |
|
|
Rhynchobatus djiddensis** | WIO, NIO | Broome, Western Australia |
|
2§ |
A. latum Yamaguti, 1952 | MPM 22637 | NR | Hemitrygon akajei | WNP | Sea of Ariake, Kyusyu, Japan |
|
4§ |
A. laurenbrownae Campbell & Beveridge, 2002 |
|
|
Pastinachus sephen | NIO | Nickol Bay, Western Australia |
|
1§ |
A. lentiginosum Vardo-Zalik & Campbell, 2011 |
|
|
Pseudobatos lentiginosus (Garman, 1880) | WCA, WNA | Gulf of Mexico |
|
1¶ |
A. lepidum Reyda & Caira, 2006 | MZUM (P) 181(h) |
|
Pateobatis uarnacoides | NIO, WCP | Off Kampung Tetabuan, Sabah, Malaysia |
|
1¶ |
A. lilium Baer & Euzet, 1962 | NR | NR | Dasyatis sp.** | ? | Ceylon Pearl Bank, Sri Lanka |
|
2§ |
A. lineatum Campbell, 1969 |
|
|
Hypanus americanus | WSA, WCA, WNA | Chesapeake Bay, Virginia, USA |
|
1‡ |
A. lintoni Goldstein, Henson & Schlicht, 1968 |
|
|
Narcine brasiliensis** | WSA | Gulf of Mexico, Texas, USA |
|
1(8,9,5)‡ |
A. lintoni† | NR |
|
Narcine brasiliensis** | WSA | Gulf of Mexico, Florida, USA |
|
– |
A. longipedunculata Meheswari, Sanaka, Lakshmi & Rao, 1985 | NR | NR | Himantura uarnak | WIO, NIO, EIO, WCP | Waltair coast, India |
|
6§ |
A. lusarmientoi Severino & Verano, 1980 | CH-MHNJP 342 | CH-MHNJP 343, 343a | Sympterygia brevicaudata (Cope, 1877) | ECP, ESP | Callao, Lima, Peru |
|
7¶ |
A. macracanthum Southwell, 1925 | NR | NR | Urogymnus sp.** | ? | Madras Coast, India |
|
6§ |
A. macrocephalum Wang & Yang, 2001 | MPM 21231 | NR | Hemitrygon akajei | WNP | Xiamen, Fujiari, China |
|
4§ |
A. macrocephalum† | MPM 21231 | MPM 21232; SYSU 20140620-1-7 | Hemitrygon akajei | WNP | Off Guanghai Port, Guangdong, China |
|
– |
A. macrocephalum† | NR | NR | Hemitrygon akajei | WNP | Sanya Fishing Port, Sanya, Hainan, China |
|
– |
A. maculatum Riser, 1955 |
|
NR | Myliobatis californicus | ENP, ECP | Monterey Bay, California, USA |
|
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 |
|
4§ |
A. makranense Maleki, Malek & Palm, 2019 | ZCUOK 130 |
ZCUOK 131–135; |
Gymnura cf. poecilura** | NIO, EIO, WCP, WNP | Chabahar coast, Gulf of Oman, Iran |
|
1¶ |
A. manteri Hassan, 1983 | IHAHE S1051/A | IHAHE S1051/B | Pastinachus sephen** | NIO | Mediterranean Sea, Egypt |
|
5§ |
A. margieae Fyler, 2011 | NMNS 6356–001 |
NMNS 6356–002, 6356–003, 6356–004, 6356–005, 6356–006, 6356–007; LRP 7468–7477; |
Orectolobus japonicus Regan, 1906 | WNP, WCP | Off Penghu Island, East China Sea, Magong, Taiwan |
|
8¶ |
A. marplatensis Ivanov & Campbell, 1998 | MLP 4025 |
MLP 4026; USNMPC 87475; |
Atlantoraja castelnaui (Miranda Ribeiro, 1907) | WSA | Mar del Plata, Buenos Aires, Argentina |
|
1‡ |
A. marquesi Rodríguez-Ibarra, Pulido-Flores, Violante-González & Monks, 2018 | CNHE 10554 |
CNHE 10555, 10556; |
Aetobatus cf. narinari** | WSA, WCA, WNA, ECA | Laguna de Términos, Ciudad del Carmen, Campeche, Mexico |
|
3¶ |
A. marquesi† | NR | NR | Aetobatus cf. narinari** | WSA, WCA, WNA, ECA | Champotón, Campeche, Mexico |
|
– |
A. martini Campbell & Beveridge, 2002 |
|
|
Myliobatis tenuicaudatus Hector, 1877 | EIO, WSP | Bunbury, Western Australia |
|
1§ |
A. maryanskii Caira & Burge, 2001 | CNHE 4171 |
CNHE 4172; LRP 2012, 2013; |
Diplobatis ommata | ECP | Loreto, Gulfo of California, Mexico |
|
5¶ |
A. marymichaelorum Twohig, Caira & Fyler, 2008 | MZUM(P) 699(H) |
MZUM(P) 700(P)–702(P); SBC P–00028; |
Brevitrygon walga (Müller & Henle, 1841) | NIO | Off Sematan, Sarawak, Malaysia |
|
1¶ |
A. marymichaelorum | NR | NR | Brevitrygon walga | NIO | Off Mukah, Sarawak, Malaysia. |
|
– |
A. masnihae Fyler & Caira, 2006 | MZUM (P) 147 |
|
Urogymnus polylepis | NIO, WCP | Kampung Abai, Kinabatangan River, Sabah, Malaysia |
|
2§ |
A. mathiasi Euzet, 1959 | NR | NR | Mustelus mustelus | ENA, MED, ECA, ESA | Sète, France |
|
1§ |
A. mathiasi* | NR | NR | Mustelus canis (Mitchill, 1815) | WNA, WCA, WSA | Sète, France |
|
– |
A. matttaylori Fyler & Caira, 2010 |
|
Hologenophore |
Rhynchobatus laevis** | NIO, WNP | Gove Harbor, Gulf of Carpentaria, Northern Territory, Australia |
|
4¶ |
A. micracantha Yamaguti, 1952 | NR | MPM 22635, 22636 | Hemitrygon akajei | WNP | Nagasaki, East China Sea, Japan |
|
4§ |
A. micracantha* | NR | NR | Gymnura micrura** | WSA, WCA, WNA, ECA | Nagasaki, East China Sea, Japan |
|
– |
A. micracantha* | NR | NR | Telatrygon zugei | WCP, WNP | Nagasaki, East China Sea, Japan |
|
– |
A. microcephalum Alexander, 1953 |
|
NR | Myliobatis californicus | ENP, ECP | Long Beach Harbor, California, USA |
|
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; |
Raja asterias Delaroche, 1809 | ENA, MED | Cap Djinet, Algérie |
|
2¶ |
A. minus† | NR | NR | Raja asterias | ENA, MED | Zemmouri El Bahri, Algérie |
|
– |
A. minus† | NR | NR | Raja asterias | ENA, MED | Bouharoun, Algérie |
|
– |
A. minusculus Marques, Brooks & Barriga, 1997 | MEPN 3030 |
MNHG 22099; |
Urobatis tumbesensis (Chirichigno F. & McEachran, 1979) | ECP | Puerto Hualtaco, Provincia de El Oro, Ecuador |
|
1‡ |
A. monksi Marques, Brooks & Barriga, 1997 | MEPN 3031 |
MNHG 22100; |
Aetobatus narinari** | WSA, WCA, WNA, ECA | Puerto Jelí, Provincia de El Oro, Ecuador |
|
1‡ |
A. mooreae Campbell & Beveridge, 2002 |
|
|
Trygonorrhina fasciata | WSP | Northhaven, South Australia |
|
2§ |
A. mujibi Bilqees, 1980 | NR |
|
Mustelus manazo | NIO, WCP, WNP | Karachi Coast, Pakistan |
|
?¶ |
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 |
|
4§ |
A. myliomaculata Srivastav, Shweta & Noopur, 1995 | DZCJ | NR | Aetomylaeus maculatus (Gray, 1834) | NIO, WCP, WNP | Madras Coast, India |
|
4§ |
A. nanogravidum Zschoche, Caira & Fyler, 2011 |
|
|
Pastinachus ater (Macleay, 1883) | WIO, NIO, EIO, WSP, WCP | Gulf of Carpentaria off Weipa, Queensland, Australia. |
|
1¶ |
A. nicoyaense Brooks & McCorquodale, 1995 |
|
|
Aetobatus narinari** | WSA, WCA, WNA, ECA | Punta Morales, Golfo de Nicoya, Costa Rica |
|
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 |
|
4¶ |
A. ningdense† | NR | NR | Hemitrygon akajei | WNP | Off Wanjichi aquatic wholesale market, Taizhou, Zhejiang Province, China |
|
– |
A. ningdense† | NR | NR | Hemitrygon akajei | WNP | 8th Seafood Market, Xiamen, Fujian Province, China |
|
– |
A. ningdense† | NR | NR | Hemitrygon akajei | WNP | Guanghai Port, Taishan, Guangdong Province, China |
|
– |
A. ningdense† | NR | NR | Hemitrygon akajei | WNP | Sanya Fishing Port, Sanya, Hainan Province, China |
|
– |
A. obuncus Marques, Brooks & Barriga, 1997 | MEPN 3032 |
MNHG 22101; |
Hypanus longus | ECP | Puerto Hualtaco, Provincia de El Oro, Ecuador |
|
6‡ |
A. ocallaghani Campbell & Beveridge, 2002 |
|
|
Aptychotrema vincentiana (Haacke, 1885 | EIO | Musgrave Shoal, South Australia |
|
2§ |
A. oceanharvestae Fyler, Caira & Jensen, 2009 |
|
|
Urogymnus acanthobothrium Last, White & Kyne, 2016 | WSP, WCP | Arafura Sea, east of Wessel Islands, Northern Territory, Australia. |
|
1¶ |
A. odonoghuei Campbell & Beveridge, 2002 |
|
|
Urolophus expansus | EIO | Holdfast Bay, South Australia |
|
1§ |
A. odonoghuei*† | NR | NR | Urolophus lobatus McKay, 1966 | EIO | Esperance, Western Australia |
|
– |
A. olseni Dailey & Mudry, 1968 |
|
NR | Pseudobatos productus (Ayres, 1854) | ENP, ECP | Newport Beach, California, USA |
|
2‡ |
A. olseni*† | NR | NR | Pseudobatos planiceps (Garman, 1880) | ECP, ESP | Lima, Chorrillos, Peru |
|
– |
A. olseni*† | NR | NR | Urobatis halleri (Cooper, 1863) | ENP, ECP | Anaheim Bay, California, USA |
|
– |
A. olseni*† | NR | NR | Urobatis halleri | ENP, ECP | Puerto Peñasco, Sonora, Mexico |
|
– |
A. omanense Maleki, Malek & Palm, 2019 | ZCUOK 117 |
ZCUOK 118–122; |
Gymnura cf. poecilura** | NIO, EIO, WCP, WNP | Chabahar coast, Gulf of Oman, Iran |
|
1¶ |
A. omanense* | NR | NR | Gymnura cf. poecilura | NIO, EIO, WCP, WNP | Bandar Abbas, Persian Gulf, Iran |
|
– |
A. parviuncinatum Young, 1954 |
|
NR | Urobatis halleri | ENP, ECP | San Diego Bays, California, USA |
|
8‡ |
A. parviuncinatum* | NR | NR | Gymnura marmorata (Cooper, 1864) | ECP | San Diego Bays, California, USA |
|
– |
A. parviuncinatum† | NR | NR | Urobatis halleri | ENP, ECP | Puerto Peñasco, Sonora, Mexico |
|
– |
A. parvum Manger, 1972 | NR | NR | Dipturus batis | ENA | Faxa Bay, Western coasts Iceland |
|
6§ |
A. paulum Linton, 1890 | NR |
|
Bathytoshia centroura | WSA, WCA, WNA | Woods Hole, Massachusetts, USA |
|
1(8,9,5)‡ |
A. paulum*† | NR | NR | Raja eglanteria | WCA, WNA | Chesapeake Bay, Virginia, USA |
|
– |
A. paulum*† | NR | NR | Hypanus americanus | WSA, WCA, WNA | Chesapeake Bay, Virginia, USA |
|
– |
A. pearsoni Williams, 1962 | NR | NR | Orectolobus maculatus (Bonnaterre, 1788) | EIO, WSP | Hastings Point NSW, Australia |
|
1§ |
A. persicum Maleki, Malek & Palm, 2019 | ZCUOK 135 |
ZCUOK 136–137; |
Gymnura cf. poecilura** | NIO, EIO, WCP, WNP | Bandar Abbas, Persian Gulf, Iran |
|
1¶ |
A. peruviense Reyda, 2008 |
|
|
Potamotrygon motoro (Müller & Henle, 1841) | WSA, WCA | Madre de Dios River at Boca Manu, Madre de Dios Department, Peru |
|
1(8)¶ |
A. pichelinae Campbell & Beveridge, 2002 |
|
|
Myliobatis tenuicaudatus | EIO, WSP | Devonport, Tasmania |
|
4§ |
A. pichelinae† | NR | NR | Myliobatis tenuicaudatus | EIO, WSP | Bunbury, Western Australia |
|
– |
A. pintanensis Wang, 1984 | NR | NR | Neotrygon kuhlii** | WSP | Fujian Province, China |
|
4¶ |
A. polytesticularis Wang & Yang, 2001 |
|
NR | Squalus sp.** | ? | Xiamen, Fujiari, China |
|
4§ |
A. ponticum Léon-Borcea, 1934 | NR | NR | Raja clavata | ENA, MED, ECA, ESA, WIO | Agigéa, Black Sea |
|
?¶ |
A. ponticum* | NR | NR | Dasyatis pastinaca | ENA, MED, ECA | Agigéa, Black Sea |
|
– |
A. popi Fyler, Caira & Jensen, 2009 |
|
|
Urogymnus acanthobothrium | WSP, WCP | Arafura Sea, east of Wessel Islands, Northern Territory, Australia. |
|
2¶ |
A. ppdeleoni Zaragoza-Tapia, Pulido-Flores & Monks, 2020 | CNHE 11253 |
CNHE 11254; |
Hypanus dipterurus | ECP | Bahía de Chamela, Jalisco, Mexico |
|
2¶ |
A. psammobati Carvajal & Goldstein, 1969 |
|
|
Psammobatis scobina (Philippi, 1857) | ESP | South Pacific Ocean, between Papudo and Talcahuano, Chile |
|
5‡ |
A. psammobati*† | NR | CH-MHNJP 342a, 342b | Sympterygia brevicaudata | ECP, ESP | Callao, Lima, Peru |
|
– |
A. puertecitense Caira & Zahner, 2001 | CNHE 4175 |
CNHE 4176; |
Heterodontus francisci | ECP, ESP | Puertecitos, Gulf of California, Mexico |
|
4¶ |
A. puntarenasense Marques, Brooks & Monks, 1995 | MNHG 20005 |
MNHG 20006–20007; |
Hypanus longus | ECP | Punta Morales, Puntarenas Province, Costa Rica |
|
2‡ |
A. quadripartitum Williams, 1968 | NR | NR | Leucoraja naevus (Müller & Henle, 1841) | ENA, MED, ECA | North Sea, off Aberdeen |
|
2§ |
A. quinonesi Mayes, Brooks & Thorson, 1978 |
|
|
Potamotrygon magdalenae (Duméril, 1865) | WCA | Magdalena River, Cienaga Jobo, vicinity of San Cristobal, Bolivar, Colombia |
|
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 |
|
– |
A. rajaebatis (Rudolphi, 1810) Euzet, 1959 | NR | NR | Dipturus batis** | ENA | Mediterranean Sea |
|
5§ |
A. rajaebatis*† | NR | NR | Dipturus oxyrinchus (Linnaeus, 1758) | ENA, MED, ECA | Sète, France |
|
– |
A. rajaebatis*† | NR | NR | Rostroraja alba (Lacepède, 1803) | ENA, MED, ECA, ESA, WIO | Sète, France |
|
– |
A. rajaebatis*† | NR | NR | Rostroraja alba | ENA, MED, ECA, ESA, WIO | Lacépède, France |
|
– |
A. rajaebatis† | NR | NR | Dipturus batis** | ENA | Sète, France |
|
– |
A. rajaebatis† | NR | NR | Dipturus batis** | ENA | Roscoff, France |
|
– |
A. rajivi Ghoshroy & Caira, 2001 | CNHE 4038 |
CNHE 4039; |
Hypanus dipterurus | ECP | Puertecitos, Gulf of California, Mexico |
|
2‡ |
A. ramiroi Ivanov, 2005 |
|
|
Potamotrygon motoro | WSA, WCA | Río Colastiné, Santa Fé, Argentina |
|
4¶ |
A. ramiroi† | NR | NR | Potamotrygon motoro | WSA, WCA | Río Coronda, Santa Fé, Argentina |
|
– |
A. regoi Brooks, Mayes & Thorson, 1981 |
|
|
Potamotrygon hystrix (Müller & Henle, 1841) | WSA | Orinoco River Delta, Orinoco River near Los Castillos, Venezuela |
|
5‡ |
A. regoi*† | NR | NR | Potamotrygon falkneri Castex & Maciel, 1963 | WSA | Paraná River, Brazil |
|
– |
A. regoi*† | NR | NR | Potamotrygon motoro | WSA, WCA | Paraná River, Brazil |
|
– |
A. rhinobati Alexander, 1953 |
|
|
Pseudobatos productus | ENP, ECP | Santa Monica Harbor, California, USA |
|
9(5)‡ |
A. rhinobati† | NR | NR | Pseudobatos productus | ENP, ECP | Ocean Park Pier, California, USA |
|
– |
A. robertsoni Campbell & Beveridge, 2002 |
|
|
Trygonorrhina fasciata | WSP | Middleton, South Australia |
|
3§ |
A. robertsoni*† | NR |
|
Pristiophorus cirratus | EIO, WSP | Port Stanvac, South Australia |
|
– |
A. robertsoni*† | NR | NR | Aptychotrema vincentiana | EIO | North Haven, South Australia |
|
– |
A. robertsoni*† | NR | NR | Aptychotrema vincentiana | EIO | Goolwa, South Australia |
|
– |
A. robertsoni*† | NR | NR | Dentiraja cerva (Whitley, 1939) | EIO, WSP | Port Stanvac, South Australia |
|
– |
A. robertsoni*† | NR | NR | Dentiraja cerva | EIO, WSP | Holdfast Bay, South Australia |
|
– |
A. robertsoni*† | NR |
|
Urolophus bucculentus Macleay, 1884 | EIO, WSP | Rapid Head, South Australia |
|
– |
A. robertsoni*† | NR |
|
Urolophus expansus | EIO | Holdfast Bay, South Australia |
|
– |
A. robertsoni*† | NR |
|
Urolophus lobatus | EIO | Esperance, Western Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | Outer Harbour, South Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | North Haven, South Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | Port Stanvac, South Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | Goolwa, South Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | Port Vincent, South Australia |
|
– |
A. robertsoni† | NR | NR | Trygonorrhina fasciata | WSP | Queenscliff, Victoria, Australia |
|
– |
A. robustum Alexander, 1953 |
|
|
Pseudobatos productus | ENP, ECP | Long Beach Harbor, California, USA |
|
4‡ |
A. robustum*† | NR | NR | Pseudobatos planiceps | ECP, ESP | Trujillo, Peru |
|
– |
A. rodmani Fyler, Caira & Jensen, 2009 |
|
|
Urogymnus acanthobothrium | WSP, WCP | Arafura Sea, east of Wessel Islands, Northern Territory, Australia. |
|
6¶ |
A. rohdei Campbell & Beveridge, 2002 |
|
|
Urolophus lobatus | EIO | Esperance, Western Australia |
|
1§ |
A. romanowi Fyler, Caira & Jensen, 2009 |
|
|
Urogymnus acanthobothrium | WSP, WCP | Arafura Sea, east of Wessel Islands, Northern Territory, Australia. |
|
1¶ |
A. rotundum Subhapradha, 1955 | NR | NR | Rhynchobatus djiddensis | WIO, NIO | Madras Coast, India |
|
4¶ |
A. royi Caira & Burge, 2001 | CNHE 4173 |
CNHE 4174; LRP 2104; |
Diplobatis ommata | ECP | Punta Arena, Gulf of California, Mexico |
|
1¶ |
A. royi† | NR | NR | Diplobatis ommata | ECP | Loreto, Gulfo of California, Mexico |
|
– |
A. rubrum Bilqees, 1980 | NR |
|
Mustelus manazo | NIO, WCP, WNP | Karachi Coast, Pakistan |
|
6§ |
A. saliki Fyler & Caira, 2006 | MZUM (P) 149 |
|
Urogymnus polylepis | NIO, WCP | Off Kampung Abai, Kinabatangan River, Sabah, Malaysia |
|
1§ |
A. santarosaliense Caira & Zahner, 2001 | CNHE 4177 |
CNHE 4178; |
Heterodontus mexicanus Taylor & Castro-Aguirre, 1972 | ECP, ESP | Santa Rosalia, Gulf of California, Mexico |
|
3¶ |
A. satyanarayanaraoi Sanaka, Vijaya Lakshmi & Hanumantha Rao, 1993 | DZAUW | NR | Glaucostegus granulatus (Cuvier, 1829) | NIO | Waltair coast, India |
|
4§ |
A. schalli Vardo-Zalik & Campbell, 2011 |
|
|
Mustelus canis | WNA, WCA, WSA | Gulf of Mexico |
|
1¶ |
A. schalli* | NR | NR | Mustelus norrisi Springer, 1939 | WNA, WCA, WSA | Gulf of Mexico |
|
– |
A. semnovesiculum Verma, 1928 | ZIMC | NR | Pastinachus sephen | NIO | Allahabad (Ganges and Jumna), India |
|
2§ |
A. semnovesiculum† | NR | NR | Pastinachus sephen** | NIO | Fog Bay, Timor Sea, North Australia |
|
– |
A. semnovesiculum† | NR | NR | Pastinachus sephen** | NIO | Nickol Bay, Western Australia |
|
– |
A. septentrionale Baer & Euzet, 1962 | NR | NR | Dipturus batis | ENA | Atlantic, Nort Sea |
|
3§ |
A. septentrionale* | NR | NR | Dipturus oxyrinchus | ENA, MED, ECA | Atlantic, Nort Sea |
|
– |
A. sinaloaensis Zaragoza-Tapia, Pulido-Flores & Monks, 2020 | CNHE 11257 |
CNHE 11258; |
Hypanus longus | ECP | La Puntilla, Mazatlán, Sinaloa, Mexico |
|
2¶ |
A. soberoni Ghoshroy & Caira, 2001 | CNHE 4040 |
CNHE 4041–4042; |
Hypanus dipterurus | ECP | Puertecitos, Gulf of California, Mexico |
|
6‡ |
A. soberoni† | NR | NR | Hypanus dipterurus | ECP | Bahía de Los Angeles, Gulf of California, Mexico |
|
– |
A. soniae Zaragoza-Tapia, Pulido-Flores, Violante-Gonzalez & Monks, 2019 | CNHE 11136 |
CNHE 11137; |
Narcine entemedor | ECP | Bahía de Acapulco, Playa Las Hamacas, Guerrero, Mexico |
|
2¶ |
A. southwelli Subhapradha, 1955 | NR | NR | Rhinobatos schlegelii** Müller & Henle, 1841 | WNP | Madras Coast, India |
|
1§ |
A. sphaera Maleki, Malek & Palm, 2013 |
|
|
Pastinachus cf. sephen** | NIO | Persian Gulf, Iran |
|
2¶ |
A. stefaniae Franzese & Ivanov, 2018 |
|
|
Discopyge tschudii Heckel, 1846 | ESP, WSA | Coastal waters off Mar Chiquita City, Buenos Aires Province |
|
1¶ |
A. stefaniae† | NR | NR | Discopyge tschudii | ESP, WSA | Coastal waters off Villa Gesell, Argentina |
|
– |
A. stefaniae† | NR | NR | Discopyge tschudii | ESP, WSA | Off San Clemente del Tuyú, Argentina |
|
– |
A. stefaniae† | NR | NR | Discopyge tschudii | ESP, WSA | Off Camarones, Argentina |
|
– |
A. stevensi Campbell & Beveridge, 2002 |
|
|
Trygonorrhina fasciata | WSP | Marion Bay, South Australia |
|
2§ |
A. stevensi† | NR | NR | Trygonorrhina fasciata | WSP | Goolwa, South Australia |
|
– |
A. stevensi† | NR | NR | Trygonorrhina fasciata | WSP | Coorong, Australia |
|
– |
A. tasajerasi Brooks, 1977 |
|
|
Styracura schmardae | WCA | Caribbean Sea, La Cienaga, Magdalena, Colombia |
|
2‡ |
A. tasajerasi*† | NR | NR | Hypanus guttatus (Bloch & Schneider, 1801) | WSA, WCA | Lake Maracaibo, Venezuela |
|
– |
A. terezae Rego & Dias, 1976 |
|
CHIO 10.847, 10.994, 31.412a-b, 31.215a-b | Potamotrygon motoro | WSA, WCA | Rio Salobra, Mato Grosso, Brazil |
|
4‡ |
A. tetabuanense Reyda & Caira, 2006 | MZUM (P) 184(h) |
|
Pateobatis uarnacoides | NIO, WCP | Off Kampung Tetabuan, Sabah, Malaysia |
|
2¶ |
A. thomasae Campbell & Beveridge, 2002 |
|
|
Aptychotrema vincentiana | EIO | Musgrave Shoal, South Australia |
|
2§ |
A. thomasae† | NR | NR | Aptychotrema vincentiana | EIO | Cowell, Australia |
|
– |
A. tortum (Linton, 1916) Baer & Euzet, 1962 | NR | NR | Aetobatus narinari | WSA, WCA, WNA, ECA | Woods Hole, Massachusetts, USA |
|
3‡ |
A. tortum† | NR | NR | Aetobatus narinari | WSA, WCA, WNA, ECA | Caimare Chico, Gulf of Venezuela |
|
– |
A. tortum† | NR |
|
Aetobatus narinari | WSA, WCA, WNA, ECA | Cape Haze Marine Laboratory, Sarasota, Florida. |
|
– |
A. triacis Yamaguti, 1952 | NR | NR | Triakis scyllium Müller & Henle, 1839 | WNP | Hamazima, Mie, Japan |
|
4¶ |
A. tripartitum Williams, 1969 | NR | NR | Raja microocellata Montagu, 1818 | ENA, ECA | English Channel, Plymouth |
|
2§ |
A. ulmeri Vardo-Zalik & Campbell, 2011 |
|
|
Raja texana | WCA | Gulf of Mexico |
|
1¶ |
A. unilateralis Alexander, 1953 |
|
NR | Myliobatis californicus | ENP, ECP | Long Beach Harbor, California, USA |
|
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 |
|
?¶ |
A. urolophi Schmidt, 1973 |
|
|
Trygonoptera testacea Müller & Henle, 1841 | WSP | Glenelg Beach near Adelaide, South Australia |
|
1§ |
A. urolophi*† | NR | NR | Urolophus paucimaculatus | EIO, WSP | Devonport, Tasmania |
|
– |
A. urotrygoni Brooks & Mayes, 1980 |
|
|
Urobatis venezuelae Schultz, 1949 | WCA | Cartagena, Colombia |
|
2‡ |
A. urotrygoni*† | NR | NR | Hypanus guttatus | WSA, WCA | Lake Maracaibo, Venezuela |
|
– |
A. urotrygoni*† | NR | NR | Hypanus guttatus | WSA, WCA | Isla Margarita, Venezuela |
|
– |
A. vargasi Marques, Brooks & Monks, 1995 | MNHG 20011 |
MNHG 20012–20013; |
Hypanus longus | ECP | Punta Morales, Puntarenas Province, Costa Rica |
|
2‡ |
A. vidali Zaragoza-Tapia, Pulido-Flores, Violante-Gonzalez & Monks, 2019 | CNHE 11134 |
CNHE 11135; |
Narcine entemedor | ECP | Bahía de Acapulco, Playa Las Hamacas, Guerrero, Mexico |
|
6¶ |
A. walkeri Campbell & Beveridge, 2002 |
|
|
Pastinachus sephen** | NIO | Nickol Bay, Western Australia |
|
2§ |
A. waltairensis Uma Maheswari, Sanaka, Vijaya Lakshmi & Hanumantha Rao, 1987 | NR | NR | Himantura uarnak | WIO, NIO, EIO, WCP | Waltair coast, India |
|
3§ |
A. wedli Robinson, 1959 | NR | DMNZ 194b,c,d, 195–197 (syntype) | Zearaja nasuta (Müller & Henle, 1841) | WSP | Petone Beach, New Zealand |
|
4§ |
A. wedli† | NR | NR | Zearaja nasuta | WSP | PortobeIlo, Otago Harbour, New Zealand |
|
– |
A. wedli† | NR | NR | Zearaja nasuta | WSP | South Island, off Lyttelton, New Zealand |
|
– |
A. westi Vardo-Zalik & Campbell, 2011 |
|
|
Raja texana | WCA | Gulf of Mexico |
|
1¶ |
A. woodsholei Baer, 1948 | NR | MNHG 40028 (syntype) | Bathytoshia centroura | WSA, WCA, WNA | Woods Hole, Massachusetts, USA |
|
2(7)‡ |
A. woodsholei† | NR | NR | Bathytoshia centroura | WSA, WCA, WNA | Western North Atlantic |
|
– |
A. xiamenensis Yang & Lin, 1994 | NR | NR | Rhynchobatus djiddensis** | WIO, NIO | Xiamen, South Fujian, China |
|
5§ |
A. zainali Fyler & Caira, 2006 | MZUM (P) 151 |
|
Urogymnus polylepis | NIO, WCP | Off Kampung Abai, Kinabatangan River, Sabah, Malaysia |
|
1§ |
A. zapterycum Ostrowski de Nuñez, 1971 |
|
NR | Zapteryx brevirostris (Müller & Henle, 1841) | WSA | Mar del Plata, Buenos Aires, Argentina |
|
2‡ |
A. zapterycum† | NR |
|
Zapteryx brevirostris | WSA | Coastal waters off Villa Gessel, Argentina |
|
– |
A. zapterycum† | NR | NR | Zapteryx brevirostris | WSA | La Lucila del Mar, Argentina |
|
– |
A. zapterycum† | NR | NR | Zapteryx brevirostris | WSA | Puerto Quequén, Argentina |
|
– |
A. zapterycum† | NR | NR | Zapteryx brevirostris | WSA | Puerto Pirámides, Argentina |
|
– |
A. zimmeri Fyler, Caira & Jensen, 2009 |
|
|
Urogymnus acanthobothrium | WSP, WCP | Arafura Sea, east of Wessel Islands, Northern Territory, Australia. |
|
1¶ |
A. zschokkei Baer, 1948 | MHNG 88/39 | NR | Torpille (common name)** | ? | Naples, Italy |
|
6§ |
A. zschokkei*† | NR | NR | Torpedo marmorata | ENA, MED, ECA, ESA | Adriatic Sea, Mediterranean Sea |
|
– |
A. zschokkei*† | NR | NR | Torpedo torpedo | ENA, MED, ECA | Sète, France |
|
– |
A. zschokkei*† | NR | NR | Torpedo torpedo | ENA, MED, ECA | Adriatic Sea, Mediterranean Sea |
|
– |
The type localities where species of Acanthobothrium have been reported is shown in Figure
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
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.
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.
The information in the Figures
In Table
According to
The categorical method developed by
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 (
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.