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Introduction
The Central Mexican Pacific (CMP) region comprises the states of Nayarit, Jalisco, Colima and Michoacán as well as their coastal (Isabel and Marietas) and oceanic islands (Marías and Revillagigedo). The region is characterized by numerous geographic features such as bays, archipelagos and islands, as well as a heterogenic bottom that combines hard and soft components. The CMP is located in the Tropical Eastern Pacific Province nearby to the Warm Temperate Northeast Pacific Province (Spalding et al., 2007) and for this reason, it shelters northern and southern fauna of the Mexican Pacific (Arriaga-Cabrera et al., 1998). Due to its relevant flora and fauna, most of the islands of the CMP are catalogued as Natural Protected Areas (CONANP, 2005; CONANP, 2007a; CONANP, 2007b; CONANP, 2019).
The study regarding the Echinodermata in the CMP began in 1840 when Gray (1840) reported the asteroids Astropecten regalis and Luidia columbia in Nayarit. Until 1891, efforts in the collection of echinoderms mostly from deep-waters were performed by foreign expeditions such as Albatross and Challenger. As a result, many new records and new species of Crinoidea (Hartlaub, 1895), Asteroidea (Sladen, 1889), Ophiuroidea (Lütken & Mortensen, 1899), Echinoidea (Agassiz, 1872; Agassiz,1898), and Holothuroidea (Ludwig, 1894) were described.
The 20th century was very productive in terms of the number of published works (75). During this period, numerous foreign expeditions (Albatross, Velero, and Zaca) collected samples from shallow and deep-waters along the region resulting in new records, new species and extensive taxonomic monographs (e.g. Ludwig, 1905; Fisher, 1928a; Fisher, 1928b; Deichmann, 1937; Ziesenhenne, 1937; Ziesenhenne, 1940; Mortensen, 1948a; Mortensen, 1948b). The first national work regarding echinoderms from the CMP was the master´s thesis of Caso (1943) who worked with the Asteroidea from México. Caso published dissertations and numerous manuscripts regarding taxonomy (Caso, 1948a; Caso, 1951; Caso, 1961; Caso, 1962a; Caso, 1965; Caso, 1966; Caso, 1974a; Caso, 1974b; Caso, 1978b; Caso, 1980; Caso,1983b), including descriptions of new species (Caso, 1954; Caso, 1983a; Caso, 1986a), and general accounts on the biodiversity of the CMP (e.g. Caso, 1948b; Caso, 1962b). Their findings helped to better understand the echinoderm fauna from shallow-waters of the region and inspired other Mexican researchers to work with echinoderms from Revillagigedo Islands (Villalobos, 1992; Chacón-Torres et al., 1993; Bautista-Romero, Reyes-Bonilla, Lluch-Cota, & Lluch-Cota, 1994), Marietas Islands (Nepote-González, 1998), Colima (Espino-Barr, Cibrián-Rodríguez, & García-Boa, 1996) and even, with species collected in several areas from the CMP that resulted as new taxa (Solís-Marín & Laguarda-Figueras, 1999). At the end of the 20th century, the first ecological studies in the CMP were published (Holguin-Quiñones, 1994; Reyes-Bonilla, 1995; Espino-Barr et al., 1996).
The first two decades of the 21st century have produced a vast number of works (55) regarding topics such as new distribution records (Frontana-Uribe, Hernández-Alcántara, & Solís-Weiss, 2000; Sotelo-Casas, Cupul-Magaña, Solís-Marín, & Rodríguez-Troncoso, 2015; Valdés de Anda, Hendrickx, & Solís-Marín, 2018), description of new species (Honey-Escandón, Solís-Marín, & Laguarda-Figueras, 2011; Solís-Marín, Arriaga-Ochoa, Galván-Villa, & Laguarda-Figueras, 2018), ecology (Holguin-Quiñones & Michel-Morfin, 2006; López-Uriarte et al., 2009; Hermosillo-Nuñez et al., 2015; Herrero-Pérezrul, Ramírez-Ortiz, Rosales-Estrada, & Reyes-Bonilla, 2015; Hermosillo-Nuñez, Rodríguez-Zaragoza, Ortiz, Calderon-Aguilera, & Cupul-Magaña, 2016; Sotelo-Casas, Cupul-Magaña, Rodríguez-Zaragoza, Solís-Marín, & Rodríguez-Troncoso, 2018), and reproduction (Sotelo-Casas, Cupul-Magaña, Solís-Marín, & Rodríguez-Troncoso, 2016). It is important to emphasize that most of these studies are focused in the compilation of species of echinoderms inhabiting in different areas of the CMP (Chávez-Dagostino, Nepote-González, Solís-Marín, & Medina-Rosas, 2000; Fuentes-Farías, Villarroel-Melo, & Solís-Marín, 2005; Honey-Escandón, Solís-Marín, & Laguarda-Figueras, 2008; Ríos-Jara, Galván-Villa, & Solís-Marín, 2008a; Ríos-Jara et al., 2008b; Santos-Beltrán & Salazar-Silva, 2011; Ríos-Jara et al., 2013; Rodríguez-Troncoso et al., 2013; Solís-Marín et al., 2013a; Granja-Fernández, Herrero-Pérezrul, López-Pérez, Hernández-Morales, & Rangel-Solís, 2015a; Solís-Marín, Laguarda-Figueras, & Durán-González, 2016b; Granja-Fernández et al., 2017; Padilla-Pérez, Rodríguez-Troncoso, Sotelo-Casas, & Cupul-Magaña, 2017; Ríos-Jara, Juárez-Carrillo, & Galván-Villa, 2017; Nava-Bravo et al., 2019).
Despite the above, the records have shown many inconsistencies such as: 1) the names of some species have been mentioned (written or registered) incorrectly across time (e.g. misspelling), 2) many of the recorded species for the CMP are invalid, and 3) the generated checklists are not complete because they are based only in museum records or material collected mostly in shallow-waters. Thus, the aim of the present manuscript is to provide an accurate and updated checklist of the valid names and synonyms of the Echinodermata from the CMP according to literature, including new distribution records of Ophiuroidea. Moreover, we provide a historical review of the study of each class of Echinodermata, as well as an updated number of species for each state and island in the region.
Materials and methods
An exhaustive search of published literature for the CMP regarding the classes Crinoidea, Asteroidea, Ophiuroidea, Echinoidea, and Holothuroidea was performed (records dating from 1840 to 2019). Moreover, we visited reference collections in order to taxonomically identify specimens of Ophiuroidea: ICML-UNAM: Colección de Equinodermos “Dra. Ma. Elena Caso”, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, Ciudad de México, México; LACM: Natural History Museum, Los Angeles County Museum, Los Angeles, United States of America (USA); and, MCZ: Museum of Comparative Zoology, Harvard University, Cambridge, USA.
When available, the geographical coordinates of the provided localities in the literature were validated using ArcMap 10.4.1 software, to corroborate distribution within the CMP maritime zone. Records were assigned to any of the following areas: coastlines of the states of Colima (COL), Jalisco (JAL), Michoacán (MICH), and Nayarit (NAY), and the islands Isabel (ISA), Marías (MAR), Marietas (MAT), and Revillagigedo (REV).
With this information, a checklist of the valid names and synonyms of Echinodermata from the CMP was constructed. The list includes those synonyms used exclusively in this area, yet some species may have larger synonym lists for other geographic areas. Systematics arrangements and valid names agree with Kroh and Mooi (2020), Mah (2020), Messing (2020), Stöhr, O´Hara, and Thuy (2020), and WoRMS (2020).
Results and discussion
Historical review
Crinoidea: Six references documented the Crinoidea from the CMP; one during the 19th, one in the 20th, and four in the 21st century. Among the Echinodermata, Crinoidea represents the less studied class of the CMP. The study of the Crinoidea from the CMP began in 1895 when Hartlaub (1895) recorded the species Florometra tanneri (as Antedon rhomboidea) for MAR at 1 236 m depth. Several years later, using a submersible, Roux (2004) collected one specimen of Hyocrinus foelli in COL at 3 030 m depth. Finally, during deep-water dredging by the TALUD cruises aboard the R/V “El Puma” of the Universidad Nacional Autónoma de México, Florometra serratissima was collected in two stations located in COL between 1 040 and 1 106 m depth (Valdés de Anda et al., 2018).
Asteroidea: 57 references addressed the Asteroidea distributed along the CMP. During the 19th century only two works were published, meanwhile the 20th and 21st centuries presented similar numbers of works, 28 and 27, respectively. The 19th century represents the era with the lowest addition of Asteroidea. The first reported Asteroidea for the CMP were Luidia columbia and Astropecten regalis, both collected in San Blas, NAY (Gray, 1840).
The 20th century was the period with the highest addition of Asteroidea to the CMP (25 species). Ludwig (1905) described Ampheraster marianus, Henricia gracilis, and Sarkaster validus (= Lophaster furcilliger) for MAR and other areas of the Eastern Pacific. According to samples from the Albatross, Fisher (1911) recorded Hippasteria lepidonotus and Henricia clarki for deep-waters from MAR and REV, respectively. Clark (1916) provided the original descriptions of Saraster insignis (= Nearchaster (Nearchaster) aciculosus) and Cyllaster seminuda (= Henricia seminudus) from Clarion Island, REV. In 1936, the Templeton Crocker Expedition collected echinoderms during the expedition of the Zaca along the north of México and as a result, Ziesenhenne (1937) mentioned Astropecten armatus, Acanthaster ellisii (= Acanthaster planci), Linckia columbiae, Pauliella aenigma (= Paulia horrida), and Sclerasterias heteropaes for shallow-waters (up to 100 m depth) from Clarion Island. During 1937-1938, the Zaca continued performing collections of shallow-water echinoderms, and Clark (1940) reported Luidia bellonae (Manzanillo, COL and Chamela Bay, JAL), Luidia foliolata, Pharia pyramidata, and Phataria unifascialis (Tenacatita Bay, JAL). Later, Caso (1943) reported the species Heliaster microbrachius and Luidia superba for Manzanillo, COL, as well as Nidorellia armata for the same locality and MAR. In later years, the addition of species for the CMP was sporadic, with mentions of just one species per work: Patiria miniata (Socorro Island, REV; Adem et al., 1960), Astropecten californicus (Manzanillo, COL; Caso, 1961), Mithrodia bradleyi (REV; Caso, 1962b), Pentaceraster cumingi (Socorro and Clarion Islands, REV; Hertlein, 1963), and Asteropsis carinifera (REV; Maluf, 1991).
During the 21st century, only eight new records for the CMP have been documented. Fuentes-Farías et al. (2005) provided a list of echinoderms for MICH including the new records of Luidia tessellata and Heliaster helianthus. Two years later, the Mexican government decreed the MAR as a Biosphere Reserve, publishing a management and conservation program (CONANP, 2007a), which provided lists of the marine and terrestrial flora and fauna, and included the record of Echinaster (Othilia) tenuispina for the area. Honey-Escandón et al. (2008) and Solís-Marín et al. (2013b) provided the most-complete checklist of echinoderms for the Mexican Pacific and the Eastern Pacific, respectively, including the new records of Heliaster polybrachius (JAL), Mediaster transfuga and Meridiastra modesta (REV), and Narcissia gracilis (NAY). Recently, Nava-Bravo et al. (2019) included the record of Astropecten verrilli for MICH.
Ophiuroidea: 59 references have been published regarding this class. The 19th century was the least productive since only one work was published. During the 20th and 21st centuries, similar numbers of manuscripts were published, 30 and 28, respectively. In 1891, the Albatross conducted an expedition along the Eastern Pacific, and as a result, Lütken and Mortensen (1899) described the first species and subspecies of ophiuroids from deep-waters of the CMP (MAR): Amphiura serpentina, Astrodia excavata, Dougaloplus notacanthus, Ophiacantha costata, Ophiacantha hirta, Ophiacantha moniliformis, Ophiernus adspersus annectens, Ophiomitra granifera, Ophiomitra partita, Ophiosphalma variabile, and Ophiura (Ophiura) scutellata.
During the 20th century a total of 30 species were added to the list of ophiuroids of the CMP, representing the highest number, in comparison to the other two centuries accounted for in this review. Döderlein (1911) conducted a revision of the world’s Euryalida, reporting Asteroschema sublaeve for MAR. With specimens collected in shallow-waters of Clarion Island, Ziesenhenne (1937) described Ophiacantha pyriformis, and reported for the first time in the CMP, the presence of Ophiactis savignyi, Ophiocoma aethiops, Ophioderma variegatum, Ophionereis annulata, and Ophiothrix galapagensis. Later, Clark (1940) found the conspicuous Diopederma daniana, Ophiolepis variegata, and Ophiothrix (Ophiothrix) spiculata in JAL and COL. In the same year, Ziesenhenne (1940) published the results obtained by the cruise Velero III performed in 1938, and described Amphichondrius laevis, Amphiodia sculptilis, Ophiolepis plateia, and Ophiophragmus lonchophorus, all of them collected in Tenacatita Bay, JAL. Later, the additions of new records of ophiuroids per work were reduced: Ophiocomella alexandri (REV; Steinbeck & Ricketts, 1941), Ophioderma teres and Ophiolepis pacifica (Puerto Vallarta, JAL; Caso, 1951), Ophioderma panamense (REV; Ziesenhenne, 1955), Ophiomyxa panamensis (REV; Hertlein, 1963), Ophiostigma tenue (ISA; Clark, 1970), Ophiomusa lymani, Ophiothela mirabilis, and Ophiuroglypha irrorata irrorata (JAL and REV; Luke, 1982), Amphichondrius unamexici (= Amphichondrius granulatus; NAY; Caso, 1983a), Amphiura seminuda and Ophiophragmus marginatus (JAL and NAY; Caso, 1986b), Amphiura (Amphiura) assimilis and Gymnophiura mollis (MAR; Maluf, 1988), Ophiactis simplex (REV; Maluf, 1991), and Ophiuroconis bispinosa (REV; Hendler, 1996).
The addition of new records of ophiuroids (22) during the 21st century has been important. Frontana-Uribe et al. (2000) collected specimens of Ophiocomella schmitti associated with coral rubble in localities from Socorro Island (REV). Fuentes-Farías et al. (2005) included the record of Ophiothrix (Ophiothrix) rudis for MICH; in the same year, Maluf and Brusca (2005) published a checklist of the echinoderms from the Gulf of California, including the general distribution of the species, and the new records of Amphipholis elevata, Astrocaneum spinosum, Microphiopholis puntarenae, Ophiocomella sexradia, Ophiocnida hispida, Ophiolepis crassa, Ophiopholis bakeri, and Ophiophthalmus diplasia (MAR and REV). The genus Ophiophthalmus is an invalid junior synonym and a new assignment is needed (Hendler, 1996), but in the present work we kept this temporary genus in order to avoid systematic confusions. Later, Honey-Escandón et al. (2008) reported 33 species of ophiuroids for the CMP, of which, the amphiurids Amphiodia occidentalis, Amphiodia platyspina, Amphiodia violacea, Amphiodia (Amphispina) digitata, Amphiodia (Amphispina) urtica, Amphipholis pugetana, and Microphiopholis platydisca, represented new records in several states and islands from the CMP. In 2011, Astrodictyum panamense (Ayala-Bocos, Reyes-Bonilla, Herrero-Pérezrul, Walther-Mendoza, & Castañeda-Fernández de Lara, 2011) and Hemipholis gracilis (Hendler, 2011) were reported in REV and JAL, respectively. García-Hernández et al. (2014) compared the ecological diversity and species composition of benthic macroinvertebrates in rocky reefs of JAL and MAT, finding new records for Amphipholis squamata and Ophionereis perplexa. Finally, Granja-Fernández et al. (2017) provided a checklist of ophiuroids from the CMP, adding the new record of Ophiophragmus papillatus.
Echinoidea: 73 references documented the presence of the Echinoidea from the CMP, representing the major number of works among all the classes. The 19th century is represented only by two publications, the 20th century for 39, and the 21st century for 32 works. In the 19th century all the work regarding Echinoidea was published by Alexander Agassiz whom documented for the first time in the CMP the species Eucidaris thouarsii and Echinometra vanbrunti, both collected in Manzanillo, COL (Agassiz, 1872), and who described Brisaster townsendi and Plexechinus cinctus for stations located in deep-waters of MAR (Agassiz, 1898).
The 20th century was by far the most productive in terms of new addition of echinoids (27) in the CMP. Clark described Hesperocidaris perplexa and Echinometra insularis (Clark, 1907; Clark, 1912), and reported Echinometra oblonga and Tripneustes gratilla from REV (Clark, 1912). With specimens collected by the Zaca, Ziesenhenne (1937) reported Clypeaster europacificus, Diadema mexicanum, and Tripneustes depressus in shallow-waters of Socorro and Clarion, REV. In 1938, was published an important work regarding fossil and living Echinoidea from the west American Cenozoic, in which were mentioned the new records of Astropyga pulvinata (JAL) and Toxopneustes roseus (ISA) (Grant & Hertlein, 1938). Posteriorly, Clark (1940) added Clypeaster ochrus and Clypeaster speciosus from ISA and NAY, respectively, and Caso (1948a) included Arbacia stellata to NAY. Clark (1948) reported the highest number of new records of echinoids for the CMP (Brissopsis pacifica, Clypeaster rotundus, Encope micropora, Encope micropora insularis, Lovenia cordiformis, Meoma ventricosa grandis, and Rhyncholampas pacificus), based on the collections of the Velero III along the Eastern Pacific during the years 1931-1941; E. micropora insularis was described in this work and its holotype corresponds to Socorro Island, REV. The records in posterior works were sporadic: Hesperocidaris asteriscus (JAL and REV; Caso, 1961), Heterocentrotus mamillatus (REV; Chan, 1974), Dendraster excentricus and Lanthonia longifissa (ISA and COL, respectively; Caso, 1980), Agassizia scrobiculata (NAY; Caso, 1983b), and Mellita notabilis (MICH; Harold & Telford, 1990).
Despite the number of published works during the 21st century is similar to the previous century, only three new records for the area have been documented during the present era. These records correspond to species observed during field work and documented in the management and conservation programs of the National Park ISA (Centrostephanus coronatus; CONANP, 2005) and the Reserve of the Biosphere MAR (Brissus obesus; CONANP, 2007a). Finally, Ríos-Jara et al. (2008a) performed collections of echinoderms in ISA at different depths finding Brissus latecarinatus in shallow-waters (< 20 m depth).
Holothuroidea: 53 references mention holothuroids from the CMP. During the 19th century only one work was published, 19 during the 20th century, and 33 during the 21st century. In the 19th century only one work was published and therefore, the number of records was low. Despite, Laetmophasma fecundum (= Pannychia moseleyi), Psolus pauper, and Ypsilothuria bitentaculata were first mentioned in Ludwig (1893), it was until 1864 when Ludwig (1894) provided an extended description of the species, reporting them in deep waters from MAR; moreover, Molpadia musculus was recorded in this work too, for the same locality.
The 20th century was the most productive in terms of the addition of new records (28). In 1926, the Templeton Crocker expedition aboard the Zaca performed dredges in order to obtain echinoderms along the Eastern Pacific, and collected Holothuria (Cystipus) inhabilis and Holothuria (Platyperona) difficilis in REV (Deichmann, 1937). From 1932 to 1938, the Allan Hancock Expedition aboard the Velero III collected several specimens of holothuroids and Deichmann (1941) found that Thyone bidentata and Thyone parafusus, both discovered in Tenacatita, JAL corresponded to new species, and that Leptopentacta panamica, Neothyone gibber, and Pentamera chierchiae were new records of species for the CMP (JAL and REV). Caso (1954) collected for the first time Holothuria (Selenkothuria) lubrica in Puerto Vallarta, JAL and found a similar species which was designated as a new species: Holothuria (Selenkothuria) portovallartensis. The biggest contribution of new records of holothuroids for the CMP (Holothuria (Halodeima) kefersteinii, Holothuria (Lessonothuria) pardalis, Holothuria (Mertensiothuria) hilla, Holothuria (Mertensiothuria) leucospilota, Holothuria (Semperothuria) imitans, Holothuria (Semperothuria) languens, Holothuria (Theelothuria) paraprinceps, Holothuria (Thymiosycia) arenicola, Holothuria (Thymiosycia) impatiens, and Isostichopus fuscus) was made by Deichmann (1958) which correspond to a continuation and expansion of the material collected by the Velero III and IV in JAL, as well as in ISA, MAR and REV. The new records per work during the rest of this century were reduced: Holothuria (Halodeima) inornata (Caso, 1961), Leptosynapta albicans (Caso, 1962b), Abyssocucumis abyssorum (Luke, 1982), Holothuria (Stauropora) fuscocinerea (Hickman, 1998), all of them collected in REV and NAY. Nepote-González (1998) conducted a research about the diversity of holothuroids of MAT, finding the new records of Chiridota rigida, Holothuria (Cystipus) rigida, Labidodemas americanum, and Pseudocnus californicus. Later, Solís-Marín and Laguarda-Figueras (1999) performed collections in several localities of the CMP (COL, NAY, MAT) and described a new species, Cucumaria flamma.
The addition of new records of holothuroids to the CMP has been important during the 21st century. As part of the first program of management and conservation of REV National Park, CONANP (2004) documented Euapta godeffroyi. Fuentes-Farías et al. (2005) and Honey-Escandón et al. (2008) provided one of the most complete checklists of echinoderms in México and added the records of Neothyone gibbosa and Labidodemas maccullochi for MICH, and Euthyonidiella zacae, Holothuria (Halodeima) atra, Holothuria (Mertensiothuria) viridiaurantia (= Holothuria (Mertensiothuria) hilla), Lissothuria ornata, Neocucumis veleronis, and Trachythyone peruana for several areas of the Mexican Pacific. Holothuria (Mertensiothuria) viridiaurantia is a recently described species which was previously assigned to H. (Mertensiothuria) hilla (Borrero-Pérez & Vanegas-González, 2019). According to a preliminary revision of the material (unpublished data) and to Borrero-Pérez and Vanegas-González (2019), in the CMP, H. (Mertensiothuria) viridiaurantia inhabits in NAY, JAL, MICH and MAT. On the other hand, based on a taxonomical review of the material, we confirm that H. (Mertensiothuria) hilla inhabits in REV, but a taxonomic confirmation of the material from ISA and MAR is required (F.A. Solís-Marín, personal communication, 01 April 2020). Honey-Escandón et al. (2011) described Holothuria (Selenkothuria) carere, collected in shallow-waters of MICH and other states of the Mexican Pacific. Santos-Beltrán and Salazar-Silva (2011) provided images and an identification key of the holothuroids from Bahía de Banderas, NAY adding the record of Afrocucumis ovulum. Ríos-Jara et al. (2013) described the taxonomic composition of the echinoderms from shallow-waters in several localities of Bahía Chamela, JAL, adding Neopentamera anexigua, Pachythyone lugubris, and Pachythyone pseudolugubris to the CMP, and Solís-Marín, Arriaga-Ochoa, Galván-Villa, and Laguarda-Figueras (2018) described Lissothuria imbricata after the collection of material from the same area. In recent years, Nava-Bravo et al. (2019) elaborated the most complete checklist of echinoderms from MICH adding the records of Chiridota aponocrita, Epitomapta tabogae, and Thyonella mexicana.
Checklist: The present updated checklist includes 187 species of Echinodermata (three Crinoidea, 35 Asteroidea, 67 Ophiuroidea, 32 Echinoidea, and 50 Holothuroidea; Table 1) resulted from the revision of 137 literature references (3 445 records) and the taxonomic revision of Ophiuroidea deposited at museum collections. Additionally, we detected 57 records in the literature (two Crinoidea, 21 Asteroidea, 14 Ophiuroidea, 15 Echinoidea, and five Holothuroidea) considered as invalid and which were removed from the updated checklist.
Table 1 Checklist of valid names (in bold) and synonyms of the Echinodermata from the Central Mexican Pacific based on literature review and museum records. All taxa are listed exactly as originally registered in the literature. Numbers refers to literature where taxa were listed, see References section. * = New distribution records
| Phylum Echinodermata Bruguière, 1791 |
|---|
| Class Crinoidea Miller, 1821 |
| Order Comatulida A.H. Clark, 1908 |
| Family Antedonidae Norman, 1865 |
| Genus Florometra A.H. Clark, 1913 |
| Florometra serratissima (A.H. Clark, 1907) COL 132 |
| Florometra tanneri (Hartlaub, 1895) MAR 90, 92 |
| Antedon rhomboideaHartlaub, 189568 |
| Orden Hyocrinida Rasmussen, 1978 |
| Familia Hyocrinidae Carpenter, 1884 |
| Genus Hyocrinus Thomson, 1876 |
| Hyocrinus foelliRoux & Pawson, 1999 COL 81, 114 |
| Class Asteroidea de Blainville, 1830 |
| Order Paxillosida Perrier, 1884 |
| Family LuidiidaeSladen, 1889 |
| Genus Luidia Forbes, 1839 |
| Luidia bellonae Lütken, 1864 JAL, COL, MICH, REV 13, 39, 104 |
| Luidia bellona Lütken, 1864 7, 92 |
| Luidia (Alternaster) bellona Lütken, 1864 91 |
| Luidia columbia (Gray, 1840) NAY, COL, MICH, REV 13, 15, 92, 104, 126, 135 |
| Luidia brevispina Lütken, 1871 8, 13 |
| Luidia columbiae (Gray, 1840) 7, 117 |
| Luidia (Petalaster) columbia (Gray, 1840) 58, 81, 91 |
| Petalaster ColumbiaGray, 184066 |
| Luidia foliolata Grube, 1866 JAL 39 |
| Luidia superba A.H. Clark, 1917 COL, MICH 8, 10, 13, 30, 124 |
| Luidia (Alternaster) superba A.H. Clark, 1917 58, 81, 91 |
| Luidia tessellata Lütken, 1859 NAY, MICH, ISA 104 |
| Luidia (Petalaster) tessellata Lütken, 1859 58, 81, 109, 110 |
| Family AstropectinidaeGray, 1840 |
| Genus Astropecten Gray, 1840 |
| Astropecten armatusGray, 1840 NAY, COL, MICH, REV 7, 8, 13, 15, 28, 30, 39 58, 81, 92, 104, 124, 135 |
| Astropecten regalisGray, 1840 NAY, COL, MICH 8, 13, 58, 66, 81, 104, 117, 124 |
| Astropecten californicus Fisher, 1906 NAY, COL 13, 81, 124 |
| Astropecten verrilli de Loriol, 1899 MICH 104 |
| Genus Psilaster Sladen, 1885 |
| Psilaster armatusLudwig, 1905 MAR 86, 90, 92 |
| Order Valvatida Perrier, 1884 |
| Family AcanthasteridaeSladen, 1889 |
| Genus Acanthaster Gervais, 1841 |
| Acanthaster planci (Linnaeus, 1758) NAY, COL, MAR, ISA, MAT, REV 7, 32, 33, 41, 44, 45, 81, 107, 108, 120, 124 |
| Acantaster planci (Linnaeus, 1758) 126 |
| Acantaster plansii (Linnaeus, 1758) 31 |
| Acanthaster elissi (Gray, 1840) 134 |
| Acanthaster elisii pseudoplanciCaso, 196279 |
| Acanthaster ellisi (Gray, 1840) 43, 45 |
| Acanthaster ellisii (Gray, 1840) 14, 15, 19, 41, 42, 72, 81, 92, 109, 110, 124, 135 |
| Acanthaster ellisii pseudoplanciCaso, 196214, 15, 41, 69 |
| Acanthaster pseudoplanciCaso, 196219 |
| Family AsterinidaeGray, 1840 |
| Genus Meridiastra O’Loughlin, 2002 |
| Meridiastra modesta (Verrill, 1867) REV 81, 124 |
| Asterina agustincasoi Caso, 1977 45 |
| Genus Patiria Gray, 1840 |
| Patiria miniata (Brandt, 1835) REV 120 |
| Asterina miniata (Brandt, 1835) 1, 7, 15, 41, 45, 81 |
| Patiria minata (Brandt, 1835) 126 |
| Family AsterodiscididaeRowe, 1977 |
| Genus Paulia Gray, 1840 |
| Paulia horridaGray, 1840 REV 41, 45, 91, 92, 126 |
| Pauliella aenigmaLudwig, 19057, 13, 15, 77, 135 |
| Family Asteropseidae Hotchkiss & A.M. Clark, 1976 |
| Genus Asteropsis Müller & Troschel, 1840 |
| Asteropsis carinifera (Lamarck, 1816) JAL, ISA, REV 41, 45, 91, 92, 126 |
| Asteropsis spinosa (Gray, 1840) 81 |
| Gymnasteria spinosaGray, 1840109, 110 |
| Family Goniasteridae Forbes, 1841 |
| Genus Hippasteria Gray, 1840 |
| Hippasteria lepidonotus (Fisher, 1905) MAR |
| Cryptopeltaster lepidonotus Fisher, 1905 13, 54, 90, 92 |
| Hippasteria pacificaLudwig, 190554 |
| Genus Mediaster Stimpson, 1857 |
| Mediaster transfugaLudwig, 1905 REV 45, 120 |
| Family Ophidiasteridae Verrill, 1870 |
| Genus Linckia Nardo, 1834 |
| Linckia columbiaeGray, 1840 NAY, JAL, MAR, REV 1, 7, 8, 13, 15, 41, 43, 45, 81, 91, 92, 126, 135 |
| Genus Narcissia Gray, 1840 |
| Narcissia gracilis A.H. Clark, 1916 NAY 81 |
| Genus Pharia Gray, 1840 |
| Pharia pyramidata (Gray, 1840) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 8, 13, 15, 39, 44, 73, 91, 92, 96, 104, 107, 113, 120, 129 |
| Pharia piramidata (Gray, 1840) 42 |
| Pharia pyramidatus (Gray, 1840) 33, 43, 45, 58, 72, 109, 110, 111, 112, 121, 124, 126 |
| Pharia pyramidatus pyramidatus (Gray, 1840) 81 |
| Pharia pyrimidata (Gray, 1840) 87 |
| Genus Phataria Gray, 1840 |
| Phataria unifascialis (Gray, 1840) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 15, 22, 29, 33, 39, 42, 43, 44, 45, 58, 72, 73, 81, 84, 87, 91, 92, 96, 104, 107, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129 |
| Phataria unifacialis (Gray, 1840) 8, 13 |
| Family Mithrodiidae Viguier, 1878 |
| Genus Mithrodia Gray, 1840 |
| Mithrodia bradleyi Verrill, 1867 NAY, COL, MICH, MAR, ISA, MAT, REV 7, 15, 33, 41, 43, 44, 45, 58, 79, 81, 87, 91, 92, 104, 107, 108, 109, 110, 120, 124, 126, 129, 134 |
| Family Oreasteridae Fisher, 1908 |
| Genus Nidorellia Gray, 1840 |
| Nidorellia armata (Gray, 1840) JAL, COL, MICH, MAR, ISA, MAT, REV 58, 81, 104, 107, 108, 109, 110, 129 |
| Nidoriella armata (Gray, 1840) 7, 8, 13 |
| Genus Pentaceraster Döderlein, 1916 |
| Pentaceraster cumingi (Gray, 1840) NAY, JAL, COL, MICH, ISA, MAT, REV 58, 81, 91, 92, 104, 107, 109, 110, 111, 113, 124, 126 |
| Oreaster occidentalis Verrill, 1867 7, 77 |
| Family Solasteridae Viguier, 1878 |
| Genus Lophaster Verrill, 1878 |
| Lophaster furcilliger Fisher, 1905 MAR |
| Lophaster validus (Ludwig, 1905) 90, 91 |
| Sarkaster validusLudwig, 190586 |
| Order Forcipulatida Perrier, 1884 |
| Family AsteriidaeGray, 1840 |
| Genus Sclerasterias Perrier, 1891 |
| Sclerasterias heteropaes Fisher, 1924 REV 7, 13, 15, 90, 92, 126, 135 |
| Family Heliasteridae Viguier, 1879 |
| Genus Heliaster Gray, 1840 |
| Heliaster helianthus (Lamarck, 1816) NAY, MICH 58, 81, 104 |
| Heliaster microbrachius Xantus, 1860 JAL, COL 8, 13, 81 |
| Heliaster polybrachius H.L. Clark, 1907 JAL 81 |
| Family Pedicellasteridae Perrier, 1884 |
| Genus Ampheraster Fisher, 1923 |
| Ampheraster marianus (Ludwig, 1905) MAR 13, 55, 56, 90 |
| Sporasterias marianaLudwig, 190586 |
| Order Spinulosida Perrier, 1884 |
| Family Echinasteridae Verrill, 1867 |
| Genus Echinaster Müller & Troschel, 1840 |
| Subgenus Othilia Gray, 1840 |
| Echinaster (Othilia) tenuispina Verrill, 1871 MAR |
| Echinaster tenuispina Verrill, 1871 43 |
| Echinaster (Othilia) teniuspinus Verrill, 1871 81 |
| Genus Henricia Gray, 1840 |
| Henricia clarki Fisher, 1910 REV 7, 13, 15, 45, 54, 56, 74, 81, 90, 92, 120, 124 |
| Henricia gracilis (Ludwig, 1905) MAR 90, 91, 92 |
| Cribrella gracilisLudwig, 190586 |
| Henricia seminudus (A.H. Clark, 1916) REV 45, 81, 120, 124 |
| Cyllaster seminuda A.H. Clark, 191613, 15, 34 |
| Henricia seminuda (A.H. Clark, 1916) 90 |
| Order Notomyotida Ludwig, 1910 |
| Family Benthopectinidae Verrill, 1899 |
| Genus Nearchaster Fisher, 1911 |
| Subgenus Nearchaster Fisher, 1911 |
| Nearchaster (Nearchaster) aciculosus (Fisher, 1910) REV |
| Nearchaster aciculosus (Fisher, 1910) 7, 90, 120 |
| Saraster insignis A.H. Clark, 191613, 15, 34 |
| Class OphiuroideaGray, 1840 |
| Order Euryalida Lamarck, 1816 |
| Family Asteronychidae Ljungman, 1867 |
| Genus Astrodia Verrill, 1899 |
| Astrodia excavata (Lütken & Mortensen, 1899) MAR 64, 92, 106 |
| Asteronyx excavataLütken & Mortensen, 189913, 38, 50, 52, 63, 81, 88 |
| Family EuryalidaeGray, 1840 |
| Genus Asteroschema Örsted & Lütken in Lütken, 1856 |
| Asteroschema sublaeveLütken & Mortensen, 1899 MAR 50, 63, 64 |
| Family Gorgonocephalidae Ljungman, 1867 |
| Genus Astrocaneum Döderlein, 1911 |
| Astrocaneum spinosum (Lyman, 1875) MAR 63, 64 |
| Asterocaneum spinosum (Lyman, 1875) 92 |
| Genus Astrodictyum Döderlein, 1927 |
| Astrodictyum panamense (Verrill, 1867) REV 6, 45, 63, 76 |
| Order Ophiurida Müller & Troschel, 1840, restricted sensu O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017 |
| Family OphiosphalmidaeO’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018 |
| Genus Ophiosphalma H.L. Clark, 1941 |
| Ophiosphalma variabile (Lütken & Mortensen, 1899) JAL, COL, MAR, REV |
| Ophiomusium variabileLütken & Mortensen, 189945, 63, 64, 81, 88, 92, 120, 123, 124 |
| Family OphiomusaidaeO’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018 |
| Genus Ophiomusa Hertz, 1927 |
| Ophiomusa lymani (Wyville-Thomson, 1873) MAR |
| Ophiomusium lymani Wyville-Thomson, 1873 63, 64, 87 |
| Family Ophiuridae Müller & Troschel, 1840 |
| Genus Ophiura Lamarck, 1801 |
| Subgenus Ophiura Lamarck, 1801 |
| Ophiura (Ophiura) scutellata (Lütken & Mortensen, 1899) MAR 63, 64 |
| Homalophiura scutellata (Lütken & Mortensen, 1899) 38, 52 |
| Ophioglypha scutellataLütken & Mortensen, 189988 |
| Ophiura scutellata (Lütken & Mortensen, 1899) 90, 92 |
| Genus Ophiuroglypha Hertz, 1927 |
| Ophiuroglypha irrorata irrorata (Lyman, 1878) MAR |
| Ophiura (Ophiuroglypha) irrorata irrorata (Lyman, 1878) 63, 64 |
| Ophiura irrorata (Lyman, 1878) 87 |
| Family Ophiopyrgidae Perrier, 1893 |
| Genus Gymnophiura Lütken & Mortensen, 1899 |
| Gymnophiura mollisLütken & Mortensen, 1899 MAR 63, 64, 90, 91, 92 |
| Order Ophioscolecida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017 |
| Family Ophioscolecidae Lütken, 1869 |
| Genus Ophiuroconis Matsumoto, 1915 |
| Ophiuroconis bispinosaZiesenhenne, 1937 JAL, COL, REV 45, 63, 64, 70, 81, 92, 120, 123, 124 |
| Order Ophiacanthida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017 |
| Family Ophiotomidae Paterson, 1985 |
| Genus Ophiomitra Lyman, 1869 |
| Ophiomitra graniferaLütken & Mortensen, 1899 NAY, MAR 63, 64, 87, 88 |
| Ophiacantha granifera (Lütken & Mortensen, 1899) 13 |
| Ophioplinthaca granifera (Lütken & Mortensen, 1899) 38, 52, 92 |
| Ophiomitra partitaLütken & Mortensen, 1899 MAR 63, 64, 88 |
| Ophiacantha partita (Lütken & Mortensen, 1899) 13 |
| Ophioplinthaca partita (Lütken & Mortensen, 1899) 38, 52, 92 |
| Family Ophiacanthidae Ljungman, 1867 |
| Genus Ophiacantha Müller & Troschel, 1842 |
| Ophiacantha costataLütken & Mortensen, 1899 MAR 13, 38, 52, 63, 64, 83, 88, 92 |
| Ophiacantha hirtaLütken & Mortensen, 1899 MAR 52, 63, 64, 81, 88, 92 |
| Ophiacantha moniliformisLütken & Mortensen, 1899 MAR, REV 38, 45, 52, 63, 64, 81 88, 92, 120, 123, 124 |
| Ophiacantha pyriformisZiesenhenne, 1937 REV 7, 13, 15, 45, 52, 63, 90, 135 |
| Genus Ophiophthalmus Matsumoto, 1917 (temporary genus) |
| Ophiophthalmus diplasia (H.L. Clark, 1911) REV |
| Ophiacantha diplasia H.L. Clark, 1911 45, 63, 92 |
| Family Ophiodermatidae Ljungman, 1867 |
| Genus Ophioderma Müller & Troschel, 1840 |
| Ophioderma panamense Lütken, 1859 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 41, 42, 77, 79, 81, 84, 91, 92, 104, 109, 110, 111, 123, 124, 137 |
| Ophiocryptus granulosus Nielsen, 1932 81 |
| Ophioderma panamensis Lütken, 1859 45, 62, 63, 64 |
| Ophioderma teres (Lyman, 1860) NAY, JAL, COL, MAR, REV* 11, 13, 62, 63, 64, 81, 92 |
| Ophioderma variegatum Lütken, 1856 NAY, JAL, COL, MAR, ISA*, REV 7, 11, 13, 15, 27, 29, 39, 41, 45, 77, 79, 81, 84, 91, 92, 111, 120, 124, 134, 135 |
| Ophioderma variegata Lütken, 1856 63, 64, 123, 137 |
| Genus Diopederma H.L. Clark, 1913 |
| Diopederma daniana (Verrill, 1867) JAL, MICH 63, 64, 104 |
| Diopederma danianum (Verrill, 1867) 11, 39, 58, 81 |
| Family Ophiomyxidae Ljungman, 1867 |
| Genus Ophiomyxa Müller & Troschel, 1842 |
| Ophiomyxa panamensisLütken & Mortensen, 1899 REV 7, 63, 77, 91, 92 |
| Family Ophiocomidae Ljungman, 1867 |
| Genus Ophiocoma L. Agassiz, 1836 |
| Ophiocoma aethiops Lütken, 1859 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 11, 13, 15, 29, 41, 42, 43, 44, 45, 58, 59, 62, 63, 64, 79, 81, 84, 87, 91, 92, 104, 109, 110, 111, 120, 121, 122, 123, 124, 130, 134, 135 |
| Genus Ophiocomella A.H. Clark, 1939 |
| Ophiocomella alexandri (Lyman, 1860) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV |
| Ophiocoma alexandri Lyman, 1860 7, 11, 13, 15, 29, 41, 42, 43, 44, 45, 49, 58, 59, 62, 63, 64, 79, 81, 84, 87, 91, 92, 104, 109, 110, 111, 120, 121, 123, 124, 130, 134 |
| Ophiocomella schmitti A.H. Clark, 1939 NAY, COL, MAR*, REV 45, 57, 63, 64, 81, 123, 124 |
| Ophiocomella sexradia (Duncan, 1887) MAR, REV 45, 63, 64, 92 |
| Order Ophioleucida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017 |
| Family OphiernidaeO’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018 |
| Genus Ophiernus Lyman, 1878 |
| Ophiernus adspersus annectensLütken & Mortensen, 1899 MAR 63, 64, 89 |
| Ophiernus adspersus adspersus Lyman, 1883 81 |
| Ophiernus polyporumLütken & Mortensen, 189938, 52 |
| Ophiernus polyporusLütken & Mortensen, 189988 |
| Order Amphilepidida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017 |
| Family Ophiolepididae Ljungman, 1867 |
| Genus Ophiolepis Müller & Troschel, 1840 |
| Ophiolepis crassa Nielsen, 1932 COL*, MAR, ISA*, MAT 63, 64, 92 |
| Ophiolepis pacifica Lütken, 1856 NAY, JAL, MICH*, MAR, MAT, REV* 62, 63, 64, 81 |
| Ophiozona pacifica (Lütken, 1856) 11, 13, 29, 92 |
| Ophiolepis plateiaZiesenhenne, 1940 JAL, ISA* 13, 52, 63, 64, 81, 115, 136 |
| Ophiolepis variegata Lütken, 1856 NAY, JAL, COL, MAR 11, 39, 63, 64, 81, 87, 92, 123, 124 |
| Family Ophionereididae Ljungman, 1867 |
| Genus Ophionereis Lütken, 1859 |
| Ophionereis annulata (Le Conte, 1851) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 13, 15, 45, 58, 59, 61, 62, 63, 64, 81, 91, 92, 104, 109, 110, 111, 120, 123, 124, 130, 135 |
| Ophiocoma annulata (Le Conte, 1851) 43 |
| Ophionereis dictiotaZiesenhenne, 194041 |
| Ophionereis dictyotaZiesenhenne, 194011, 45, 104, 123, 124, 134 |
| Ophionereis perplexaZiesenhenne, 1940 JAL, MAT 59 |
| Family Amphiuridae Ljungman, 1867 |
| Genus Amphiura Forbes, 1843 |
| Amphiura seminudaLütken & Mortensen, 1899 NAY, REV 28, 63, 92 |
| Amphiura serpentinaLütken & Mortensen, 1899 MAR 52, 63, 64, 92, 131 |
| Amphiura serpentina var. a Lütken & Mortensen, 189988 |
| Subgenus Amphiura (Amphiura) Forbes, 1843 |
| Amphiura (Amphiura) assimilis Lütken & Mortensen, 1899MAR |
| Amphiodia assimilis (Lütken & Mortensen, 1899) 63, 64, 90 |
| Amphiura assimilisLütken & Mortensen, 189992 |
| Genus Amphichondrius Nielsen, 1932 |
| Amphichondrius granulatus (Lütken & Mortensen, 1899) NAY |
| Amphichondrius unamexiciCaso, 198325, 28, 90, 92 |
| Amphichondrius laevisZiesenhenne, 1940 JAL 52, 63, 64, 81, 87, 115, 136 |
| Genus Amphiodia Verrill, 1899 |
| Amphiodia occidentalis (Lyman, 1860) NAY 63, 64, 81 |
| Amphiodia platyspina Nielsen, 1932 NAY 63, 64, 81 |
| Amphiodia psara H.L. Clark, 1935 NAY* |
| Amphiodia sculptilisZiesenhenne, 1940 JAL 52, 63, 64, 81, 136 |
| Amphiodia tabogae Nielsen, 1932 NAY* |
| Amphiodia violacea (Lütken, 1856) NAY 63, 64, 81 |
| Subgenus Amphispina Nielsen, 1932 |
| Amphiodia (Amphispina) digitata Nielsen, 1932 NAY 63, 64, 81 |
| Amphiodia (Amphispina) urtica (Lyman, 1860) NAY, MAR 63, 64, 81 |
| Genus Amphipholis Ljungman, 1866 |
| Amphipholis elevata Nielsen, 1932 MAR 63, 64, 92 |
| Amphipholis pugetana (Lyman, 1860) NAY, REV* 63, 64, 81 |
| Amphipholis squamata (Delle Chiaje, 1828) NAY, JAL, COL, MAT, ISA* 59, 63, 64 |
| Genus Dougaloplus A.M. Clark, 1970 |
| Dougaloplus notacanthus (Lütken & Mortensen, 1899) MAR 63, 64, 92 |
| Amphiura notacanthaLütken & Mortensen, 189952, 81, 88 |
| Genus Microphiopholis Turner, 1985 |
| Microphiopholis geminata (Le Conte, 1851) NAY* |
| Microphiopholis platydisca (Nielsen, 1932) NAY, COL 63, 64 |
| Amphipholis platydisca Nielsen, 1932 81 |
| Microphiopholis puntarenae (Lütken, 1856) MAR 63, 64 |
| Amphipholis puntarenae (Lütken, 1856) 92 |
| Genus Ophiocnida Lyman, 1865 |
| Ophiocnida hispida (Le Conte, 1851) NAY*, JAL, COL, MAR, MAT 59, 62, 63, 64, 92 |
| Genus Ophiophragmus Lyman, 1865 |
| Ophiophragmus lonchophorusZiesenhenne, 1940 NAY, JAL 13, 52, 63, 64, 90, 136 |
| Ophiophragmus marginatus (Lütken, 1856) NAY, JAL 28, 63, 64, 81, 111 |
| Ophiophragmus papillatusZiesenhenne, 1940 NAY, JAL, MICH, MAT, REV* 64, 104 |
| Ophiophragmus paucispinus Nielsen, 1932 MAR* |
| Genus Ophiostigma Lütken, 1856 |
| Ophiostigma tenue Lütken, 1856 JAL, MAR, ISA, MAT 35, 63, 64, 92 |
| Family Ophiactidae Matsumoto, 1915 |
| Genus Ophiactis Lütken, 1856 |
| Ophiactis savignyi (Müller & Troschel, 1842) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 11, 13, 23, 29, 45, 58, 59, 62, 63, 64, 81, 91, 104, 111, 120, 121, 122, 123, 124, 130, 135 |
| Ophiactis simplex (Le Conte, 1851) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 58, 59, 62, 63, 64, 81, 91, 92, 104, 111 |
| Genus Hemipholis Lyman, 1865 |
| Hemipholis gracilis Verrill, 1867 NAY, JAL 63, 64, 71 |
| Family OphiopholidaeO‘Hara, Stöhr, Hugall, Thuy, & Martynov, 2018 |
| Genus Ophiopholis Müller & Troschel, 1842 |
| Ophiopholis bakeri McClendon, 1909 REV 63, 92 |
| Family Ophiotrichidae Ljungman, 1867 |
| Genus Ophiothrix Müller & Troschel, 1840 |
| Ophiothrix galapagensisLütken & Mortensen, 1899 MAR, REV 7, 13, 15, 45, 63, 64, 91, 92, 135 |
| Subgenus Ophiothrix A.M. Clark, 1966 |
| Ophiothrix (Ophiothrix) rudis Lyman, 1874 NAY, JAL, MICH, MAR, MAT, REV 62, 63, 64 |
| Ophiothrix rudis Lyman, 1874 58, 81, 92, 104 |
| Ophiotrix rudis Lyman, 1874 44 |
| Ophiothrix (Ophiothrix) spiculata Le Conte, 1851 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 62, 63, 64, 104 |
| Ophiothrix spiculata Le Conte, 1851 11, 13, 29, 39, 58, 59, 81, 87, 91, 92, 111, 123, 124 |
| Genus Ophiothela Verrill, 1867 |
| Ophiothela mirabilis Verrill, 1867 NAY, JAL, COL, MICH, ISA, MAT 58, 59, 62, 63, 64, 81, 104, 109, 110, 111 |
| Ophiothrix mirabilis (Verrill, 1867) 87 |
| Class Echinoidea Leske, 1778 |
| Order Cidaroida Claus, 1880 |
| Family Cidaridae Gray, 1825 |
| Genus Eucidaris Pomel, 1883 |
| Eucidaris thouarsii (L. Agassiz & Desor, 1846) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 9, 13, 15, 21, 32, 33, 39, 41, 42, 43, 44, 45, 59, 65, 72, 73, 75, 77, 81, 84, 87, 92, 104, 107, 108, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129, 134, 135 |
| Cidaris Thouarsii L. Agassiz & Desor, 1846 2 |
| Eucidaris Thouarsii (L. Agassiz & Desor, 1846) 40 |
| Eucidaris thourarsii (L. Agassiz & Desor, 1846) 91 |
| Eucidaris thowarsii (L. Agassiz & Desor, 1846) 79 |
| Genus Hesperocidaris Mortensen, 1928 |
| Hesperocidaris asteriscus H.L. Clark, 1948 JAL, MICH, ISA, REV 7, 13, 15, 21, 45, 58, 92, 104, 111, 113, 126 |
| Hesperocidaris perplexa (H.L. Clark, 1907) REV 7, 13, 15, 21, 45, 65, 81, 92, 97, 120, 124, 135 |
| Tretocidaris perplexa H.L. Clark, 190736, 51 |
| Order Diadematoida Duncan, 1889 |
| Family Diadematidae Gray, 1855 |
| Genus Astropyga Gray, 1825 |
| Astropyga pulvinata (Lamarck, 1816) JAL, COL, MICH, MAR, MAT, REV 9, 21, 33, 41, 44, 45, 58, 65, 73, 81, 87, 91, 92, 104, 107, 113 |
| Astropygia pulvinata (Lamarck, 1816) 126 |
| Genus Centrostephanus Peters, 1855 |
| Centrostephanus coronatus (Verrill, 1867) JAL, COL, MICH, MAR, ISA, MAT 42, 43, 44, 72, 73, 81, 96, 107, 109, 110, 111, 112, 113, 124, 129 |
| Genus Diadema Gray, 1825 |
| Diadema mexicanum A. Agassiz, 1863 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 1, 5, 7, 13, 15, 21, 22, 29, 31, 33, 41, 42, 43, 44, 45, 53, 58, 72, 73, 74, 75, 79, 81, 84, 91, 92, 96, 98, 104, 107, 108, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129, 134 |
| Centrechinus mexicanus (A. Agassiz, 1863) 7, 9, 40, 65, 77, 135 |
| Diadema mexicanus A. Agassiz, 1863 31 |
| Order Arbacioida Gregory, 1900 |
| Family Arbaciidae Gray, 1855 |
| Genus Arbacia Gray, 1835 |
| Arbacia stellata (Blainville, 1825; ?Gmelin, 1788) NAY, JAL, MAR, ISA |
| Arbacia incisa (A. Agassiz, 1863) 9, 13, 21, 42, 43, 81, 84, 109, 110, 111, 113 |
| Order Camarodonta Jackson, 1912 |
| Family Echinometridae Gray, 1855 |
| Genus Echinometra Gray, 1825 |
| Echinometra insularis H.L. Clark, 1912 REV 37, 51, 100 |
| Echinometra oblonga (Blainville, 1825) REV 1, 7, 13, 15, 21, 31, 37, 40, 41, 45, 74, 79, 81, 91, 92, 95, 120, 124, 126, 134, 135 |
| Echinometra mathei oblonga (Blainville, 1825) 75 |
| Echinometra vanbrunti A. Agassiz, 1863 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 1, 7, 15, 31, 33, 41, 42, 43, 44, 45, 58, 73, 79, 81, 84, 87, 91, 92, 104, 107, 108, 109, 110, 111, 113, 120, 121, 124, 126, 129, 134 |
| Echinometra van brunti A. Agassiz, 1863 13, 21, 22, 29 |
| Echinometra Van brunti A. Agassiz, 1863 2, 9 |
| Echinometra van-brunti A. Agassiz, 1863 77 |
| Echinometra VanBrunti A. Agassiz, 1863 40 |
| Echinometra vanbruntii A. Agassiz, 1863 80 |
| Heliocidaris stenopora H.L. Clark, 19129 |
| Genus Heterocentrotus Brandt, 1835 |
| Heterocentrotus mamillatus (Linnaeus, 1758) REV 7, 32, 108 |
| Family Toxopneustidae Troschel, 1872 |
| Genus Toxopneustes L. Agassiz, 1841 |
| Toxopneustes roseus (A. Agassiz, 1863) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 13, 15, 21, 22, 29, 33, 40, 41, 42, 43, 44, 45, 58, 65, 73, 79, 81, 87, 91, 92, 96, 104, 107, 109, 110, 111, 112, 113, 121, 124, 126, 129 |
| Genus Tripneustes L. Agassiz, 1841 |
| Tripneustes depressus A. Agassiz, 1863 JAL, COL, MICH, MAR, ISA, MAT, REV 1, 7, 13, 15, 18, 20, 21, 31, 33, 40, 41, 42, 43, 44, 45, 65, 73, 75, 79, 80, 81, 91, 92, 99, 104, 107, 108, 109, 110, 111, 113, 120, 121, 124, 126, 129, 130, 134, 135 |
| Tripneustes gratilla (Linnaeus, 1758) REV 37 |
| Order Cassiduloida Claus, 1880 |
| Family Cassidulidae L. Agassiz & Desor, 1847 |
| Genus Rhyncholampas A. Agassiz, 1869 |
| Rhyncholampas pacificus (A. Agassiz, 1863) MAR, REV 93 |
| Cassidulus pacifica (A. Agassiz, 1863) 7, 91, 92 |
| Cassidulus pacificus (A. Agassiz, 1863) 13, 15, 26, 40, 101 |
| Order Clypeasteroida A. Agassiz, 1872 |
| Family Clypeasteridae L. Agassiz, 1835 |
| Genus Clypeaster Lamarck, 1801 |
| Clypeaster europacificus H.L. Clark, 1914 REV 7, 13, 15, 24, 40, 51, 81, 92, 120, 124, 135 |
| Clypeaster (Alexandria) europacificus H.L. Clark, 1914 102 |
| Clypeaster ochrus H.L. Clark, 1914 MAR, ISA, REV 13, 24, 39, 81, 92, 93, 120, 124, 126 |
| Clypeaster (Stolonoclypus) ochrus H.L. Clark, 1914 91 |
| Clypeaster rotundus (A. Agassiz, 1863) JAL, MAR, ISA, REV 7, 15, 24, 40, 43, 77, 92 |
| Clypeaster (Stolonoclypus) rotundus A. Agassiz, 1863 91, 102 |
| Clypeaster speciosus Verrill, 1870 NAY, COL, REV 7, 13, 15, 24, 39, 40, 41, 45, 77, 92, 126 |
| Clypeaster (Stolonoclypus) speciosus Verrill, 1870 91, 102 |
| Family Dendrasteridae Lambert, 1900 |
| Genus Dendraster L. Agassiz in Agassiz & Desor, 1847 |
| Dendraster excentricus (Eschscholtz, 1831) ISA 24 |
| Family Mellitidae Stefanini, 1912 |
| Genus Mellita L. Agassiz, 1841 |
| Mellita notabilis H.L. Clark, 1947 MICH 58, 67, 81, 104 |
| Genus Encope L. Agassiz, 1840 |
| Encope micropora L. Agassiz, 1841 JAL, MICH |
| Encope fragilis H.L. Clark, 194840 |
| Encope micropora fragilis H.L. Clark, 194824 |
| Encope perspectiva L. Agassiz, 1841 40, 58, 81, 104 |
| Encope wetmorei A.H. Clark, 1946 58, 81, 104 |
| Encope micropora insularis H.L. Clark, 1948 REV 24, 102 |
| Encope insularis H.L. Clark, 194813, 15, 40, 51 |
| Genus Lanthonia Coppard, 2016 |
| Lanthonia longifissa (Michelin, 1858) COL |
| Mellita longifissa Michelin, 1858 24, 81, 124 |
| Order Holasteroida Durham & Melville, 1957 |
| Family Plexechinidae Mooi & David, 1996 |
| Genus Plexechinus A. Agassiz, 1898 |
| Plexechinus cinctus A. Agassiz, 1898 MAR 3, 4, 13, 26, 51, 90, 92 |
| Order Spatangoida L. Agassiz, 1840 |
| Family Schizasteridae Lambert, 1905 |
| Genus Brisaster Gray, 1855 |
| Brisaster townsendi (A. Agassiz, 1898) MAR |
| Briaster townsendi (A. Agassiz, 1898) 81 |
| Brisaster towsendi (A. Agassiz, 1898) 13, 26 |
| Schizaster townsendi A. Agassiz, 189851 |
| Schizaster Townsendi A. Agassiz, 18983, 4 |
| Family Prenasteridae Lambert, 1905 |
| Genus Agassizia Valenciennes, 1846 |
| Agassizia scrobiculata Valenciennes, 1846 NAY 26, 81 |
| Family Brissidae Gray, 1855 |
| Genus Brissopsis L. Agassiz, 1840 |
| Brissopsis pacifica (A. Agassiz, 1898) REV 7, 13, 15, 26, 40, 91, 92 |
| Genus Brissus Gray, 1825 |
| Brissus latecarinatus (Leske, 1778) ISA 109, 110 |
| Brissus obesus Verrill, 1867 MAR 43 |
| Genus Meoma Gray, 1851 |
| Meoma ventricosa grandis Gray, 1851 JAL, REV 81, 91, 93, 120 |
| Meoma grandis Gray, 1851 7, 26, 40, 77, 92, 103 |
| Meoma ventricosa (Lamarck, 1816) 45, 124, 126 |
| Family Loveniidae Lambert, 1905 |
| Genus Lovenia Desor in Agassiz & Desor, 1847 |
| Lovenia cordiformis A. Agassiz, 1872 REV 13, 15, 26, 40, 41, 45, 77, 81, 92, 93, 124 |
| Lovenia cardiformis A. Agassiz, 18727 |
| Lovenia (Lovenia) cordiformis A. Agassiz, 187291 |
| Class Holothuroidea Selenka, 1867 |
| Order Dendrochirotida Grube, 1840 |
| Family Psolidae Burmeister, 1837 |
| Genus Lissothuria Verrill, 1867 |
| Lissothuria imbricataSolís-Marín, Arriaga-Ochoa, Galván-Villa, & Laguarda-Figueras, 2018 JAL 125 |
| Lissothuria ornata Verrill, 1867 JAL, MICH, MAT 104, 111, 113, 125, 127 |
| Thyonepsolus beebeiDeichmann, 193781 |
| Genus Psolus Oken, 1815 |
| Psolus pauperLudwig, 1893 MAR 85, 94 |
| Family Sclerodactylidae Panning, 1949 |
| Genus Afrocucumis Deichmann, 1944 |
| Afrocucumis ovulum (Selenka, 1867) NAY, JAL, MICH, MAT 104, 113, 116, 127 |
| Genus Neothyone Deichmann, 1941 |
| Neothyone gibber (Selenka, 1867) NAY, JAL, MICH, MAT, REV 7, 44, 47, 81, 84, 90, 104, 105, 111, 112, 113, 128 |
| Thyone gibber (Selenka, 1867) 13, 15 |
| Neothyone gibbosaDeichmann, 1941 JAL, MICH 58, 81, 104, 111, 113 |
| Genus Pachythyone Deichmann, 1941 |
| Pachythyone lugubris (Deichmann, 1939) JAL 111, 113 |
| Pachythyone pseudolugubrisDeichmann, 1941 JAL, MICH, MAT 104, 111, 113, 127 |
| Genus Neopentamera Deichmann, 1941 |
| Neopentamera anexiguaDeichmann, 1941 JAL 111, 113 |
| Genus Euthyonidiella Heding & Panning, 1954 |
| Euthyonidiella zacae (Deichmann, 1938) REV 120 |
| Phyllophorus zacae Deichmann, 1938 45, 81, 124 |
| Family Phyllophoridae Östergren, 1907 |
| Genus Pentamera Ayres, 1852 |
| Pentamera chierchiae (Ludwig, 1886) JAL, MICH, MAT, REV |
| Pentamera chierchia (Ludwig, 1886) 7, 13, 15, 45, 47, 81, 91, 92, 104, 111, 113, 124, 126, 128 |
| Genus Thyone Oken, 1815 |
| Thyone bidentataDeichmann, 1941 JAL 13, 47, 113, 119 |
| Thyone parafususDeichmann, 1941 JAL 13, 47, 90, 92, 119, 130 |
| Family CucumariidaeLudwig, 1894 |
| Genus Abyssocucumis Heding, 1942 |
| Abyssocucumis abyssorum (Théel, 1886) NAY 87 |
| Genus Cucumaria de Blainville, 1830 |
| Cucumaria flammaSolís-Marín & Laguarda-Figueras, 1999 NAY, JAL, COL, MICH, MAT 44, 58, 73, 81, 104, 107, 111, 113, 118, 121, 124, 128, 129 |
| Pattalus mollis Selenka, 1868 44, 105 |
| Genus Neocucumis Deichmann, 1944 |
| Neocucumis veleronis (Deichmann, 1941) JAL, COL, MICH, MAT 81, 104, 111, 113, 124, 127 |
| Genus Pseudocnus Panning, 1949 |
| Pseudocnus californicus (Semper, 1868) NAY, JAL, COL, MICH, MAT 44, 58, 81, 104, 105, 111, 113, 116, 124, 128 |
| Genus Trachythyone Studer, 1876 |
| Trachythyone peruana (Semper, 1868) NAY, JAL 81, 113 |
| Genus Thyonella Verrill, 1872 |
| Thyonella mexicana (Deichmann, 1941) MICH 104 |
| Genus Leptopentacta H.L. Clark, 1938 |
| Leptopentacta panamicaDeichmann, 1941 JAL 13, 47, 90 |
| Family Ypsilothuriidae Heding, 1942 |
| Genus Ypsilothuria Perrier, 1886 |
| Ypsilothuria bitentaculata (Ludwig, 1893) MAR 92, 94 |
| Shpaerothuria bitentaculataLudwig, 189385 |
| Order HolothuriidaMiller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 |
| Family Holothuriidae Burmeister, 1837 |
| Genus Holothuria Linnaeus, 1767 |
| Subgenus Cystipus Haacke, 1880 |
| Holothuria (Cystipus) inhabilis Selenka, 1867 REV 91, 126 |
| Holothuria inhabilis Selenka, 1867 7, 46, 92, 130 |
| Holothuria parinhabilis Cherbonnier, 1951 15 |
| Jaegerothuria inhabilis (Selenka, 1867) 77 |
| Holothuria (Cystipus) rigida (Selenka, 1867) NAY, JAL, COL, MICH, MAT 44, 58, 81, 104, 105, 113, 124 |
| Holothuria rigida (Selenka, 1867) 44, 84, 111 |
| Subgenus Halodeima Pearson, 1914 |
| Holothuria (Halodeima) atra Jaeger, 1833 NAY, JAL, ISA 81 |
| Holothuria atra Jaeger, 1833 109, 110 |
| Holothuria (Halodeima) inornata Semper, 1868 NAY, JAL, MICH, ISA, MAT, REV 104, 107, 113, 116, 119, 120, 126, 127 |
| Holothuria inornata Semper, 1868 13, 15, 111, 112, 129 |
| Holothuria (Halodeima) kefersteinii (Selenka, 1867) NAY, JAL, MICH, MAR, ISA, MAT, REV 104, 107 |
| Holothuria kefersteiini (Selenka, 1867) 43 |
| Holothuria kefersteini (Selenka, 1867) 44, 45, 84, 92 |
| Holothuria kefersteinii (Selenka, 1867) 129 |
| Holothuria kefersteni (Selenka, 1867) 7 |
| Holothuria (Halodeima) kefersteini (Selenka, 1867) 58, 81, 91, 105, 109, 110, 116, 119, 120, 124, 126 |
| Holothuria (Paraholothuria) riojai Caso, 1963 58 |
| Ludwigothuria kefersteini (Selenka, 1867) 48, 77 |
| Subgenus Lessonothuria Deichmann, 1958 |
| Holothuria (Lessonothuria) pardalis Selenka, 1867 JAL 91, 113 |
| Holothuria pardalis Selenka, 1867 13, 90 |
| Lessonothuria pardalis (Selenka, 1867) 48 |
| Subgenus Mertensiothuria Deichmann, 1958 |
| Holothuria (Mertensiothuria) hilla Lesson, 1830 MAR, ISA, REV 120, 126 |
| Brandtothuria gyrifer (Selenka, 1867) 48 |
| Holothuria gyrifer (Selenka, 1867) 13, 16 |
| Holothuria hilla Lesson, 1830 45, 111 |
| Holothuria (Thymiosycia) hilla Lesson, 1830 81, 109, 110, 124 |
| Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) COL, MAR, REV 81, 91, 120, 124, 126 |
| Holothuria leucospilota (Brandt, 1835) 7, 13, 15, 31, 41, 45, 79, 90, 92 |
| Mertensiothuria leucospilota (Brandt, 1835) 48 |
| Holothuria (Mertensiothuria) viridiaurantia Borrero-Pérez & Vanegas-González, 2019NAY, JAL, MICH, MAT |
| Holothuria hilla Lesson, 1830 111, 129 |
| Holothuria (Mertensiothuria) hilla Lesson, 1830 104, 113, 116, 127 |
| Holothuria (Thymiosycia) hilla Lesson, 1830 81 |
| Subgenus Platyperona Rowe, 1969 |
| Holothuria (Platyperona) difficilis Semper, 1868 NAY, JAL, MICH, MAR, MAT, REV 81, 91, 104, 116, 120, 124, 126, 127 |
| Holothuria difficilis Semper, 1868 7, 13, 15, 45, 46, 112, 129, 130 |
| Microthele difficilis (Semper, 1868) 48 |
| Subgenus Selenkothuria Deichmann, 1958 |
| Holothuria (Selenkothuria) carere Honey-Escandón, Solís-Marín, & Laguarda-Figueras, 2011MICH 82, 104 |
| Holothuria (Selenkothuria) lubrica Selenka, 1867 NAY, JAL, MICH, MAR, ISA, REV 58, 81, 91, 104, 109, 110, 113, 116, 120, 124, 126 |
| Holothuria lubrica Selenka, 1867 7, 12, 13, 15, 45, 58, 92, 111, 112, 121 |
| Holothuria pseudolubrica Cherbonnier, 1951 15 |
| Selenkothuria lubrica (Selenka, 1867) 41, 48 |
| Holothuria (Selenkothuria) portovallartensis Caso, 1954NAY, JAL, MICH, REV 58, 81, 91, 104, 116, 119, 126 |
| Holothuria portovallartensisCaso, 195412, 13, 92 |
| Selenkothuria portovallartensis (Caso, 1954) 48 |
| Subgenus Semperothuria Deichmann, 1958 |
| Holothuria (Semperothuria) imitans Ludwig, 1875 NAY, JAL, COL, MICH, ISA, MAT, REV 81, 91, 104, 105, 109, 110, 116, 120, 124, 126 |
| Holothuria imitans Ludwig, 1875 7, 13, 15, 44, 45, 92, 129 |
| Holothuria imitans var. polymorphaCaso, 196215 |
| Semperothuria imitans (Ludwig, 1875) 48, 77 |
| Holothuria (Semperothuria) languens Selenka, 1867 JAL, MICH, MAR 81, 104 |
| Holothuria languens Selenka, 1867 13 |
| Semperothuria languens (Selenka, 1867) 48 |
| Subgenus Stauropora Rowe, 1969 |
| Holothuria (Stauropora) fuscocinerea Jaeger, 1833 JAL, MICH, MAR, ISA, REV |
| Holothuria fuscocinerea Jaeger, 1833 45, 73, 78 |
| Holothuria fuscoscinerea Jaeger, 1833 43 |
| Holothuria (Mertensiothuria) fuscocinerea Jaeger, 1833 81, 109, 110, 124 |
| Holothuria (Stauropora) pluricuriosaDeichmann, 1937120, 126 |
| Subgenus Theelothuria Deichmann, 1958 |
| Holothuria (Theelothuria) paraprinceps Deichmann, 1937REV 126 |
| Holothuria paraprincepsDeichmann, 193792 |
| Theelothuria paraprinceps (Deichmann, 1937) 48, 77 |
| Subgenus Thymiosycia Pearson, 1914 |
| Holothuria (Thymiosycia) arenicola Semper, 1868 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 44, 58, 81, 91, 104, 105, 107, 109, 110, 113, 116, 119, 120, 124, 126 |
| Brandtothuria arenicola (Semper, 1868) 42, 48, 77 |
| Holothuria arenicola Semper, 1868 7, 15, 44, 45, 73, 84, 92, 111, 121, 129 |
| Holothuria (Thymiosycia) impatiens (Forskål, 1775) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 58, 81, 91, 104, 105, 107, 109, 110, 113, 116, 120, 124 |
| Brandtothuria impatiens (Forskål, 1775) 42, 48, 77 |
| Holothuria impatiens (Forskål, 1775) 7, 15, 43, 44, 45, 58, 84, 92, 111, 112, 124, 129 |
| Genus Labidodemas Selenka, 1867 |
| Labidodemas americanum Deichmann, 1938 NAY, JAL, COL, MICH, MAT, REV 44, 45, 81, 104, 105, 113, 116, 120, 124, 126, 129 |
| Labidodemas americanus Deichmann, 1938 58 |
| Labidodemas maccullochi (Deichmann, 1958) JAL, COL, MICH 111, 113 |
| Holothuria (Irenothuria) maccullochi (Deichmann, 1958) 58 |
| Labidodemas macullochi (Deichmann, 1958) 81, 124 |
| Order SynallactidaMiller, Kerr, Paulay, Reich, Wilson, Carvajal, & Rouse, 2017 |
| Family Stichopodidae Haeckel, 1896 |
| Genus Isostichopus Deichmann, 1958 |
| Isostichopus fuscus (Ludwig, 1875) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV 7, 42, 43, 44, 45, 48, 58, 72, 73, 81, 84, 91, 92, 104, 105, 107, 109, 110, 111, 112, 113, 119, 120, 121, 124, 126, 129, 133 |
| Isostichopus fuscis (Ludwig, 1875) 96 |
| Stichopus fuscus Ludwig, 1875 15, 17, 78 |
| Order Elasipodida Théel, 1882 |
| Family Laetmogonidae Ekman, 1926 |
| Genus Pannychia Théel, 1882 |
| Pannychia moseleyi Théel, 1882 JAL, COL, MAR 92, 94 |
| Laetmophasma fecundumLudwig, 189385 |
| Order Molpadida Haeckel, 1896 |
| Family Molpadiidae Müller, 1850 |
| Genus Molpadia Cuvier, 1817 |
| Molpadia musculus Risso, 1826 MAR 92 |
| Ankyroderma danielsseni Théel, 1886 85 |
| Order Apodida Brandt, 1835 |
| Family Chiridotidae Östergren, 1898 |
| Genus Chiridota Eschscholtz, 1829 |
| Chiridota aponocrita A.H. Clark, 1920 MICH 104 |
| Chiridota rigida Semper, 1867 NAY, MICH, MAT 44, 81, 104, 105 |
| Family Synaptidae Burmeister, 1837 |
| Genus Epitomapta Heding, 1928 |
| Epitomapta tabogae Heding, 1928 MICH 104 |
| Genus Euapta Östergren, 1898 |
| Euapta godeffroyi (Semper, 1868) NAY, JAL, COL, MICH, MAR, ISA, REV 41, 43, 45, 60, 73, 81, 109, 110, 113 |
| Genus Leptosynapta Verrill, 1867 |
| Leptosynapta albicans (Selenka, 1867) REV 7, 15, 90, 94 |
COL = Colima; ISA = Isabel Island; JAL = Jalisco; MAR = Marías Islands; MAT = Marietas Islands; MICH = Michoacán; NAY = Nayarit; REV = Revillagigedo Islands.
Invalid and doubtful records: The records considered as invalid (52) or doubtful (five) are based on: 1) erroneous or doubtful distributions, 2) incorrect taxonomic identifications, or 3) entries in faunal list not backed up by records in the primary literature.
There are some records considered invalid because of inconsistencies in their geographical distributions. The ophiuroids Ophiocomella ophiactoides and Astrophyton muricatum, and the holothuroid Isostichopus badionotus have been reported in the CMP (Honey-Escandón et al., 2008; Solís-Marín et al., 2016b; CONANP, 2019), yet the species distribution range is exclusively for the Caribbean and the Gulf of México (Hendler, Miller, Pawson, & Kier, 1995; Solís-Marín et al., 2013a), meanwhile the echinoids Echinothrix calamaris and Echinothrix diadema inhabits the Indian Ocean, but have been reported in REV (Sotelo-Casas & Rodríguez-Troncoso, 2014). The holothuroid Pentamera chiloensis is reported in Barra de Navidad, JAL (Rodríguez-Troncoso et al., 2013); this species distributes only in South America (Perú and Chile; Solís-Marín et al., 2013a). Records previously mentioned correspond to erroneous records and must be invalid (R.C. Sotelo-Casas and F.A. Solís-Marín, personal communication, 01 March 2020).
The crinoid Hyocrinus foelli (Solís-Marín et al., 2013a; Solís-Marín et al., 2013b; Solís-Marín, Laguarda-Figueras, & Honey-Escandón, 2014) and the holothuroid Psychronaetes hanseni (Maluf, 1991; Bautista-Romero et al., 1994; Massin & Hendrickx, 2011) have been reported in Clarion Island, REV, but according to their original descriptions (Pawson, 1983; Roux & Pawson, 1999) and the validation of the geographical coordinates, all the records corresponds to the Clarion-Clipperton Fracture Zone which is located approximately 1 200 km far away from REV. The echinoids Aporocidaris milleri and Kamptosoma asterias were reported in NAY (station 96, 2 988-3 001 m depth) by Parker (1963), yet the validation of the geographic coordinates revealed that this station is located in the entrance of the Gulf of California, and far away from the maritime zone of NAY. Moreover, K. asterias is distributed in the Central Pacific Ocean, Chile and the Antarctica (Mooi, Constable, Lockhart, & Pearse, 2004).
Furthermore, some records had inconsistencies in their taxonomic identifications. Ophionereis albomaculata (ophiuroid) has records in ISA and MAR (Caso, 1961; Maluf, 1991; Maluf & Brusca, 2005; Granja-Fernández et al., 2015a; Granja-Fernández et al., 2017); during our revision of museum collections, we taxonomically reviewed all the material known for O. albomaculata (Galápagos Islands and Panamá), and none corresponded to México. The material identified as O. albomaculata from México must relate to another species of Ophionereis. The collected material of the holothuroid Holothuria (Lessonothuria) pardalis from REV (Solís-Marín et al., 2013a; Sotelo-Casas & Rodríguez-Troncoso, 2014), was previously identified as H. (Lessenothuria) pardalis, yet it corresponds to a new species in process of description (F.A. Solís-Marín, personal communication, 01 April 2020). It is important to mention that in the CMP, the species only distributes in Tenacatita, JAL (Deichmann, 1958).
Most of the entries are based on unbacked up records of primary references. Bautista-Romero et al. (1994) constructed a checklist of echinoderms from REV according to previous records (Fisher, 1911; Ziesenhenne, 1937; Caso, 1962b; Maluf, 1988), and reported: Florometra serratissima (crinoid); Amphiaster insignis, Astropecten verrilli, Echinaster (Othilia) tenuispina, Heliaster kubiniji, Henricia aspera, Henricia asthenactis, Henricia leviuscula, Henricia polyacantha, Leiaster teres, Luidia foliolata, Luidia phragma, Mediaster aequalis, Narcissia gracilis, Odontaster crassus, Tethyaster canaliculatus, and Thrissacanthias penicillatus (asteroids); Amphiodia (Amphispina) urtica, Amphipholis squamata, Amphiura arcystata, Astrocaneum spinosum, Diopederma daniana, Ophiolepis crassa, Ophionereis eurybrachiplax, Ophiopaepale diplax and Ophiura luetkenii (ophiuroids); and Arbacia stellata (as Arbacia incisa), Centrostephanus coronatus, Encope grandis, Encope micropora, Lytechinus pictus, and Strongylocentrotus fragilis (echinoids). We consulted the previous literature and we did not find these records on them or in other literature references, therefore these records are invalid.
Based on literature records, Lophaster furcilliger (asteroid), Astrocaneum spinosum (ophiuroid), and Centrostephanus coronatus (echinoid) have been recorded in the program of management and conservation of REV (CONANP, 2019). The only records of L. furcilliger in the CMP correspond to MAR (Ludwig, 1905; Maluf, 1988; Maluf, 1991), therefore their inclusion in REV is incorrect. Despite it is very probable that A. spinosum and C. coronatus distributes in REV, we did not find any record in publications to back it up. Other species reported in a program of management and conservation (ISA; CONANP, 2005) are the echinoids Clypeaster europacificus and Lovenia cordiformis, but again no primary references supporting these records were found, and therefore are considered as invalid records.
Finally, based on a revision of literature, the asteroids Paulia horrida and Luidia bellonae were reported in MAR by Maluf (1988) and Maluf and Brusca (2005), respectively; nevertheless, we did not find any record of these species for MAR but for REV. Similarly, the asteroid Cnemidaster nudus has only one mention for COL (Solís-Marín et al., 2016b), but it corresponds to an erroneous record (F.A. Solís-Marín, personal communication, 01 April 2020). For these reasons, we consider both records as invalid.
The following records were considered as doubtful. The ophiuroid Amphiodia periercta inhabits in the North Pacific Ocean (from Aleutian Islands, Alaska to central California; Lambert & Austin, 2007), yet it has a report in Clarion Island, REV (Maluf, 1988; Bautista-Romero et al., 1994). Due to the northern affinity of A. periercta, its distribution in REV is unlikely. The record of Hymenaster pentagonalis in REV was obtained by the Expedition Ocean Exploration Trust/Nautilus Live 2017 (CONANP, 2019), yet this species distributes only in Hawaii (Mah, 2020).
The echinoids Caenocentrotus gibbosus (Clark, 1925; Grant & Hertlein, 1938; Maluf, 1988; Maluf,1991; Bautista-Romero et al., 1994) and Lytechinus semituberculatus (Clark, 1925; Clark,1948; Mortensen, 1943b; Caso, 1962b; Maluf, 1988; Bautista-Romero et al., 1994) have many mentions in Clarion Island, REV, yet in all these references the locality is treated as doubtful. It is important to emphasize that both species inhabit in South America (Galápagos Islands, Perú, and Chile; Grant & Hertlein, 1938; Solís-Marín et al., 2013a), thus their distribution in México is unlikely. Similarly, the holothuroid Holothuria (Selenkothuria) theeli is recorded from REV as doubtful (Hertlein, 1963; Maluf, 1991; Maluf & Brusca, 2005). In the Eastern Pacific, H. (Selenkothuria) theeli inhabits in lower latitudes (i.e. Costa Rica-Perú; Solís-Marín et al., 2013a) and in México it has been collected only in Guerrero (Honey-Escandón et al., 2008). For these reasons, we determined these records as doubtful.
New records: Following review of museum material, we add 16 new records of Ophiuroidea to different states (four NAY, one COL, and one MICH) and islands (two MAR, four ISA, and four REV) from the CMP (appendix 1). All the species were previously recorded in areas from the CMP (Table 1), excepting Amphiodia psara, Amphiodia tabogae, Microphiopholis geminata, and Ophiophragmus paucispinus, which represent new records for the CMP.
Amphiodia psara has previous records in USA (California; Clark, 1935) and México (Gulf of California and Guerrero; Maluf, 1988; Maluf & Brusca, 2005; Granja-Fernández et al., 2015b). After the taxonomic revision of A. psara in different museum collections, we did not find material from Guerrero, but for the Pacific of Baja California and Baja California Sur, the Gulf of California (Ángel de la Guarda Island, Tiburón Island, Consag Rock, and Sonora), and NAY. The records from Baja California, Baja California Sur and NAY are new, being Punta Mita, Nayarit, the southernmost area of its geographical distribution.
Amphiodia tabogae, M. geminata, and O. paucispinus distribute along the Eastern Pacific (México to Galápagos Islands; Solís-Marín et al., 2013a). Specifically, in México, the three species inhabit the Gulf of California; A. tabogae has been reported also in Guerrero (Granja-Fernández et al., 2015b). The finding of these species in Punta Mita, NAY (A. tabogae and M. geminata), and in MAR (O. paucispinus), represents new records in the CMP, and fills the distribution gap along the Mexican Pacific.
Diversity and distribution: The diversity of the Echinodermata from the CMP is represented by 187 species of echinoderms belonging to 27 orders, 63 families, 110 genera, and 16 subgenera (Table 2). Compared to the total number of echinoderm species worldwide (7 437 species; WoRMS, 2020), the CMP shelters 2.5 %. The orders with the highest number of species in the CMP were Amphilepidida (Ophiuroidea, 40 species), Holothuriida and Dendrochirotida (Holothuroidea, 21 species, respectively), Ophiacanthida (Ophiuroidea, 16 species), and Valvatida (Asteroidea, 15 species), meanwhile the best represented families were Amphiuridae (Ophiuroidea, 26 species), and Holothuriidae (Holothuroidea, 21 species) (Table 1). On the other hand, the orders Ophiurida (45 species), Aspidochirotida (22 species), and Dendrochirotida (22 species) were the best represented in the Southern Mexican Pacific (SMP; Granja-Fernández et al. 2015b). Differences in the numbers of recorded orders between the CMP and the SMP are due to recent changes in the systematics of the Ophiuroidea (O´Hara, Stöhr, Hugall, Thuy & Martynov, 2018) in which most of the members previously included in the order Ophiurida, now are in Amphilepidida, and in the Holothuroidea, where most of the members of Aspidochirotida were changed to Holothuriida (Miller et al., 2017). If we compare both faunas in the same classification system, similar trends are shown, suggesting that both areas possess similar faunas in terms of orders and families of echinoderms.
Table 2 Taxonomic categories of echinoderms from the Central Mexican Pacific
| Class | Orders | Families | Genera | Subgenera | Species |
| Crinoidea | 2 | 2 | 2 | 0 | 3 |
| Asteroidea | 5 | 16 | 24 | 2 | 35 |
| Ophiuroidea | 6 | 20 | 35 | 4 | 67 |
| Echinoidea | 8 | 14 | 23 | 0 | 32 |
| Holothuroidea | 6 | 11 | 26 | 10 | 50 |
| TOTAL | 27 | 63 | 110 | 16 | 187 |
In the CMP, the class Ophiuroidea was the most numerous with 67 species, followed by Holothuroidea (50), Asteroidea (35), Echinoidea (32), and Crinoidea (three) (Table 2). In general, the number of species of Holothuroidea, Asteroidea and Echinoidea were similar between the CMP and the SMP, but Ophiuroidea was represented by more species in the CMP (67) than to the SMP (46); meanwhile, the class Crinoidea had not records in the SMP (Granja-Fernández et al., 2015b). Differences on the Ophiuroidea are related to the recent sampling effort directed in the CMP (Granja-Fernández et al., 2017), and the extensive revision of samples from museum collections (reflected in the new records of species in this work). Nine species of Crinoidea from México to Perú (Solís-Marín et al., 2013a) and at least seven in the northern part of the American continent (Lambert & Austin, 2007) are reported in the literature therefore, finding crinoids in the CMP and SMP is likely. Despite the above, a null to a scarce number of species of Crinoidea are reported in both areas. We consider that the explanation of this pattern (and in general of the Eastern Pacific) is not due to the natural biogeographic distribution of the Crinoidea in the Eastern Pacific, yet to the scarce prospection in deep waters (the natural environment of the class in the Eastern Pacific), as well as to the low interest and lack of taxonomical specialists in the class.
As previously mentioned, the CMP is represented by 187 species of echinoderms; of those, 122 inhabit in ~ 640 km of the coastline of the studied states, and 142 in an extensive maritime area of the islands (~ 155 000 km2). The SMP has not extensive islands such as the CMP, therefore if only their coastlines are compared, the latter possesses a lower number of echinoderms species than the SMP (162 species; Granja-Fernández et al., 2015b). In order to have a better comprehension of the number of species between both areas, a standardization per area is required, therefore considering the whole CMP (~ 640 km) has 5.2 species/km, and the SMP (~ 1 200 km) possesses 7.4 species/km. The collecting effort in more areas, habitats, and depths could explain higher biodiversity of echinoderms in the coastline of CMP, but further collections of material and biogeographic analysis are required in order to better understand their distribution pattern.
In terms of spatial distribution, the coast of the state of JAL presented the highest number of species (84), followed by the coast of NAY (74), MICH (63), and COL (55); among the islands, REV showed the major number of species (85) followed by MAR (81), MAT (48), and ISA (44) (Table 3). Compared to the most complete and recent works of echinoderms by states (Honey-Escandón et al., 2008; Rodríguez-Troncoso et al., 2013; Solís-Marín et al., 2016b; Granja-Fernández et al., 2017; Ríos-Jara et al., 2017; Nava-Bravo et al., 2019) and islands (CONANP, 2005; CONANP, 2007a; CONANP, 2007b; CONANP, 2019; Honey-Escandón et al., 2008; Solís-Marín et al., 2013a; Granja-Fernández et al., 2017), this work increases the number of recorded species in 27 % for NAY, 49 % for JAL, 1.5 % for MICH, 70 % for MAR, 64 % for ISA, 48 % for MAT, 38 % for REV, and finally 45 % for the entire CMP region. We found the same number of species for COL as its most recent account of echinoderms (Solís-Marín et al., 2016b). Differences in the numbers of species among previous studies and the present one is attributed to a more intense literature review, the validation of geographical coordinates, the correction of records, and the addition of new ones.
Table 3 Number of species of echinoderms from the Central Mexican Pacific
| NAY | JAL | COL | MICH | MAR | ISA | MAT | REV | |
| Crinoidea | 0 | 0 | 2 | 0 | 1 | 0 | 0 | 0 |
| Asteroidea | 13 | 10 | 13 | 11 | 12 | 8 | 6 | 19 |
| Ophiuroidea | 33 | 27 | 18 | 12 | 39 | 13 | 16 | 25 |
| Echinoidea | 7 | 12 | 9 | 10 | 14 | 12 | 7 | 21 |
| Holothuroidea | 21 | 35 | 13 | 30 | 15 | 11 | 19 | 20 |
| TOTAL | 74 | 84 | 55 | 63 | 81 | 44 | 48 | 85 |
COL = Colima; ISA = Isabel Island; JAL = Jalisco; MAR = Marías Islands; MAT = Marietas Islands; MICH = Michoacán; NAY = Nayarit; REV = Revillagigedo Islands.
JAL, MAR and REV were the areas with the highest number of species and have been prospected in several shallow and deep-water localities/stations since late 1800´s by foreign (Albatross, Velero, and Zaca; e.g. Agassiz, 1898; Clark, 1940; Deichmann, 1958) and national expeditions (e.g. Caso, 1943b; Caso, 1961; Caso, 1962b). In contrast, the state of COL, as well as ISA and MAT islands have the lowest number of species, and their poor prospections initiated in the late 1900´s (e.g. Caso, 1961; Caso, 1980), and have been mostly performed during the 2000´s, strictly in shallow-waters (e.g. Chávez-Dagostino et al., 2000; Ríos-Jara et al., 2008a, Ríos-Jara et al., 2013; Sotelo-Casas et al., 2018). On the other hand, MAR and REV (the unique oceanic islands of the CMP) display unique characteristics such as particular oceanographic and topographic conditions, high productivity, mesophotic reefs, great depths, among others, which promote high levels of general marine biodiversity and endemism (CONANP, 2007a; CONANP, 2019), explaining their highest number of echinoderms in the region. In conclusion, the obtained data shows that the number of species of echinoderms in the CMP region is mostly associated with a sampling effort, yet specific conditions of particular areas (i.e. MAR and REV) can be influencing their high biodiversity, but more collections of material and further biogeographic analysis are required in order to sustain this statement.
Final considerations: The CMP is located in an oceanographic transitional zone, therefore fauna from the North and South regions of the Mexican Pacific (Arriaga-Cabrera et al., 1998), the Eastern Pacific (e.g. Solís-Marín et al., 2013a), and even of higher latitudes (e.g. Lambert & Austin, 2007), can inhabit in the region. In this sense, MAR and REV are important as they are considered a stepping stone to import organisms from the Indo-Pacific species into the Tropical Eastern Pacific (Reyes-Bonilla & López-Pérez, 1998), as well as in the connection for dispersal species to and from northern (Gulf of California) and the mainland CMP (López-Pérez et al., 2015).
All the islands considered in the present work possess their own programs of management and conservation (CONANP, 2005; CONANP, 2007a; CONANP, 2007b; CONANP, 2019) in which detailed descriptions of geographical, physical, demographical, social and environmental characteristics (among others) are provided. Despite the above, we detected that all these programs underestimated the number of species of echinoderms since they reported 16 species for ISAB (CONANP, 2005), 24 for MAR (CONANP, 2007a), 25 for MAT (CONANP, 2007b), and 52 for REV (CONANP, 2019). The inclusion of all the species reported in the present work on the lists of these natural protected areas will further emphasize the importance of the islands.
Due to the importance of the islands of the CMP, most of the research of echinoderms in the region have been conducted on them (e.g. Ríos-Jara et al., 2008a; Sotelo-Casas et al., 2018) relegating the mainland of the states of NAY, JAL, COL, and MICH, which remain almost unstudied and which have a high potential of finding numerous species of echinoderms since they have complex environments (extensive bays, coastal lagoons, estuaries, mangroves and coral reefs; Arriaga-Cabrera et al., 1998). Regardless the differences on the diversity of echinoderms and exploration of the mainland and islands from the CMP, all of them are important due to 34 species collected in the region have been described as new species (five Asteroidea, 16 Ophiuroidea, five Echinoidea, and eight Holothuroidea; Table 4). This highlights the potential of the CMP in finding more new species. Although the present manuscript provides the most-completed checklist of the echinoderms in the CMP, we encourage to intensify research in shallow and deep-waters and across different habitats of all the islands and mainland of the states of the region in order to better understand its biodiversity and distributional patterns. Finally, the preservation of the marine fauna of the entire CMP is key in the maintenance of the entire Mexican and Tropical Pacific; therefore, we encourage ensuring its effective protection through inventories of marine and coastal species.
Table 4 Described species and subspecies of Echinodermata based on material collected in the Central Mexican Pacific
| Class | Species | Locality | Reference |
| Asteroidea | Ampheraster marianus | Marías Islands | Ludwig, 1905 |
| Astropecten regalis | San Blas, Nayarit | Gray, 1840 | |
| Henricia gracilis | Marías Islands | Ludwig, 1905 | |
| Henricia seminudus | Revillagigedo Islands | Clark, 1916 | |
| Psilaster armatus | Marías Islands | Ludwig, 1905 | |
| Ophiuroidea | Amphichondrius laevis | Tenacatita Bay, Jalisco | Ziesenhenne, 1940 |
| Amphiodia sculptilis | Tenacatita Bay, Jalisco | Ziesenhenne, 1940 | |
| Amphiura serpentina | Marías Islands | Lütken & Mortensen, 1899 | |
| Astrodia excavata | Marías Islands | Lütken & Mortensen, 1899 | |
| Dougaloplus notacanthus | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiacantha costata | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiacantha hirta | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiacantha moniliformis | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiacantha pyriformis | Revillagigedo Islands | Ziesenhenne, 1937 | |
| Ophiernus adspersus annectens | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiolepis plateia | Tenacatita Bay, Jalisco | Ziesenhenne, 1940 | |
| Ophiomitra granifera | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiomitra partita | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiophragmus lonchophorus | Tenacatita Bay, Jalisco | Ziesenhenne, 1940 | |
| Ophiosphalma variabile | Marías Islands | Lütken & Mortensen, 1899 | |
| Ophiura (Ophiura) scutellata | Marías Islands | Lütken & Mortensen, 1899 | |
| Echinoidea | Brisaster townsendi | Marías Islands | Agassiz, 1898 |
| Echinometra insularis | Revillagigedo Islands | Clark, 1912 | |
| Encope micropora insularis | Revillagigedo Islands | Clark, 1948 | |
| Hesperocidaris perplexa | Revillagigedo Islands | Clark, 1907 | |
| Plexechinus cinctus | Marías Islands | Agassiz, 1898 | |
| Holothuroidea | Cucumaria flamma | Colima, Nayarit, Marietas Islands | Solís-Marín & Laguarda-Figueras, 1999 |
| Lissothuria imbricata | Jalisco | Solís-Marín et al., 2018 | |
| Holothuria (Selenkothuria) carere | Michoacán | Honey-Escandón et al., 2011 | |
| Holothuria (Selenkothuria) portovallartensis | Puerto Vallarta, Jalisco | Caso, 1954 | |
| Psolus pauper | Marías Islands | Ludwig, 1894 | |
| Thyone bidentata | Tenacatita Bay, Jalisco | Deichmann, 1941 | |
| Thyone parafusus | Tenacatita Bay, Jalisco | Deichmann, 1941 | |
| Ypsilothuria bitentaculata | Marías Islands | Ludwig, 1894 |
Ethical statement: authors declare that they all agree with this publication and made significant contributions; that there is no conflict of interest of any kind; and that we followed all pertinent ethical and legal procedures and requirements. All financial sources are fully and clearly stated in the acknowledgements section. A signed document has been filed in the journal archives.
