The phylum Echinodermata is represented approximately by 7 000 living species, and 13 000 fossil species, and comprises five living classes (Pawson, 2007). In the Mexican Pacific, the taxonomy and ecology of the echinoderm species have been studied for more than a hundred years, being the Gulf of California the most extensively studied area (Solís-Marín et al., 2013).

The Southern Mexican Pacific (SMP) comprises the States of Guerrero, Oaxaca and Chiapas. This is a region with a high marine biodiversity and endemism, besides it has a large number of habitats (i.e. coral reefs, mangroves, coastal lagoons) and represents the main dispersal route to Mexico for species from Central America (Arriaga-Cabrera et al., 1998; Reyes-Bonilla & López-Pérez, 1998). A large amount of information has been published regarding the echinoderm fauna of the SMP, resulting in a research history that can be divided into three periods.

The first expedition carried out at the coasts of the SMP was Albatross in 1891. The leading scientist of the expedition was Alexander Agassiz, who collected marine specimens from the Eastern Pacific. The expedition resulted in the reports of Ludwig (1894) (Holothuroidea), Agassiz (1898, 1904) (Echinoidea), Lütken and Mortensen (1899) (Ophiuroidea) and Ludwig (1905) (Asteroidea). Although Albatross was the only expedition into the SMP during the 19th century, numerous taxonomic contributions were produced for this area mostly for Asteroidea (Gray, 1871; Perrier, 1875, 1878; Sladen, 1899) and Echinoidea (Agassiz, 1863, 1872, 1873, 1881, 1898; Lockington, 1876; Rathbun, 1886).

During the 20th century (in 1904 and 1905) the Albatross made a final expedition to the SMP. This expedition resulted in the extensive work of H.L. Clark (1920) about the Eastern Tropical Pacific Asteroidea. Along with the Albatross, during 1931-1941, the Velero III surveyed the SMP. As a result of the Velero III expeditions, Deichmann (1941, 1958) (Holothuroidea), H.L. Clark (1948) (Echinoidea) and Ziesenhenne (1940) (Ophiuroidea) published extensive works on the echinoderms of the area. During 1937-1938, the New York Zoological Society organized an expedition to the Eastern Pacific (Zaca) under the direction of William Beebe, which generated the works of Deichmann (1938) (Holothuroidea) and H.L. Clark (1940) (Asteroidea, Ophiuroidea, Echinoidea). In addition, during this century, there was an increase in systematic studies of echinoderms from the SMP. In particular, researchers such as H.L. Clark (1907a, 1907b, 1914, 1915, 1917, 1920, 1922, 1925, 1940, 1948), Deichmann (1936, 1937, 1938, 1941, 1958) and Caso (1944, 1945, 1946, 1948a, 1948b, 1949, 1951, 1954, 1958, 1962, 1963, 1964, 1965, 1967a, 1967b, 1970, 1977, 1978a, 1978b, 1979, 1980a, 1980b, 1983, 1986) produced a vast number of publications.

Recently, the study of echinoderms in the Southern Mexican Pacific has not only focused on systematics, but also on ecology (Benítez-Villalobos, 2001; Lirman et al., 2001; Calderón-Aguilera & Reyes-Bonilla, 2006; López-Pérez et al., 2008; Zamorano & Leyte-Morales, 2005a, 2005b, 2009), bioerosion (Herrera-Escalante et al., 2005; Benítez-Villalobos et al. 2008b), reproduction (Benítez-Villalobos & Martínez-García, 2012; Benítez-Villalobos et al., 2012, 2013), and symbiosis (Granja-Fernández et al., 2013b).

Notwithstanding that the knowledge of the echinoderms in the SMP has increased since the 19th century, the large amount of work in the area has generated a myriad of names that not only have often been misspelled, but have also created longstanding taxonomic problems. To overcome this trend, the main goal of this contribution is to present the first updated checklist of all valid and synonymous names of echinoderms from the SMP based on published literature and from specimens deposited in collections, which will be useful as a baseline for future taxonomic, ecological and marine resource management studies.

MATERIALS AND METHODS

We present a list of all known valid names and synonyms of echinoderm species (classes Asteroidea, Ophiuroidea, Echinoidea and Holothuroidea) from Guerrero (GRO), Oaxaca (OAX), Chiapas (CHIS) and the Gulf of Tehuantepec (GT), México. The Gulf of Tehuantepec comprises the States of Oaxaca and Chiapas; some of the records in the area do not mention a specific State, thus we refer to them as Gulf of Tehuantepec.

The checklist is based on an exhaustive published literature search (131 references) and records of specimens deposited in museums or properly curated reference collections (AMNH-American Museum of Natural History, New York; AMS-Australian Museum, Sydney; CAS-California Academy of Sciences, San Francisco; FMNH-The Field Museum, Chicago; ICML-UNAM-Colección Nacional de Equinodermos “Dra. Ma. Elena Caso”, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, México City; LACM-Los Angeles County Museum, Los Angeles; MACN-Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires; MCZ-Museum of Comparative Zoology, Cambridge; UMAR-Colección de equinodermos de la Universidad del Mar, Puerto Ángel, México; USNM-National Museum of Natural History, Washington; YPM-Yale Peabody Museum of Natural History, New Haven).

Systematic arrangements follow the criteria of A.M. Clark (1989, 1993, 1996), A.M. Clark and Mah (2001) and Mah (2014) for the Asteroidea; Smith et al. (1995), Stöhr et al. (2014) for the Ophiuroidea; Mortensen (1928, 1935, 1943a, 1943b, 1948, 1950, 1951), Kroh and Smith (2010) and Kroh and Mooi (2014) for the Echinoidea; Pawson and Fell (1965), Rowe (1969), Solís-Marín et al. (2009), Smirnov (2012) and WoRMS (2014) for the Holothuroidea. The taxonomic status of all species (valid names and synonyms) was tracked in the literature back to the original description, and was constructed with the help of systematics experts. The current checklist includes only those synonyms that have been used in the SMP, but some species can have a larger synonym list in other geographic areas.

Results and discussion

Historical review

Asteroidea: A total of 39 references addressed the Asteroidea in the SMP. During the 19th century, only six works were written; in the 20th century 24 references were published and during the current one, nine works have been published (Fig. 1A). The authors who contributed with the highest number of works were H.L. Clark (1907b, 1920, 1940), Caso (1944, 1945, 1948b, 1970, 1977, 1979) and A.M. Clark (1989, 1993, 1996).

The study of Asteroidea in the SMP began in 1867 when Verrill (1867a, 1867b) mentioned the presence of Phataria unifascialis in Acapulco, Guerrero. Four years later, Gray (1871) described Luidia latiradiata as Platasterias latiradiata for the Gulf of Tehuantepec, which was thought to be a living fossil species of Somasteroidea, an extinct group from the Orvidician (Fell, 1962b). The species was further transfered to Luidiidae and no longer considered as a survivor member of this archaic group (Blake, 1972). Based on specimens donated by the Museum of Comparative Zoology, Cambridge to the Museum of Natural History of Paris, Perrier (1875) recorded Nidorellia armata, Pharia pyramidatus, Heliaster kubiniji and Heliaster microbrachius from Acapulco, Guerrero.

The 20th century was the period with the highest addition of Asteroidea to the SMP fauna (20 species). Ludwig (1905) described Eremicaster pacificus, Pseudarchaster pulcher, Mediaster transfuga, Hymenaster violaceus, Zoroaster hirsutus and Freyella insignis from Acapulco and other areas of the Tropical Eastern Pacific. Ludwig (1905) also reported Porcellanaster waltharii from Acapulco, now considered as a synonym of Eremicaster crassus (A.M. Clark, 1989). It is important to note that P. waltharii has been incorrectly spelled as P. waltherii (A.M. Clark, 1989), in contrast with the original spelling (Ludwig, 1905). Döderlein (1936) added Pentaceraster cumingi from Acapulco to the list, which had also been named as Oreaster occidentalis in some studies (Clark H.L., 1940; Salcedo-Martínez et al., 1988). Caso (1977) described Asterina agustincasoi based on specimens collected in Ixtapa Island (Guerrero) but O´Loughlin (2002) synonymized this species with Meridiastra modesta. Luke (1982) published a catalog of the benthic invertebrate collections of the SCRIPPS institution of oceanography and recorded for the first time Ceramaster leptoceramus for the SMP, which was collected in the Gulf of Tehuantepec. Salcedo-Martínez et al. (1988) published an inventory of benthic macroinvertebrates in rocky bottoms of Zihuatanejo (Guerrero). This work represented the first inventory of marine fauna from Guerrero and added the species Mithrodia bradleyi. Years later, Solís-Marín et al. (1993) recorded Luidia bellonae, Luidia columbia, Linckia columbiae, Astropecten ornatissimus and Heliaster helianthus for Guerrero and Oaxaca. The 20th century era concluded with the work of Gamboa-Contreras and Tapia-García (1998) from the Gulf of Tehuantepec, the authors included the record of Luidia foliolata, Astropecten armatus, Astropecten regalis and Astropecten verrilli. It is important to emphasize that A. regalis has been reported in the Greater Antilles, the Caribbean, the Gulf of México as well as in areas of the Eastern Pacific such as Costa Rica, Panamá and México (Solís-Marín et al., 2005; Honey-Escandón et al., 2008; Pawson et al., 2009; Alvarado et al., 2010; Miloslavich et al., 2010). Astropecten verrilli was called Astropecten californicus (e.g. Verrill, 1914; Honey-Escandón et al., 2008); but according to Döderlein (1917) the valid species is the former.

So far, the current century is the era with the lowest number of new records of Asteroidea for the SMP. A.M. Clark and Mah (2001) reported Heliaster polybrachius, while Benítez-Villalobos et al. (2008a) and Honey-Escandón et al. (2008) added Luidia superba, Luidia tessellata and Asteropsis carinifera The species Gymnasteria spinosa has been reported in the SMP as Asteropsis spinosa (Benítez-Villalobos et al., 2008a; Honey-Escandón et al. 2008), but according to Perrier (1875) the valid species is A. carinifera. According to Benítez-Villalobos et al. (2008a) and Bastida-Zavala et al. (2013), Leiaster teres inhabits Oaxaca (USNM E6635). This specimen was collected by Ricketts between 1940-1943 in Puerto Escondido during an expedition conducted in the Gulf of California; therefore the record is not valid for Oaxaca.

All species of Asteroidea that have been collected from the SMP are deposited in all the consulted museums, except the YPM. The collections with a major number of deposited species were ICML-UNAM (13), LACM (9) and the USNM (9) (Table 1).

 

Ophiuroidea: The Ophiuroidea is represented by the lowest number of publications (27) in the SMP. In the 19th century there were published only four works, in the 20th century, 15 and in the 21st century, eight (Fig. 1B). The authors who have largely contributed to the study of Ophiuroidea in the area were Lyman (1860), Verrill (1867a, 1867b), Lütken and Mortensen (1899), Clark (1915, 1940), Boone (1926, 1938), Tortonese (1933a), Ziesenhenne (1937, 1940), Caso (1951, 1979, 1986), Downey (1969), Rowe and Pawson (1977), Luke (1982), Salcedo-Martínez et al. (1988), Gamboa-Contreras and Tapia-García (1998), Benítez-Villalobos (2001), Zamorano and Leyte-Morales (2005a), Honey-Escandón et al. (2008), Granja-Fernández and López-Pérez (2011, 2012), Benítez-Villalobos et al. (2012) and Granja-Fernández et al. (2013b, 2014).

The brittle star Ophiocoma alexandri was the first ophiuroid reported in the SMP (Lyman, 1860). Ophioderma teres, Ophioderma panamensis and Ophiocoma aethiops were recorded in the SMP by Verrill (1867a, 1867b), since then the species have been frequently reported by a large number of authors (Table 1). One of the most seminal works on Eastern Pacific Ophiuroidea is that of Lütken and Mortensen (1899) who described almost 50 species for this area. They described six new species (Ophiacantha moniliformis, Amphiophiura superba, Ophiernus adspersus annectens, Amphiura serpentina, Dougaloplus gastracanthus, Ophiomusium variabile, all of them collected in deep waters (902 to 4 082 m depth) from Guerrero, and they also documented for the first time in the area the cosmopolitan Ophiactis savignyi. The species Dougaloplus gastracanthus was originally described as Amphiura gastracanthus, nevertheless Clark (1970) transferred the species to the genus Dougaloplus.

During the 20th century, a larger number of papers were published about Ophiuroidea in the SMP. Clark (1915) reported for the first time Ophiernus seminudus, Ophiothrix (Ophiothrix) spiculata and Ophionereis annulata for this area. Twenty five years later, Clark (1940) added seven reports of brittle stars (Amphiura arcystata, Amphichondrius granulatus, Amphiodia (Amphispina) urtica, Ophioderma variegata, Diopederma daniana, Ophiolepis fulva and Ophiolepis variegata) to the area. Ziesenhenne (1940) described an important number of new species of Ophiuroidea (19) from the Eastern Pacific. He described the species Ophiomisidium leurum, Amphichondrius laevis, Amphiodia sculptilis and Ophiophragmus papillatus for the SMP (Guerrero and Oaxaca) in depths up to 91 m. Caso (1951) recorded Ophiothrix (Ophiothrix) rudis and Ophiolepis pacifica in Zihuatanejo, Guererro. Luke (1982) added Ophiura (Ophiuroglypha) irrorata irrorata, Amphiura seminuda, Stegophiura ponderosa, Ophiactis simplex and Ophiomusium glabrum, but except by O. simplex all species were collected from deep waters (up to 3 610 m depth). Finally, Caso (1986) recorded the presence of Ophiophragmus marginatus, and Salcedo-Martínez et al. (1988) cited Amphipholis squamata to the SMP.

During the current century, the unique work with new additions of Ophiuroidea to the SMP was published by Honey-Escandón et al. (2008) who reported 10 species (Asteronyx longifissus, Ophiomyxa panamensis, Ophiotoma paucispina, Ophiernus adspersus adspersus, Amphioplus coniortodes, Amphiodia platyspina, Amphiodia tabogae, Ophiocnida hispida, Ophiothela mirabilis and Ophionereis perplexa). It is important to note that both O. adspersus adspersus and A. coniortodes are species distributed in the Caribbean and the Gulf of México (Lyman, 1883; H.L.Clark, 1918; Pawson et al., 2009; Miloslavich et al., 2010), therefore the record of the species in the SMP should be treated with caution. Finally, Ophiacantha paucispina originally described by Lütken and Mortensen (1899) was transferred to the genus Ophiotoma (Clark, 1915; Martynov, 2010). The ophiuroid Amphioplus (Unioplus) daleus was reported in the Gulf of Tehuantepec at 590 km south of Puerto Ángel, Oaxaca (Turner & Hallan, 2011; Granja-Fernández & López-Pérez, 2012), however, since the original locality of recollection was off Mexican waters (Station 3414; 10 °14´ N - 96° 28´ W) (Lütken & Mortensen, 1899), we excluded the record from the current checklist.

Almost all the consulted museums had records of Ophiuroidea from the SMP, except CAS, MACN, AMNH and YPM. The museums with the highest number of species from the study area were LACM (24) and ICML-UNAM (23) (Table 1). Museum review render two new reports of Ophiuroidea, Microphiopholis platydisca from Acapulco, Guerrero at 20.1 m depth (LACM 38-136) and Chacahua Bay, Oaxaca from 73.1 to 91.44 m depth (MCZ 5666), and Ophiostigma tenue from Petatlán Bay, Guerrero at 45.7 m depth (LACM 34.138), and Chacahua Bay, Oaxaca from 73.1 to 91.44 m depth (MCZ 5668). The brittle star M. platydisca was originally described by Nielsen (1932) as Amphipholis platydisca, posteriorly the species was transferred to the genus Microphiopholis (Turner, 1985).

 

Echinoidea: The Echinoidea is the class with the highest number of published works (64) from the SMP, most of them of Agassiz (1863, 1872, 1873, 1881, 1898, 1904), Caso (1946, 1948a, 1949, 1978a, 1978b, 1979, 1980a, 1980b, 1983) and Clark (1914, 1917, 1925, 1940, 1948). During the 19th century a total of 11 works were produced, while in the 20th and 21st century 36, and 17 were published, respectively (Fig. 1C).

During the 19th century, 10 species of echinoids were found for the first time in the SMP, all of them by Alexander Agassiz. Agassiz (1863), described the species Diadema mexicanum, Echinometra vanbrunti, Toxopneustes roseus, Rhyncholampas pacificus and Clypeaster rotundus, from Acapulco and recorded for the first time Meoma ventricosa grandis in the area. Agassiz (1872) also added Mellita longifissa for Acapulco. Finally, Agassiz (1898) described Aporocidaris milleri, Cystechinus loveni and Cystocrepis setigera, all of them from Acapulco at a depth of 3 436 m.

The 20th century had the highest addition of records of Echinoidea in the SMP. Agassiz (1904) did his last contribution adding to the list Brisaster townsendi from Acapulco, a species previously described from the Gulf of Panamá and 500 miles south Guaymas, Sonora. Clark (1914) described Clypeaster ochrus based in a specimen collected at Acapulco by the Hassler expedition in 1872. Grant and Hertlein (1938) in their important work on the West American Cenozoic Echinoidea reported for the first time the species Astropyga pulvinata, Clypeaster europacificus, Encope micropora and Encope perspectiva from Guerrero. Clark (1940) also recorded for the first time the species Eucidaris thouarsii in Zihuatanejo, Guerrero, and in 1948 the same author recorded Mesocentrotus franciscanus, Strongylocentrotus purpuratus, Encope micropora fragilis and Encope wetmorei from Guerrero. It is paramount to emphasize that Clark (1948: 279) commented about M. franciscanus: “There is in the present collection a very good adult specimen from Petatlán Bay, but it seems quite improbable and needs confirmation”, and about Strongylocentrotus purpuratus (Clark, 1948: 281): “The Velero has secured specimens at 46 stations, of which the southernmost at Petatlán Bay, México. As the only other stations south of the United States at which purpuratus was secured are at or near Cedros Island, the Petatlán Bay record needs confirmation. The specimens labeled Petatlán are undoubtedly purpuratus but, as in the case of franciscanus, the locality is hard to credit”. Years later, Caso (1978b) reported M. franciscanus and S. purpuratus from White Friars and Petatlán (Guerrero). In the Mexican Pacific, both species are recorded in the northernmost area of the country (Baja California and Baja California Sur) and in the Gulf of California (Solís-Marín et al., 2005; Honey-Escandón et al., 2008), moreover there is no record of the species elsewhere in the Eastern Pacific (Alvarado et al. 2010); therefore this record should be treated with caution. Although the name Strongylocentrotus franciscanus has been recently used for authors as Honey-Escandón et al. (2008), this species was replaced to the genus Mesocentrotus (Tatarenko & Poltaraus, 1993). Otherwise, Caso (1978a, 1980a, 1980b, 1983) recorded for the first time in the area the species Hesperocidaris asteriscus, Centrostephanus coronatus, Mellita notabilis, and Brissus latecarinatus. Harold and Telford (1990) reported Mellita grantii and Mellita kanakoffi from Acapulco. Mellita kanakoffi is an echinoid with both recent and fossil records (Upper Pleistocene), but in the studied area only living specimens have been added (Harold & Telford, 1990). Finally, Gamboa-Contreras and Tapia-García (1998) reported the species Clypeaster elongatus from the Gulf of Tehuantepec. The other known report of this species is in the Eastern Pacific by Clark (1948) from the Galápagos Islands.

During the current century, Benítez-Villalobos et al. (2008a) and Honey-Escandón et al. (2008) reported Tripneustes depressus, Lovenia cordiformis and Agassizia scrobiculata. Further, Zamorano and Leyte-Morales (2009) conducted a study of echinoderms associated to coral reefs from Zihuatanejo and Acapulco and recorded for the first time the species Echinometra mathaei oblonga in the SMP. In the Eastern Pacific (México, Panamá and Galápagos Islands), Echinometra mathaei oblonga was named as Echinometra oblonga (Lessios, 2005; Solís-Marín et al., 2005; Honey-Escandón et al., 2008; Alvarado et al., 2010) but since 1943 Mortensen indicated that the correct name for this species was the former.

The echinoids from the SMP are deposited in all the museums, except the FMNH. Most of the species are deposited in the ICML-UNAM (17 spp.) and MCZ (16 spp.). During museum revision, two new records were added to the fauna of the SMP, Arbacia stellata from Acapulco (MCZ 471), and Encope laevis from Bahía de Petatlán, Guerrero at depths of 9.1 to 18.3 m (CAS-IZ 87348.00). Along the Eastern Pacific (México, El Salvador, Nicaragua, Costa Rica, Panamá, Colombia, Perú), A. stellata has currently named as Arbacia incisa (Hooker et al., 2005; Neira & Cantera, 2005; Solís-Marín et al., 2005; Honey-Escandón et al., 2008; Alvarado et al. 2010), but according to Mortensen (1935) the valid name is A. stellata. Meanwhile, E. laevis is an echinoid distributed in Nicaragua, Costa Rica and Colombia (Neira & Cantera, 2005; Alvarado et al., 2010), therefore this is the first record of this species in the Mexican Pacific.

 

Holothuroidea: To date, the current number of references for the class Holothuroidea in the SMP is 35. During the 19th century there were only four works, during the 20th century 22 papers were published and only nine works during the current century (Fig. 1D). Caso (1954, 1958, 1962, 1963, 1964, 1965, 1967a, 1967b, 1986) and Deichmann (1936, 1937, 1938, 1941, 1958) were the authors with most of the published works on Holothuroidea in the study area.

The 19th century addressed the lowest number of Holothuroidea reports in the SMP. Verrill (1867b) was the first to record Holothuroidea in the area; he reported Afrocucumis ovulum, Holothuria (Halodeima) kefersteini and Holothuria (Selenkothuria) lubrica from Acapulco. Twenty-seven years later, Ludwig (1894) added Pseudostichopus mollis, Pannychia moseleyi, Oneirophanta mutabilis mutabilis, Benthodytes incerta, Benthodytes sanguinolenta and Molpadia musculus from Guerrero.

The highest addition of records occurred during the 20th century. Deichmann (1937) reported Isostichopus fuscus from Acapulco. One year later, Deichmann (1938) made an important contribution on the holothurians from Lower California, Central America and the Galápagos islands. In this work, Deichmann reported for the first time in the study area Lissothuria ornata, Neothyone gibber, Euthyonidiella zacae, Pentamera chierchiae, Pentamera zacae, Pseudocnus californicus, Holothuria (Semperothuria) languens, Holothuria (Thymiosycia) arenicola and Holothuria (Thymiosycia) impatiens. Euthyonidiella zacae and P. zacae were originally described from Tangola-Tangola Bay, Oaxaca, México. In 1941, Deichmann reported Neothyone gibbosa, Pachythyone pseudolugubris, Trachythyone peruana, Thyone strangeri and Leptopentacta panamica from the studied area, but the type localities are located in the Gulf of California and Colombia (Deichmann, 1941). On the other hand, T. strangeri is the unique species with its type locality in the SMP (Isla Grande, Guerrero at 12.8 m depth). It is important to highlight that Deichmann (1941) made a mistake when recorded the latitude of the type locality as “170° 46´ N”, instead of “17° 46´ N”. Caso (1958) reported Holothuria (Halodeima) inornata from Zihuatanejo and Deichmann (1958) reported three more species of the genus Holothuria (Holothuria (Mertensiothuria) hilla, Holothuria (Selenkothuria) portovallartensis and Holothuria (Semperothuria) imitans) from Guerrero and Oaxaca. Caso (1965) carried out the most important work on Holothuroidea from Zihuatanejo reporting a total of six new records (Holothuria (Cystipus) inhabilis, Holothuria (Cystipus) rigida, Holothuria (Halodeima) atra, Holothuria (Mertensiothuria) leucospilota, Holothuria (Platyperona) difficilis and Labidodemas maccullochi) for the SMP. Luke (1982) reported for the first time in the area Abyssocucumis abyssorum and Ypsilothuria bitentaculata, which are distributed in deep waters (more than 3000 m depth) (Solís-Marín et al., 2009; Massin & Hendrickx, 2011). Salcedo-Martínez et al. (1988) recorded Thyone parafusus in Zihuatanejo, Guerrero, which correspond to the only record of the species for the entire Eastern Pacific, being Salcedo-Martínez et al. (1988) and Honey-Escandón et al. (2008) the only authors addresing this species. Herrero-Pérezrul and Reyes-Bonilla (1997) added Euapta godeffroyi from Zihuatanejo. Finally, Solís-Marín and Laguarda-Figueras (1999) described Cucumaria flamma after material collected from Zihuatanejo.

In contrast to the other classes of echinoderms, the Holothuroidea have the highest number of records (12) during the 21st century. Despite the type localities for the species Molpadiodemas neovillosus are Caroline and Galápagos Islands, O’Loughlin and Ahearn (2005) reviewed material from Punta Maldonado, Guerrero (3 436 m). In the same year, Zamorano and Leyte-Morales (2005a) added Holothuria (Selenkothuria) theeli and Holothuria (Stauropora) fuscocinerea to the SMP. The work of Honey-Escandón et al. (2008) may be the most important work since it added 6 holothuroid records (Psolidium dorsipes, Pachythyone lugubris, Neopentamera anexigua, Holothuria (Theelothuria) paraprinceps, Labidodemas americanum and Chiridota aponocrita) for the study area. Massin and Hendrickx (2011) recorded the deep-water species Psolidium gracile (740-790 m depth) and Laetmogone scotoeides (1 180 - 1 420 m depth) from Guerrero.

The Holothuroidea of the SMP are deposited in the collections of CAS, ICML-UNAM, LACM, MCZ, UMAR and USNM. The collection with the highest number of deposited species is the ICML-UNAM, with 32 species. The revision of the material in the ICML-UNAM added two new records for the Southern Mexican Pacific: Thyone bidentata from Playa del Hotel Caleta, Guerrero (ICML-UNAM 5.66.1), and Chiridota rigida from Playa Contramar, Guerrero (ICML-UNAM 5.74.11), Playa Las Gatas, Guerrero (ICML-UNAM 5.74.12), Estacahuite, Oaxaca (ICML-UNAM 5.74.14), Playa Panteón, Oaxaca (ICML-UNAM 5.74.15) and La Mina, Oaxaca (ICML-UNAM 5.74.16).

 

Diversity and distribution: The checklist of echinoderms from the Southern Mexican Pacific includes 162 species, 96 genera, 54 families, 20 orders and four classes (Table 1). The Holothuroidea were the most numerous with 52 species followed by the Ophiuroidea (46 species), Echinoidea (34) and Asteroidea (30) (Table 2). No records in the literature or in museum collections refer to the Crinoidea from the SMP. The orders with the highest number of species were Ophiurida (Ophiuroidea, 45 species), Aspidochirotida (Holothuroidea, 22), Dendrochirotida (Holothuroidea, 22), Paxillosida (Asteroidea, 13) and Clypeasteroida (Echinoidea, 13); while the orders Velatida and Brisingida (Asteroidea), Euryalida (Ophiuroidea), Arbacioida and Cassiduloida (Echinoidea), and Molpadida (Holothuroidea) were represented by a single species. The best represent families were Holothuriidae (Holothuroidea, 19 species) and Amphiuridae (Ophiuroidea, 18 species). Both families are considered the most diverse and dominant in shallow tropical waters (O´Loughlin et al., 2007; Stöhr et al., 2012).

For the entire Mexican Pacific, Solís-Marín et al. (2013) reported 216 species, the Ophiuroidea being the most numerous with 63 species followed by the Asteroidea (59), Holothuroidea (53), Echinoidea (37) and Crinoidea (4) (Table 3). Similar trend is observed for the Gulf of California (232 species, Solís-Marín et al., 2013) and the Pacific side of Central America (282 species, Alvarado et al., 2010) where the Ophiuroidea and Holothuroidea are the most diverse (Table 3). It is important to note the following between studies: 1) the databases structures are different; while the present study and Alvarado et al. (2010) contain reports from published resources and museums, Solís-Marín et al. (2013) includes only museum records; 2) the studied areas are highly contrasting regarding coastal extension; while the present study includes approximately 1 200 km along the coast of the States of Guerrero, Oaxaca and Chiapas, the entire Mexican Pacific has 7 800 km, the Gulf of California, 3 000 km and Central America, 3 800 km. The results suggest that in just 40 % of the coastal line, the SMP comprises 70 % of the total echinoderm fauna compared with, for example, the Gulf of California. This suggests that the echinoderm biodiversity from the SMP is not small considering that the Gulf of California is extensively and far better studied.

According to our results, Guerrero was the State with the highest number of recorded species (135 spp.), followed by Oaxaca (94 spp.), Chiapas (15 spp.) and the Gulf of Tehuantepec (14 spp.). Specifically, the Holothuroidea were the most diverse in Guerrero (48 spp.) and Oaxaca (33 spp.), while the Echinoidea (7 spp.) and the Asteroidea (8 spp.) were the most diverse in Chiapas and the Gulf of Tehuantepec, respectively (Fig. 2).

Holothuroidea have not been recorded in the Gulf of Tehuantepec, but there are records of Trachythyone peruana and Holothuria (Halodeima) kefersteini from Chiapas (Fig. 2). Of the 30 species of Asteroidea, 14 spp. were only reported in one State, most of them from Guerrero; while, Astropecten armatus and Heliaster microbrachius were widespread in the entire region. Twenty-seven species of Ophiuroidea were reported in one State, mostly for Guerrero, while Diopederma daniana was the only species distributed in the entire SMP. Fifteen species of Echinoidea were reported in one State (mostly in Guerrero), but Astropyga pulvinata, Echinometra vanbrunti, Clypeaster europacificus and Encope micropora were widespread in the entire region. Finally, the Holothuroidea were distributed mostly in one State (22 spp.).

Respect to the most complete and recent works of echinoderms from the Mexican Pacific (Honey-Escandón et al. 2008) and Oaxaca (Bastida-Zavala et al., 2013), this work increases the number of known species in 35% for Guerrero, 62 % for Oaxaca, and 7 % for Chiapas. Compared with previous works, we found a substantial increase in regional diversity after the compilation of information from 131 references and the review of several specimens deposited in 11 museums and reference collections.

The historic revision of SMP echinoderms has allowed us to identify valid names, synonyms, misspelled names and erroneous distributions. The checklist provided in this study is an important baseline for future studies in western México, but also for the entire Eastern Tropical Pacific, given that most of the included species are widespread in the area and inhabit multiple substrates and habitats within the region.

Altough the SMP is considered a poorly studied region within the Mexican Pacific (Bastida-Zavala et al., 2013), echinoderm studies have lasted over a century (1860-2013) and have resulted from intensive collaborative effort of national and overseas researches in all disciplines but mainly in systematics. This contribution reveals that the SMP harbors a large number of echinoderm species (chiefly Ophiuroidea and Holothuroidea) being comparable to larger regions such as the Mexican Pacific and the Gulf of California (Table 3). Recent studies addressing the biodiversity of echinoderms in the area indicate, for example, that the States of Oaxaca and Chiapas could harbor two to three times the observed biodiversity (Granja-Fernández & López-Pérez, 2012; López-Pérez et al., 2012a).

Although the historical revision provides valuable information for systematics and ultimately for species management, it also provides an insight of the completeness and gaps regarding the knowledge of the echinoderm fauna in the SMP. In this regard, we encourage a full field survey for the entire region at different depths and across different habitats, especially in soft and rocky bottoms from Chiapas since ongoing in this area suggests its potential to render new records and species yet to be discovered. Currently, the States of Oaxaca and Chiapas have experienced an increase in anthropogenic pressure, mostly in the zones of Zihuatanejo and Acapulco (Guerrero), and Puerto Escondido and Bahías de Huatulco (Oaxaca) (CONANP, 2003; López-Pérez et al., 2012b), therefore these areas are required to be monitored.

ACKNOWLEDGMENTS

The authors are very grateful to S. Stöhr (Swedish Museum of Natural History), A. Kroh (Natural History Museum Vienna) and C. Mah (USNM) for assistance in systematics matters. Special thanks to A. Baldinger and P. Benson (MCZ), G. Hendler (LACM), P. Greenhall (USNM), C. LeBeau (AMNH), E. Rodríguez (AMNH), S. Keable (AMS), R. Mooi (CAS), A. Tablado (MACN) and A. Martínez-Melo (ICML-UNAM) for sharing museum records. We thank M. Hendrickx, R. Bastida-Zavala, A. Durán-González, P. Medina-Rosas, A. Hernández-Morales and C. Aparicio-Cid for providing valuable literature. This project was founded by CONABIO (HJ029, JF047). This work is part of the PhD thesis of RGF at UAM. While this work was done, RGF was supported by a scholarship from CONACYT (336853). We thank the reviewers for commenting and improving the manuscript.

RESUMEN

Listado de los equinodermos (Echinodermata) del Pacífico sur de México: una revisón histórica. Los equinodermos del Pacífico Sur mexicano han sido estudiados por tres siglos, sin embargo discrepancias en la nomenclatura de algunas especies han prevalecido a través del tiempo. El objetivo de este trabajo es presentar la primera lista de especies actualizada de todos los nombres válidos y sinonimias, así como una revisión histórica del estudio de los equinodermos en el Pacífico Sur mexicano. La lista de especies se basa en una exhaustiva búsqueda de registros de literatura y de especímenes depositados en colecciones de referencia. Existen 162 especies válidas de equinodermos en el Pacífico Sur mexicano incluidas en 96 géneros, 54 familias y 20 órdenes. El estado de Guerrero presentó 135 especies, Oaxaca, 94 y Chiapas, 15. Se actualizó la lista de especies y se añadieron cinco nuevos registros de especies (Microphiopholis platydisca, Ophiostigma tenue, Arbacia stellata, Thyone bidentata, Chiridota rigida) para el Pacífico Sur mexicano y uno para el Pacífico mexicano (Encope laevis). Esta lista de especies incrementa el número de especies para el área de estudio, sin embargo, los estudios sugieren que aún existen más especies por ser descubiertas.

 

Palabras claves: Asteroidea, Ophiuroidea, Echinoidea, Holothuroidea, México.

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TABLE 1

Checklist of valid names (in bold) and synonyms of the echinoderms from the Southern Mexican Pacific. Distribution in the Southern Mexican Pacific and museum records in brackets. Numbers following brackets refer to literature where the taxa were listed, see references section. For acronyms see material and methods section.

All taxa are listed as originally reported in the literature

 

CUADRO 1

Lista de nombres válidos (en negritas) y sinonimias de los equinodermos del Pacífico Sur Mexicano. La distribución dentro del Pacífico Sur Mexicano y los registros de museos se presentan en corchetes. Los números después de los corchetes se refieren a la literatura en donde se reportó a los taxa, ver la sección de referencias. Para los acrónimos

ver la sección de material y métodos. Todos los taxa se enlistan tal cual fueron reportados en la literatura

 

Phylum Echinodermata Brugiére, 1791

Class Asteroidea Blainville, 1830

Order Paxillosida Perrier, 1884

Family Luidiidae Sladen, 1889

Genus Luidia Forbes, 1839

Luidia bellonae Lütken, 1865 [GRO, OAX; LACM] 8, 116

Luidia (Alternaster) bellonae Lütken, 1865 13, 87

Luidia columbia (Gray, 1840) [GRO, OAX, GT; CAS, LACM, MACN, MCZ] 8

Luidia brevispina Lütken, 1871 75

Luidia columbiae (Gray, 1840) 75

Luidia (Petalaster) columbia (Gray, 1840) 13, 87

Luidia foliolata (Grube, 1866) [GT] 75

Luidia latiradiata (Gray, 1871) [OAX, CHIS, GT; MACN, MCZ] 8

Astropecten latiradiatus (Gray, 1871) 103, 104

Luidia (Platasterias) latiradiata (Gray, 1871) 13, 44, 87, 114, 116

Platasterias latiradiata Gray, 1871 19, 20, 23, 34, 66, 71, 82, 112, 123

Luidia superba A.H. Clark, 1917 [OAX] 8

Luidia (Alternaster) superba A.H. Clark, 1917 13, 87

Luidia tessellata Lütken, 1859 [GRO, OAX, CHIS; CAS] 8

Luidia (Petalaster) tessellata Lütken, 1859 13, 87, 114

Family Astropectinidae Gray, 1840

Genus Astropecten Gray, 1840

Astropecten armatus Gray, 1840 [GRO, OAX, CHIS, GT; CAS, ICML-UNAM, LACM, MACN, AMNH, USNM] 8, 13, 75, 87, 114

Astropecten ornatissimus Fisher, 1906 [GRO; CAS, ICML-UNAM] 87, 116

Astropecten regalis Gray, 1840 [GRO, CHIS, GT; ICML-UNAM, LACM] 75, 87, 114

Astropecten verrilli de Loriol, 1899 [GT]

Astropecten californicus Fisher, 1906 75

Family Porcellanasteridae Sladen, 1883

Genus Eremicaster Fisher, 1905

Eremicaster crassus (Sladen, 1883) [GRO, OAX; USNM] 8

Eremicaster crassus gracilis (Sladen, 1883) 13

Porcellanaster waltharii Ludwig, 1905 93

Eremicaster pacificus (Ludwig, 1905) [GRO; USNM] 44

Porcellanaster pacificus Ludwig, 1905 93

Family Pseudarchasteridae Sladen, 1889

Genus Pseudarchaster Sladen, 1889

Pseudarchaster pulcher Ludwig, 1905 [GRO] 73, 93

 

Order Valvatida Perrier, 1884

Family Asterinidae Gray, 1840

Genus Meridiastra O’Loughlin, 2002

Meridiastra modesta (Verrill, 1870) [GRO; ICML-UNAM] 87

Asterina agustincasoi Caso, 1977 35, 45, 101, 116

Family Goniasteridae Forbes, 1841

Genus Ceramaster Verrill, 1899

Ceramaster leptoceramus (Fisher, 1905) [GT] 84, 94

Genus Mediaster Stimpson, 1857

Mediaster transfuga Ludwig, 1905 [GRO; MCZ, USNM] 73, 84, 93

Family Oreasteridae Fisher, 1911

Genus Nidorellia Gray, 1840

Nidorellia armata (Gray, 1840) [GRO, OAX; ICML-UNAM, UMAR] 8, 13, 67, 87, 103, 104, 110, 116

Genus Pentaceraster Döderlein, 1916

Pentaceraster cumingi (Gray, 1840) [GRO, OAX; ICML-UNAM, LACM, UMAR] 8, 13, 67, 87, 129

Oreaster occidentalis Verrill, 1870 56, 94, 110

Family Asteropseidae Hotchkiss & A.M. Clark, 1976

Genus Asteropsis Müller & Troschel, 1840

Asteropsis carinifera (Lamarck, 1816) [OAX; ICML-UNAM, UMAR] 8

Asteropsis spinosa (Gray, 1840) 13, 87

Family Mithrodiidae Viguier, 1878

Genus Mithrodia Gray, 1840

Mithrodia bradleyi Verrill, 1870 [GRO, OAX; ICML-UNAM, UMAR] 8, 13, 87, 110

Family Ophidiasteridae Verrill, 1870

Genus Linckia Nardo, 1834

Linckia columbiae Gray, 1840 [GRO] 116

Genus Pharia Gray, 1840

Pharia pyramidatus (Gray, 1840) [GRO, OAX; AMS, ICML-UNAM, LACM, MCZ, AMNH, UMAR] 8, 10

Pharia pyramidata (Gray, 1840) 38, 56, 94, 127, 129

Pharia pyramidatus pyramidatus (Gray, 1840) 13, 87

Ophidiaster pyramidatus Gray, 1840 103

Genus Phataria Gray, 1840

Phataria unifascialis (Gray, 1840) [GRO, OAX; AMS, CAS, ICML-UNAM, LACM, MCZ, AMNH] 8, 13, 38, 56, 87, 94, 110, 112, 121, 127, 129

Linckia unifascialis Gray, 1840 103, 104, 124, 125

 

Order Velatida Perrier, 1884

Family Pterasteridae Perrier, 1875

Genus Hymenaster Wyville-Thomson, 1873

Hymenaster violaceus Ludwig, 1905 [GRO; USNM] 46, 53, 73, 93, 116

 

Order Forcipulatida Perrier, 1884

Family Heliasteridae Viguier, 1878

Genus Heliaster Gray, 1840

Heliaster helianthus (Lamarck, 1816) [GRO; ICML-UNAM] 116

Heliaster kubiniji Xantus, 1860 [GRO; AMS, ICML-UNAM, LACM, MCZ, AMNH] 38, 49, 87, 103, 104, 116

Heliaster microbrachius Xantus, 1860 [GRO, OAX, CHIS, GT; AMS, FMNH, ICML-UNAM, LACM, MCZ, USNM] 8, 13, 38, 49, 87, 114, 116

Heliaster microbrachia Xantus, 1860 97

Heliaster polybrachius H.L. Clark, 1907 [GRO] 47

Family Zoroasteridae Sladen, 1889

Genus Zoroaster Wyville-Thomson, 1873

Zoroaster hirsutus Ludwig, 1905 [GRO; CAS, USNM] 47, 53, 74, 93, 116

 

Order Brisingida Fisher, 1928

Family Freyellidae Downey, 1986

Genus Freyella Perrier, 1885

Freyella insignis Ludwig, 1905 [GRO] 93

 

Class Ophiuroidea Gray, 1840

Order Euryalida Lamarck, 1816

Family Asteronychidae Verrill, 1899

Genus Asteronyx Müller & Troschel, 1842

Asteronyx longifissus Döderlein, 1927 [GRO; ICML-UNAM] 87

 

Order Ophiurida Müller & Troschel, 1840

Family Ophiomyxidae Ljungman, 1867

Genus Ophiomyxa Müller & Troschel, 1840

Ophiomyxa panamensis Lütken & Mortensen, 1899 [GRO; ICML-UNAM] 87

Family Ophiacanthidae Ljungman, 1867

Genus Ophiacantha Müller & Troschel, 1842

Ophiacantha moniliformis Lütken & Mortensen, 1899 [GRO; USNM] 69, 87, 95

Genus Ophiotoma Lyman, 1883

Ophiotoma paucispina (Lütken & Mortensen, 1899) [OAX; ICML-UNAM] Ophiacantha paucispina Lütken & Mortensen, 1899 8, 87

Family Ophiuridae Müller & Troschel, 1840

Genus Ophiura Lamarck, 1801

Ophiura (Ophiuroglypha) irrorata irrorata (Lyman, 1878) [OAX; LACM] Ophiura irrorata (Lyman, 1878) 77, 94

Genus Ophiomisidium Koehler, 1914

Ophiomisidium leurum Ziesenhenne, 1940 [OAX; LACM] 69, 77, 131

Genus Amphiophiura Matsumoto, 1915

Amphiophiura superba (Lütken & Mortensen, 1899) [GRO; USNM]

Ophioglypha superba Lütken & Mortensen, 1899 95

Genus Ophiernus Lyman, 1878

Ophiernus adspersus adspersus Lyman, 1883 [GRO] 87

Ophiernus adspersus annectens Lütken & Mortensen, 1899 [GRO; USNM] Ophiernus polyporus Lütken & Mortensen, 1899 95

Ophiernus polyporum Lütken & Mortensen, 1899 69

Ophiernus seminudus Lütken & Mortensen, 1899 [GT; AMS] 51, 77

Genus Stegophiura Matsumoto, 1915

Stegophiura ponderosa (Lyman, 1878) [OAX] 77

Amphiophiura ponderosa (Lyman, 1878) 94

Family Amphiuridae Ljungman, 1867

Genus Amphioplus Verrill, 1899

Amphioplus coniortodes H.L. Clark, 1918 [GRO; USNM] 87

Genus Amphiura Forbes, 1843

Amphiura arcystata H.L. Clark, 1911 [GRO; MCZ] 56

Amphiura seminuda Lütken & Mortensen, 1899 [OAX, GT; LACM] 75, 77, 94

Amphiura serpentina Lütken & Mortensen, 1899 [GRO; USNM] 69, 95

Genus Amphichondrius Nielsen, 1932

Amphichondrius granulatus (Lütken & Mortensen, 1899) [GRO, OAX; MCZ] 77

Amphichondrius granulosus Nielsen, 1932 56

Amphichondrius laevis Ziesenhenne, 1940 [OAX; LACM] 69, 77, 131

Genus Amphiodia Verrill, 1899

Amphiodia (Amphispina) urtica (Lyman, 1860) [OAX; MCZ] 77

Amphiodia urtica (Lyman, 1860) 56

Amphiodia platyspina Nielsen, 1932 [GRO; ICML-UNAM] 87

Amphiodia sculptilis Ziesenhenne, 1940 [GRO, OAX; AMS, LACM, MCZ] 69, 77, 108, 131

Amphiodia tabogae Nielsen, 1932 [GRO; ICML-UNAM, LACM] 87

Genus Amphipholis Ljungman, 1866

Amphipholis squamata (Delle Chiaje, 1828) [GRO, OAX; LACM] 77, 110

Genus Dougaloplus Clark A.M., 1970

Dougaloplus gastracanthus (Lütken & Mortensen, 1899) [GRO; USNM]

Amphioplus gastracanthus (Lütken & Mortensen, 1899) 51

Amphiura gastracantha Lütken & Mortensen, 1899 69, 87, 95

Genus Microphiopholis Turner, 1985

Microphiopholis platydisca (Nielsen, 1932) [GRO, OAX; LACM, MCZ]

Genus Ophiocnida Lyman, 1865

Ophiocnida hispida (Le Conte, 1851) [GRO, OAX; ICML-UNAM, UMAR] 77, 80, 87

Genus Ophiophragmus Lyman, 1865

Ophiophragmus marginatus (Lütken, 1856) [OAX; ICML-UNAM, LACM] 8, 42, 77, 87

Ophiophragmus papillatus Ziesenhenne, 1940 [GRO, OAX; LACM] 69, 77, 80, 131

Genus Ophiostigma Lütken, 1856

Ophiostigma tenue Lütken, 1856 [GRO, OAX; LACM, MCZ]

Family Ophiotrichidae Ljungman, 1867

Genus Ophiothrix Müller & Troschel, 1840

Ophiothrix (Ophiothrix) rudis Lyman, 1874 [GRO, OAX; ICML-UNAM, LACM, UMAR] 8, 76, 77, 80

Ophiothrix rudis Lyman, 1874 9, 25, 87, 94, 110, 127

Ophiothrix (Ophiothrix) spiculata Le Conte, 1851 [GRO, OAX; ICML-UNAM, LACM, MCZ, USNM] 8, 76, 77, 80

Ophiothrix spiculata Le Conte, 1851 25, 38, 42, 51, 87, 110, 127

Genus Ophiothela Verrill, 1867

Ophiothela mirabilis Verrill, 1867 [GRO, OAX; ICML-UNAM, UMAR] 8, 76, 77, 80, 87

Family Ophiactidae Matsumoto, 1915

Genus Ophiactis Lütken, 1856

Ophiactis savignyi (Müller & Troschel, 1842) [GRO, OAX; ICML-UNAM, LACM, UMAR, USNM)] 8, 9, 17, 25, 38, 56, 76, 77, 80, 87, 94, 95, 110, 127

Ophiactis simplex (Le Conte, 1851) [GRO, OAX; ICML-UNAM, LACM, UMAR] 8, 9, 76, 77, 80, 87, 94, 110, 127

Family Ophionereididae Ljungman, 1867

Genus Ophionereis Lütken, 1859

Ophionereis annulata (Le Conte, 1851) [GRO, OAX; FMNH, ICML-UNAM, LACM, MCZ, UMAR] 8, 51, 76, 77, 79, 80, 87, 110, 121, 127

Ophionereis dictyota Ziesenhenne, 1940 25

Ophionereis perplexa Ziesenhenne, 1940 [GRO; ICML-UNAM] 87

Family Ophiocomidae Ljungman, 1867

Genus Ophiocoma L. Agassiz, 1835

Ophiocoma aethiops Lütken, 1859 [GRO, OAX; FMNH, ICML-UNAM, LACM, UMAR] 8, 11, 25, 38, 51, 56, 76, 77, 80, 87, 94, 110, 121, 124, 125, 127

Ophiocoma alexandri Lyman, 1860 [GRO, OAX; ICML-UNAM, LACM, MCZ, UMAR] 8, 11, 15, 25, 38, 51, 56, 69, 76, 77, 80, 87, 94, 96, 110, 124, 125, 127, 130

Family Ophiodermatidae Ljungman, 1867

Genus Ophioderma Müller & Troschel, 1840

Ophioderma panamensis Lütken, 1859 [GRO, OAX; ICML-UNAM, LACM, UMAR] 76, 77, 80

Ophiocryptus granulosus Nielsen, 1932 8, 9, 25, 77, 87

Ophioderma panamense Lütken, 1859 21, 47, 52, 81

Ophiura panamensis (Lütken, 1859) 124, 125

Ophioderma teres (Lyman, 1860) [GRO, OAX; ICML-UNAM, LACM, UMAR] 8, 25, 51, 76, 77, 80, 87

Ophiura teres Lyman, 1860 125

Ophioderma variegata Lütken, 1856 [GRO, OAX; ICML-UNAM, LACM] 8, 77

Ophioderma variegatum Lütken, 1856 25, 56, 87

Genus Diopederma H.L. Clark, 1913

Diopederma daniana (Verrill, 1867) [GRO, OAX, CHIS, GT; ICML-UNAM, LACM, UMAR, USNM] 8, 77

Diopederma danianum (Verrill, 1867) 25, 56, 75, 87, 94, 114

Family Ophiolepididae Ljungman, 1867

Genus Ophiolepis Müller & Troschel, 1840

Ophiolepis fulva H.L. Clark, 1940 [OAX; LACM, MCZ] 56, 69, 77

Ophiolepis pacifica Lütken, 1856 [GRO, OAX; ICML-UNAM, UMAR] 8, 76, 77, 80, 87

Ophiozona pacifica (Lütken, 1856) 25

Ophiolepis variegata Lütken, 1856 [GRO, OAX, GT; ICML-UNAM, LACM, USNM] 8, 25, 38, 42, 56, 75, 77, 87, 94

Genus Ophiomusium Lyman, 1869

Ophiomusium glabrum Lütken & Mortensen, 1899 [OAX; ICML-UNAM] 8, 77, 87, 94

Ophiomusium variabile Lütken & Mortensen, 1899 [GRO; MCZ, USNM] 51, 69, 87, 95

 

Class Echinoidea Leske, 1778

Order Cidaroida Claus, 1880

Family Ctenocidaridae Mortensen, 1928

Genus Aporocidaris A. Agassiz & H.L. Clark, 1907

Aporocidaris milleri (A. Agassiz, 1898) [GRO; ICML-UNAM, USNM] 36, 87, 100, 116

Porocidaris milleri A. Agassiz, 1898 5, 6

Family Cidaridae Gray, 1825

Genus Eucidaris Pomel, 1883

Eucidaris thouarsii (L. Agassiz & Desor, 1846) [GRO, OAX; ICML-UNAM, LACM, MCZ, USNM] 8, 13, 18, 22, 36, 56, 87, 94, 116, 127, 128

Genus Hesperocidaris Mortensen, 1928

Hesperocidaris asteriscus H.L.Clark, 1948 [GRO, OAX; ICML-UNAM, UMAR] 8, 13, 36, 110, 116, 127, 129

 

Order Diadematoida Duncan, 1889

Family Diadematidae Gray, 1855

Genus Astropyga Gray, 1825

Astropyga pulvinata (Lamarck, 1816) [GRO, OAX, CHIS; CAS, ICML-UNAM, LACM, MCZ] 8, 13, 22, 36, 38, 56, 57, 81, 87, 114, 116

Genus Centrostephanus Peters, 1855

Centrostephanus coronatus (Verrill, 1867) [GRO; ICML-UNAM] 36, 87, 116, 129

Genus Diadema Gray, 1825

Diadema mexicanum A. Agassiz, 1863 [GRO, OAX; ICML-UNAM, LACM, MCZ, UMAR, USNM, YPM] 1, 2, 3, 4, 6, 7, 8, 9, 13, 14, 18, 36, 55, 58, 59, 60, 68, 85, 87, 89, 90, 91, 106, 109, 110, 116, 124, 125, 126, 127, 128, 129

Centrechinus mexicanus (A. Agassiz, 1863) 23, 57, 81, 130

 

Order Arbacioida Gregory, 1900

Family Arbaciidae Gray, 1855

Genus Arbacia Gray, 1835

Arbacia stellata (Blainville, 1825; ?Gmelin, 1788) [GRO; MCZ]

 

Order Camarodonta Jackson, 1912

Family Echinometridae Gray, 1855

Genus Echinometra Gray, 1825

Echinometra mathaei oblonga (Blainville, 1825) [GRO]

Echinometra oblonga (Blainville, 1825) 129

Echinometra vanbrunti A. Agassiz, 1863 [GRO, OAX, CHIS; CAS, ICML-UNAM, LACM, MACN, MCZ, UMAR, USNM] 1, 8, 13, 18, 56, 57, 68, 87, 94, 114, 116, 121, 127, 128, 129

Echinometra van brunti A. Agassiz, 1863 2, 22, 37, 38, 106, 110, 124, 125, 126

Heliocidaris stenopora H.L. Clark, 1912 22, 81

Toxocidaris mexicana (L. Agassiz & Desor, 1846) 1, 2

Family Toxopneustidae Troschel, 1872

Genus Toxopneustes L. Agassiz, 1841

Toxopneustes roseus (A. Agassiz, 1863) [GRO, OAX; ICML-UNAM, LACM, MCZ, UMAR, USNM] 8, 13, 18, 24, 37, 38, 55, 57, 81, 87, 94, 109, 110, 116, 127, 128, 129

Boletia rosea A. Agassiz, 1863 1, 2

Lytechinus roseus (A. Agassiz, 1863) 124, 125

Genus Tripneustes L. Agassiz, 1841

Tripneustes depressus A. Agassiz, 1863 [OAX; UMAR] 8, 13

Family Strongylocentrotidae Gregory, 1900

Genus Mesocentrotus Tatarenko & Poltaraus, 1993

Mesocentrotus franciscanus (A. Agassiz, 1863) [GRO; LACM]

Strongylocentrotus franciscanus (A. Agassiz, 1863) 37, 57

Genus Strongylocentrotus Brandt, 1835

Strongylocentrotus purpuratus (Stimpson, 1857) [GRO] 37, 57

 

Order Cassiduloida Claus, 1880

Family Cassidulidae L. Agassiz & Desor, 1847

Genus Rhyncholampas A. Agassiz, 1869

Rhyncholampas pacificus (A. Agassiz, 1863) [GRO, OAX; AMNH, AMS, CAS, ICML-UNAM, MCZ, USNM] 8, 87, 126

Cassidulus pacificus (A. Agassiz, 1863) 13, 24, 41, 57, 81, 109, 116

Pygorhyncus pacificus A. Agassiz, 1863 1, 68, 124, 125

Rhyncholampas pacifica (A. Agassiz, 1863) 55, 122

Rhynchopygus pacificus (A. Agassiz, 1863) 2, 105, 106

 

Order Clypeasteroida A. Agassiz, 1872

Family Clypeasteridae L. Agassiz, 1835

Genus Clypeaster Lamarck, 1801

Clypeaster elongatus H.L. Clark, 1948 [GT]

Clypeaster elongata H.L. Clark, 1948 75

Clypeaster europacificus H.L. Clark, 1914 [GRO, OAX, CHIS; CAS, ICML-UNAM, LACM, MCZ] 40, 81, 87, 114, 116

Clypeaster ochrus H.L. Clark, 1914 [GRO, OAX; MCZ] 40, 50, 56, 57, 68, 81, 116

Clypeaster rotundus (A. Agassiz, 1863) [GRO, CHIS; MCZ] 2, 40, 57, 81, 94, 116, 119

Stoloniclypeus rotundus A. Agassiz, 1863 124, 125

Stolonoclypus rotundus A. Agassiz, 1863 1, 2, 68

Family Mellitidae Stefanini, 1912

Genus Mellita L. Agassiz, 1841

Mellita grantii Mortensen, 1948 [GRO; MCZ] 83

Mellita kanakoffi Durham, 1961 [GRO; MCZ] 83

Mellita longifissa Michelin, 1858 [GRO, CHIS; CAS, ICML-UNAM, MCZ] 2, 21, 40, 70, 81, 83, 87, 106, 107, 114, 116, 117

Mellita notabilis H.L. Clark, 1947 [GRO, CHIS; CAS, ICML-UNAM, USNM] 83, 87, 114, 117

Mellita eduardobarrosoi Caso, 1981 39, 40, 88, 116

Genus Encope L. Agassiz, 1840

Encope laevis H.L. Clark, 1948 [GRO; CAS]

Encope micropora L. Agassiz, 1841 [GRO, OAX, CHIS; ICML-UNAM, MCZ] 8, 13, 40, 81, 87, 114, 116

Encope micropora fragilis H.L. Clark, 1948 [GRO, GT; CAS, LACM, MCZ] 40

Encope fragilis H.L.Clark, 1948 57, 68, 71, 75, 116

Encope perspectiva L. Agassiz, 1841 [GRO, OAX; CAS, LACM] 40, 57, 81

Encope wetmorei A.H. Clark, 1946 [GRO; ICML-UNAM, LACM, USNM] 38, 40, 57, 87, 116

 

Order Holasteroida Durham & Melville, 1957

Family Urechinidae Duncan, 1889

Genus Cystechinus A. Agassiz, 1879

Cystechinus loveni A. Agassiz, 1898 [GRO, OAX; MCZ, USNM] 5, 6, 68

Urechinus loveni (A. Agassiz, 1898) 41, 52, 99, 116

Family Pourtalesiidae A. Agassiz, 1881

Genus Cystocrepis Mortensen, 1907

Cystocrepis setigera (A. Agassiz, 1898) [GRO; MCZ, USNM]

Echinocrepis setigera A. Agassiz, 1898 5, 6, 68

 

Order Spatangoida Claus, 1876

Family Schizasteridae Lambert, 1905

Genus Brisaster Gray, 1855

Brisaster townsendi (A. Agassiz, 1898) [GRO]

Schizaster townsendi A.Agassiz, 1898 6

Family Prenasteridae Lambert, 1905

Genus Agassizia Valenciennes, 1846

Agassizia scrobiculata Valenciennes, 1846 [GRO, OAX; ICML-UNAM] 8, 13, 87

Family Brissidae Gray, 1855

Genus Brissus Gray, 1825

Brissus latecarinatus (Leske, 1778) [GRO; ICML-UNAM] 41, 87, 116

Genus Meoma Gray, 1851

Meoma ventricosa grandis Gray, 1851 [GRO, OAX; CAS, ICML-UNAM, LACM, MCZ, UMAR] 8, 13, 43, 87, 115, 116

Kleinia nigra A. Agassiz, 1863 1, 2

Meoma grandis Gray, 1851 2, 3, 4, 6, 24, 41, 55, 56, 81, 109, 119, 122, 126

Meoma nigra (A. Agassiz, 1863) 124, 125

Family Loveniidae Lambert, 1905

Genus Lovenia Desor, in Agassiz & Desor, 1847

Lovenia cordiformis A. Agassiz, 1872 [OAX; UMAR] 8, 13

 

Class Holothuroidea Selenka, 1867

Order Dendrochirotida Grube, 1840

Family Psolidae Burmeister, 1837

Genus Lissothuria Verrill, 1867

Lissothuria ornata Verrill, 1867 [GRO, OAX; CAS, ICML-UNAM, LACM] 8

Thyonepsolus beebei Deichmann, 1937 63, 64, 87, 110

Genus Psolidium Ludwig, 1887

Psolidium dorsipes Ludwig, 1887 [GRO, OAX; ICML-UNAM] 87

Psolidium gracile Ludwig, 1894 [GRO] 98

Family Sclerodactylidae Panning, 1949

Genus Afrocucumis Deichmann, 1944

Afrocucumis ovulum (Selenka, 1867) [GRO, OAX; ICML-UNAM, LACM, MCZ] 8, 87, 118

Euthyonidium ovulum (Selenka, 1867) 33, 63, 64, 110

Stolus ovulum Selenka, 1867 111, 125

Thyone ovulum (Selenka, 1867) 61, 63, 120

Genus Neothyone Deichmann, 1941

Neothyone gibber (Selenka, 1867) [GRO, OAX; ICML-UNAM, MCZ] 64, 87, 110, 118

Thyone gibber (Selenka, 1867) 63

Neothyone gibbosa Deichmann, 1941 [GRO, OAX; ICML-UNAM, MCZ, UMAR] 64, 87, 110

Genus Pachythyone Deichmann, 1941

Pachythyone lugubris (Deichmann, 1939) [GRO; LACM, USNM] 87

Pachythyone pseudolugubris Deichmann, 1941 [GRO, OAX; ICML-UNAM, LACM] 64, 87

Genus Neopentamera Deichmann, 1941

Neopentamera anexigua Deichmann, 1941 [GRO; ICML-UNAM] 87, 118

Genus Euthyonidiella Heding & Paning, 1954

Euthyonidiella zacae (Deichmann, 1938) [OAX; MCZ] 8

Phyllophorus (Urodemella) zacae Deichmann, 1938 118

Phyllophorus zacae Deichmann, 1938 63, 64

Family Phyllophoridae Östergren, 1907

Genus Pentamera Ayres, 1852

Pentamera chierchiae (Ludwig, 1887) [GRO, OAX; ICML-UNAM, MCZ] 110

Pentamera chierchia (Ludwig, 1887) 63, 87

Pentamera zacae Deichmann, 1938 [OAX; MCZ] 8, 63, 64

Genus Thyone Jaeger, 1833

Thyone bidentata Deichmann, 1941 [GRO; ICML-UNAM]

Thyone parafusus Deichmann, 1941 [GRO; ICML-UNAM] 87, 110

Thyone strangeri Deichmann, 1941 [GRO; MCZ]

Family Cucumariidae Ludwig, 1894

Genus Abyssocucumis Heding, 1942

Abyssocucumis abyssorum (Théel, 1886) [GRO, OAX; ICML-UNAM, MCZ, USNM] 94

Staurocucumis abyssorum (Théel, 1886) 8, 87

Genus Cucumaria Blainville, 1830

Cucumaria flamma Solís-Marín & Laguarda-Figueras, 1999 [GRO; ICML-UNAM, USNM] 87, 113, 118

Genus Pseudocnus Panning, 1949

Pseudocnus californicus (Semper, 1868) [GRO, OAX; ICML-UNAM, LACM, MCZ, UMAR, USNM] 8, 87

Cucumaria californica Semper, 1868 27, 31, 63, 64, 110

Genus Trachythyone Studer, 1876

Trachythyone peruana (Semper, 1868) [GRO, CHIS; ICML-UNAM, LACM] 87, 114

Pentacta peruana (Semper, 1868) 42, 64

Genus Leptopentacta H.L. Clark, 1938

Leptopentacta panamica Deichmann, 1941 [OAX; LACM] 64

Family Ypsilothuriidae Heding, 1942

Genus Ypsilothuria Perrier, 1886

Ypsilothuria bitentaculata (Ludwig, 1893) [GRO, OAX; MCZ] 98

Sphaerothuria bitentaculata Ludwig, 1893 94

 

Order Aspidochirotida Grube, 1840

Family Holothuriidae Burmeister, 1837

Genus Holothuria Linnaeus, 1767

Holothuria (Cystipus) inhabilis Selenka, 1867 [GRO]

Holothuria parinhabilis Cherbonnier, 1951 31

Jaegerothuria inhabilis (Selenka, 1867) 110

Holothuria (Cystipus) rigida (Selenka, 1867) [GRO; ICML-UNAM] 87

Holothuria rigida (Selenka, 1867) 31

Holothuria (Halodeima) atra Jaeger, 1833 [GRO, OAX; ICML-UNAM] 87, 97

Holothuria atra Jaeger, 1833 31

Holothuria (Halodeima) inornata Semper, 1868 [GRO]

Holothuria inornata Semper, 1868 27, 31

Holothuria (Halodeima) kefersteini (Selenka, 1867) [GRO, OAX, CHIS; ICML-UNAM, LACM, MCZ, UMAR, USNM] 87, 114, 118

Holothuria kefersteinii (Selenka, 1867) 16, 54, 119

Holothuria (Paraholothuria) riojai Caso, 1963 29, 31

Ludwigothuria kefersteinii (Selenka, 1867) 28, 65, 110

Stichopus kefersteinii Selenka, 1867 111, 120, 125

Holothuria (Mertensiothuria) hilla Lesson, 1830 [GRO, OAX; CAS, ICML-UNAM, LACM, UMAR, USNM]

Brandtothuria gyrifer (Selenka, 1867) 65

Holothuria gyrifer (Selenka, 1867) 31

Holothuria (Thymiosycia) hilla Lesson, 1830 87

Microthele (Paramicrothele) zihuatanensis Caso, 1964 30, 31

Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) [GRO; ICML-UNAM] 87, 97

Holothuria leucospilota (Brandt, 1835) 31

Holothuria (Platyperona) difficilis Semper, 1868 [GRO, OAX; ICML-UNAM, UMAR] 87, 97

Holothuria difficilis Semper, 1868 31

Microthele difficilis (Semper, 1868) 110

Holothuria (Selenkothuria) lubrica Selenka, 1867 [GRO, OAX; ICML-UNAM, LACM, MCZ, UMAR] 8, 87, 118

Cucumaria lubrica (Selenka, 1867) 110

Holothuria lubrica Selenka, 1867 26, 27, 31, 62, 111, 120, 125

Holothuria pseudolubrica Cherbonnier, 1951 27, 31

Selenkothuria lubrica (Selenka, 1867) 65, 110, 127

Holothuria (Selenkothuria) portovallartensis Caso, 1954 [GRO, OAX; ICML-UNAM, UMAR] 8, 87

Holothuria portovallartensis Caso, 1954 127

Selenkothuria portovallartensis (Caso, 1954) 65

Holothuria (Selenkothuria) theeli (Deichmann, 1938) [GRO, OAX; ICML-UNAM, UMAR] 8, 87

Holothuria theeli (Deichmann, 1938) 127

Holothuria (Semperothuria) imitans Ludwig, 1875 [GRO, OAX; ICML-UNAM, UMAR] 8, 87, 97

Holothuria imitans Ludwig, 1875 127

Holothuria imitans var. polymorpha Caso, 1962 31

Semperothuria imitans (Ludwig, 1875) 65

Holothuria (Semperothuria) languens Selenka, 1867 [OAX]

Holothuria languens Selenka, 1867 63

Holothuria (Stauropora) fuscocinerea Jaeger, 1833 [GRO, OAX; ICML-UNAM, UMAR] 8

Holothuria fuscocinerea Jaeger, 1833 12, 127

Holothuria (Mertensiothuria) fuscocinerea Jaeger, 1833 87

Holothuria (Theelothuria) paraprinceps Deichmann, 1937 [GRO; ICML-UNAM] 87

Holothuria (Thymiosycia) arenicola Semper, 1868 [GRO, OAX; ICML-UNAM, UMAR] 8, 87

Holothuria arenicola Semper, 1868 63, 127

Holothuria (Thymiosycia) impatiens (Forskål, 1775) [GRO, OAX; CAS, ICML-UNAM, LACM, UMAR, USNM] 8, 9, 87, 97

Brandtothuria impatiens (Forskål, 1775) 65, 110

Holothuria impatiens (Forskål, 1775) 27, 31, 63, 127

Genus Labidodemas Selenka, 1867

Labidodemas americanum Deichmann, 1938 [GRO, OAX; ICML-UNAM] 87

Labidodemas maccullochi (Deichmann, 1958) [GRO, OAX; ICML-UNAM] 87, 118

Irenothuria maccullochi Deichmann, 1958 31

Family Stichopodidae Haeckel, 1896

Genus Isostichopus Deichmann, 1958

Isostichopus fuscus (Ludwig, 1875) [GRO, OAX; ICML-UNAM, MCZ, UMAR] 8, 87, 110, 127, 129

Stichopus fuscus Ludwig, 1875 27, 31, 32, 62

Family Synallactidae Ludwig, 1894

Genus Molpadiodemas Heding, 1935

Molpadiodemas neovillosus O’Loughlin & Ahearn, 2005 [GRO; USNM] 102

Genus Pseudostichopus Théel, 1886

Pseudostichopus mollis Théel, 1886 [GRO] 92

 

Order Elasipodida Théel, 1882

Family Laetmogonidae Ekman, 1926

Genus Pannychia Théel, 1882

Pannychia moseleyi Théel, 1882 [GRO; USNM] 98

Laetmophasma fecundum Ludwig, 1893 92

Genus Laetmogone Théel, 1879

Laetmogone scotoeides (H.L. Clark, 1913) [GRO] 98

Family Deimatidae Théel, 1882

Genus Oneirophanta Théel, 1879

Oneirophanta mutabilis mutabilis Théel, 1879 [GRO, OAX]

Oneirophanta mutabilis Théel, 1879 92, 94

Family Psychropotidae Théel, 1882

Genus Benthodytes Théel, 1882

Benthodytes incerta Ludwig, 1893 [GRO; USNM] 92

Benthodytes sanguinolenta Théel, 1882 [GRO] 92

 

Order Molpadida Haeckel, 1896

Family Molpadiidae Müller, 1850

Genus Molpadia Cuvier, 1817

Molpadia musculus Risso, 1826 [GRO, OAX] 48, 94, 98

Ankyroderma danielsseni Théel, 1886 92

 

Order Apodida Brandt, 1835

Family Chiridotidae Öestergren, 1898

Genus Chiridota Eschscholtz, 1829

Chiridota aponocrita H.L. Clark, 1920 [GRO, OAX; ICML-UNAM, UMAR] 87

Chiridota rigida Semper, 1868 [GRO, OAX; ICML-UNAM]

Family Synaptidae Burmeister, 1837

Genus Euapta Östergren, 1898

Euapta godeffroyi (Semper, 1868) [OAX, GRO] 78, 86