Eunicidae Classification Essay

On By In 1

Marine biodiversity of Costa Rica: Class Polychaeta (Annelida)

Harlan K. Dean

Department of Invertebrates. Museum of Comparative Zoology. 26 Oxford St. Harvard University. Cambridge, MA 02138 (USA).


Received 22-IV-2003. Corrected 28-VII-2004. Accepted 13-VIII-2004.

Abstract

Three hundred and thirty two species of polychaetes belonging to 50 families are reported for Costa Rica based on previously published reports. In a few families, available material was re-examined and appropriate changes in species identifications were made. Only three species, (Glycera oxycephala Ehlers, 1887; Hemipodia pustulata (Friedrich 1956); and Scolepis (Scolepis) squamata (Müller, 1806)) have been reported from the Caribbean coast of Costa Rica, and these three species have also been reported from the Pacific Coast. The families with more than 15 species are the Spionidae (26), Cirratulidae (24), Nereididae (21), Lumbrineridae (19) and Paraonidae (16). Despite extensive sampling on the Pacific coast over the last 25 years, the Costa Rican polychaete fauna, especially that of the Caribbean coast, remains poorly known.

Key words: Annelida, Polychaeta, biodiversity, polychaete worms, Gulf of Nicoya, Golfo Dulce, Costa Rica.


The first published reports of polychaetes collected in Costa Rica were described in a series of reports by Grube (1856, 1857, 1859) on species collected mainly from Puntarenas. Only a few scattered reports were published in subsequent years (Mörch 1863, Augener 1922) until Treadwell 1928a) published a list of species collected from Cocos Island (Treadwell 1928a) and a new species of Phyllodoce from the Gulf of Nicoya (Treadwell 1928b). This was followed by a series of papers by Hartman (1939a, b, 1940, 1944a, b) reporting on the Allan Hancock cruises in the eastern Pacific with many subtidal polychaete records from Costa Rica. Fauchald (1973) and Dexter (1974) later identified intertidal polychaetes, mostly from sandy beaches on both the Caribbean and Pacific coasts of Costa Rica. Nichols Driscoll (1976) provided the first records of the subtidal polychaete fauna from Golfo Dulce. In a survey of eastern Pacific Spirorbidae, Knight-Jones et al. (1979) included several species collected at various sites along the Pacific coast of Costa Rica. In the early 1980’s the cooperative efforts of the University of Delaware and the Universidad de Costa Rica led to the publication of several long lists of mainly subtidal polychaetes from the Gulf of Nicoya (Golfo de Nicoya) estuary (Vargas et al. 1985, Maurer et al. 1988, and Dean 1996a). Vargas (1987, 1988, 1989, 1996) also published several lists of polychatetes as part of his analysis of the infaunal community of the intertidal sandflats of Punta Morales. Dean (1996b) and Leon-Morales and Vargas (1998) expanded on the earlier work of Nichols Driscoll (1976) in Golfo Dulce with a list of polychaetes collected by the R/V Victor Hensen cruises in 1993 and 1994. More recent published works of selected polychaete taxa collected in Costa Rica include Dean (1998a, b, 2001a, b) and Böggemann (2002).

This list of published records of the polychaete fauna of Costa Rica is an attempt to both summarize what is known of the polychaete fauna of Costa Rica and to correct improper or misleading identifications. Many of the original species designations have been modified by modern workers and every attempt has been made to include all synonomies and nomenclatural changes in the list. Also, some of the recent material collected in the Gulf of Nicoya and Golfo Dulce was re-examined and necessary corrections were made in the original species designations for some families.

It is noted in the remarks section for many of these species that the original species designation may be in error and the material, if available, should be re-examined. This is most often due to the reported species occurrence for Costa Rica being well outside the published range for that species. In other cases, a more intensive analysis by modern workers may have revealed that what was originally considered a single species with a wide geographic range is actually two or more recognizable similar species. Finally, some species reported from Costa Rican waters by early workers have been poorly described, the type material has been lost, and they have not been collected since their original description The validity of these species is, therefore, doubtful.

The present list of species includes 332 species of polychaetes reported from Costa Rica belonging to 50 of the approximately 80 recognized families. It must be noted, however, that this number may be somewhat high since many species have only been identified to the genus level (or sometimes just to family). Many of those specimens are probably occurrences of more fully identified species already included in the list. Any overestimates due to the inclusion of these incompletely identified species are probably compensated for by the incomplete sampling efforts. For example, there have only been 3 species of polychaetes (the glycerids Glycera oxycephala and Hemipodia pustulata and the spionid Scolepis (Scolepis) squamata) reported from the Caribbean coast of Costa Rica (all three have also been reported from Pacific Costa Rica as well). If more intensive sampling, especially on the Caribbean coast, was conducted it would be even more apparent that the polychaete fauna of Costa Rica is very high.

The families with the greatest number of species recorded from Costa Rica are the Spionidae, Cirratulidae, Nereididae, and Lumbrineridae. These families are usually well represented in the benthos often with a high abundance of species but, again, there may be a samping effect involved here too. The higher number of species reported in the families Cirratulidae, Nereididae, Capitellidae and Pilargiidae may be at least partially due to those families having been sampled more intensively from intertidal sites and/or having undergone more rigorous taxonomic analysis (Dean 1998a, 2001a, b.). If material from some other families were to be re-examined, perhaps many more species would be recognized.

Despite the shortcomings in the data it is apparent from this list that the polychaete fauna of Costa Rica is highly diverse and, in all like-lihood, is more diverse on the Pacific coast.

Hopefully, this review will facilitate future work by acting as a much needed reference to what is presently known about this fauna. The polychaetes usually make up approximately 50% of the species present in most marine benthic communities and are considered an important food source for many other organisms. Also, as pointed out by Blake (1997a), their often rapid generation times and high reproductive potential allow the polychaetes to respond very quickly to changes in their environment. This has resulted in the utilization of opportunistic polychaete species such as Mediomastus, Polydora, Streblospio, etc., as so-called "indicator" species in the monitoring of ecological disturbances such as pollution (Reish 1979, Blake 1997a). Any analysis of the health of the benthos requires knowledge of the polychaete fauna and it is hoped that this list will facilitate the ecological assessment of marine environments well as stimulate future research in Costa Rica.


Acoetidae
Acoetes mortenseni (Monro 1928)

Costa Rican records: Golfo de Nicoya, Vargas et al. 1985: 337, Maurer et al. 1988: 48 (both as Panthalis marginata), Dean 1996a: 75 (as Panthalis mortenseni)

Habitat: Subtidal, muddy sand, 18 m.

Distribution: Known from California (USA) to Pacific Panama (Pettibone 1989).

Panthalis marginata Hartman 1939a = Acoetes mortenseni 

Panthalis mortenseni (Monro 1928) = Acoetes mortenseni 

Polyodontes oculeus (Treadwell 1901)

Costa Rican records: Golfo de Nicoya, Maurer et al. 1988: 48, Dean 1996a: 75 (both as Polyodontes oculea).

Habitat: Subtidal, muddy sand to sand, 13- 18 m.

Distribution: Reported in the Caribbean Sea and from the Gulf of California (Mexico) to Panama and Ecuador in the Eastern Pacific (Pettibone 1989).

Acrocirridae
Acrocirrus sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 290 (listed in the Cirratulidae).

Habitat: Subtidal, 64 m.

Ampharetidae

Amphicteis scaphobranchiata Moore 1906

Costa Rican records: Golfo Dulce – Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal, mud (?).

Distribution: Reported from Washington State (USA) to Baja California (Mexico) in the eastern Pacific as well as the Gulf of Mexico (Hilbig 2000a).

Amphicteis sp.

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 334, Maurer et al. 1988: 46.

Habitat: Subtidal, mud.

 Amphicteis obscurior Chamberlin 1919 = Amphicteis sp.

Isolda bipinnata (Fauchald 1977) = Mellina sp.

Mellina sp.

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 334, Maurer et al. 1988: 46, Dean 1996a: 71 (all as Isolda bipinnata Fauchald).

Habitat: Subtidal, sandy mud, 15-22 m.

Amphinomidae
Chloeia entypa Chamberlin 1919

Costa Rican record: Chatham Bay, Cocos Island - Hartman 1939b: 8.

Habitat: Subtidal, mostly sand.

Distribution: C. entypa has been recorded from California to Ecuador in the eastern Pacific (Hartman 1939b), however, Kudenov (1995) has indicated that C. pinnata Moore 1911 may be the senior synonym of this species.

Chloeia euglochis Ehlers 1887 = Chloeia viridis

Chloeia pseudeuglochis Augener 1922

Costa Rican record: Culebra Bay, Golfo de Papagayo - Augener 1922: 39.

Habitat: Intertidal to shallow subtidal

Remarks: Augener made no drawings and differentiated this species from C. euglochis only by the presence of distally entire neurosetae. The neurosetae of C. pinnata have small teeth proximal to the small prong of the forked setae but the tines themselves are smooth (Kudenov 1995). It is probable Augener’s specimen was C. pinnata.

Chloeia viridis Schmarda 1861

Costa Rican records: Cocos Island –Treadwell 1928a: 450, off Nuez Island (Isla Manuelita), Hartman 1940: 205. Golfo de Santa Elena, Salinas Bay - Hartman 1940: 205.

Habitat: Subtidal, 11-90 m, rock, coral & corallines

Distribution: C. viridis has been reported from the western Atlantic and Caribbean Sea as well as the eastern Pacific from the Gulf of California to Panama (Fauchald 1977, Gathof 1984).

Eurythoe complanata (Pallas 1776)

Costa Rican records: Cocos Island, Puerto Culebra - Hartman 1940: 202. Golfo de Santa Elena, Playa Blanca & Parker Bay – Hartman 1940: 202, Treadwell 1941: 18.

Habitat: Intertidal to shallow subtidal.

Distribution: Considered circumtropical (Salazar-Vallejo 1996-1997).

Eurythoe sp.

Costa Rican record: Golfo de Nicoya -Maurer et al. 1984: 51

Habitat: Collected by shrimp trawl.

Hermodice carunculata (Pallas 1766)

Costa Rican record: Golfo de Santa Elena, Port Parker - Treadwell 1941: 17.

Habitat: Intertidal to shallow subtidal

Distribution: H. carunculata is characterized as a warm-water Atlantic species (Salazar-Vallejo 1996-1997) and Treadwell’s report from the eastern Pacific is doubtful.

Linopherus annulata (Hartmann-Schröder 1965) = Linopherus spiralis

Linopherus kristiani Salazar-Vallejo 1987

Costa Rican record: Golfo Dulce – Dean 1996b: 83.

Habitat: Subtidal mud.

Distribution: Previously described from western Mexico by Salazar-Vallejo (1987)

Linopherus spiralis (Wesenberg-Lund 1949)

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 334, Maurer et al. 1988: 46 (both as L. annulata), Vargas 1989: 1909.

Habitat: Intertidal sandy mud.

Distribution: This species was described by Wesenberg-Lund (1949) from the Iranian Gulf, and Hartmann-Schröder (1992) later tentatively identified material from the central Atlantic. This report from the Pacific needs to be verified.

Notopygos crinita Grube 1855

Costa Rican record: Cocos Island - Treadwell 1928a: 450

Habitat: Subtidal, 183 m.

Distribution: Salazar-Vallejo (1996-1997) characterized this species as trans-Atlantic and Treadwell’s identification of material from Cocos Island is dubious.

Notopygos ornata Grube 1856

Costa Rican records: Puntarenas, Golfo de Nicoya - Grube 1856: 53. Golfo de Santa Elena, Parker Bay - Hartman 1940: 207, Treadwell 1941: 18.

Habitat: intertidal to shallow subtidal? Corals.

Distribution: Fauchald (1977) reports this species to occur in warm waters in the western Atlantic and the eastern Pacific. Hartman (1940) reported its occurrence from the Gulf of California (Mexico) to the Galapagos in the eastern Pacific.


Aphroditidae
Aphrodita sp.

Costa Rican record: SE of Judas Point (Punta Judas) - Treadwell 1941: 21.

Habitat: Subtidal, mud, 90 m.

Aphrodita japonica Marenzeller 1879

Costa Rican record: Cabo Blanco, Nicoyan Penninsula - Dean 1996a: 71.

Habitat: Subtidal.

Distribution: Fauchald (1977) reported this species from the North Pacific and Eastern Pacific and Hartman (1939a) reported its distribution in the eastern Pacific Ocean from California (USA) to Ecuador


Capitellidae
Capitella capitata (Fabricius 1780) species complex

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 334, Maurer et al. 1988: 46, Dean 1996b: 83 (as Paraheteromastides sp.), Dean 2001a: 71-73.

Habitat: subtidal mud, 40-43 m, intertidal muddy sand and amongst mangrove roots.

Distribution: Cosmopolitan with equatorial submergence (Warren 1991) but known to occur as a complex of sibling species (Grassle and Grassle 1976).

Dasybranchus lumbricoides Grube 1878

Costa Rican record: Golfo de Nicoya –Dean 2001a: 73, Dittman & Vargas 2001: 283.

Habitat: Intertidal muddy sand

Distribution: Reported from the eastern and western Pacific as well as the western Atlantic and Gulf of Mexico. It has been collected in the eastern Pacific from California to Panama as well as the Galapagos Islands in intertidal or shallow waters (Dean 2001a).

Decamastus nudus Thomassin 1970

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335, Maurer et al. 1988: 46, Dean (1996a): 71 (all as N. tenuis Moore), Dean 2001a: 73-75.

Habitat: Subtidal muds to sands, 11-42 m, intertidal muddy sand.

Distribution: Originally described from Madagascar (Thomassin 1970), but since reported from the Gulf of California, Mexico (Hernández-Alcántara and Solís-Weiss 1998) and the Pacific Coast of Costa Rica (Dean 2001a) in the eastern Pacific.

 Heteromastus filiformis (Claparéde 1864)

Costa Rican record: Golfo de Nicoya –Dean 2001a: 75.

Habitat: Intertidal, muddy sand and sand.

Distribution: Reported from numerous sites in the Atlantic and Pacific Ocean as well as the Mediterranean Ocean and Australia (Blake 2000a).

Leiochrides cf. pallidior (Chamberlin 1918)

Costa Rican record: Golfo de Nicoya –Dean 2001a: 75-76.

Habitat: Subtidal muddy sand, 18 m.

Distribution: Reported from intertidal and shallow subtidal waters in California and the Pacific coast of Costa Rica (Dean 2001a).


Mastobranchus variabilis
Ewing 1984

Costa Rican record: Golfo de Nicoya –Dean 2001a: 76.

Habitat: Subtidal muddy sand, 18 m.

Distribution: Originally described from the Gulf of Mexico (Ewing 1984b) and since reported from the Gulf of California, Mexico (Hernández-Alcántara and Solís-Weiss 1998) and the Golfo de Nicoya, Costa Rica (Dean 2001a).


Mediomastus ambiseta
(Hartman 1947)

Costa Rican records: Golfo de Nicoya –Dean 1996a: 71 (as M. californiensis Hartman (in part)), Dean 2001a: 76-78. Golfo Dulce –Dean 1996b: 83.

Habitat: Subtidal mud to sand, 11-50 m, Intertidal muddy sand and amongst mangrove roots.

Distribution: Known from the East and West coasts of North America as well as the Gulf of Mexico (Warren et al. 1994). M. ambiseta is known from California to Golfo Dulce in Costa Rica (Dean 2001a).

Mediomastus californiensis Hartman 1944

Costa Rican records: Golfo de Nicoya –Maurer & Vargas 1984: 101, Vargas et al. 1985: 335, Vargas 1987: 304, 1988: 294, 1989: 1909, Maurer et al. 1987: 138, Maurer et al. 1988: 46, Dean 1996a: 71, Vargas 1996: 359, Dittman & Vargas 2001: 283, 287, Dean 2001a: 78-80. Golfo Dulce - Dean 1996b: 83, Leòn-Morales & Vargas 1998: 85.

Habitat: Subtidal mud to sand, 9-200 m intertidal muddy sand and sand.

Distribution: M. californiensis has been reported from both coasts of North America and the Gulf of Mexico (Ewing 1984a), numerous sites from the Pacific coast of Mexico (Salazar-Vallejo 1981, de León-González 1994a, Hernández-Alcántara and Solís-Weiss 1998), and the Golfo de Nicoya and Golfo Dulce in Costa Rica (Dean 2001a).

Notomastus hemipodus Hartman 1947

Costa Rican records: —Golfo de Nicoya –Maurer & Vargas 1984: 101 (as N. luridis Verrill), Maurer et al. 1987: 138 (as N. luridis Verrill), Vargas 1987: 304, Maurer et al. 1988: 46 (as N. luridis Verrill), Dean 1996a: 71 (as N. luridis Verrill), Dean 2001a: 80.

Habitat: Subtidal mud to sand, 9-46 m, intertidal muddy sand and amongst mangrove roots.

Distribution: Both coasts of North America and the Gulf of Mexico (Ewing 1984a), the Gulf of California, Mexico (Hernández-Alcántara and Solís-Weiss 1998) and Golfo de Nicoya, Costa Rica (Dean 2001a).

Notomastus lineatus Claparéde 1870

Costa Rican record: Golfo de Nicoya, Punta Morales - Dean 2001: 81-82.

Habitat: intertidal fine sand

Distribution: Cosmopolitan, numerous reports from Pacific Mexico (Salazar-Vallejo 1985, Hernández-Alcántara and Solís-Weiss 1998) and the Golfo de Nicoya, Costa Rica (Dean 2001a).

Notomastus tenuis Moore 1909

Costa Rican record: Golfo de Nicoya –Dean 2001a: 83.

Habitat: Subtidal muddy sand to sand, 14- 24 m.

Distribution: Known in the Eastern Pacific from Canada to Mexico with an additional "questionable" record from the Gulf of Mexico (Dean 2001a).

Notomastus tenuis (sensu Maurer et al. 1988) = Decamastus nudis

Mediomastus sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 290.

Habitat: Subtidal, sand, 64 m.

Chaetopteridae

Mesochaetopterus alipes Monro 1928 = Spiochaetopterus costarum

Mesochaetopterus sp. =Spiochaetopterus costarum

Phyllochaetopterus sp. = Spiochaetopterus costarum

Phyllochaetopterus socialis (Claparéde 1870) = Spiochaetopterus costarum

Spiochaetopterus costarum (Claparéde 1870)

Material examined: Sta. 28, 9º52’16" N, 84º45’30" W, 26 m. Golfo Dulce, intertidal. Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335, Maurer et al. 1988: 46 (both as Mesochaetopterus sp. & Phyllochaetopterus sp. ), Maurer & Vargas 1984: 101 ( as Phyllochaetopterus sp.), Dean 1996a: 71 (as Mesochaetopterus alipes and Phyllochaetopterus socialis).

Habitat: Subtidal, mud to sand, 11-35 m, Intertidal, mud.

Distribution: This species is widespread and has been reported from Canada to California in the eastern Pacific (Blake 2000b).


Chrysopetalidae
Chrysopetalum macrophthalmum Hartmann-Schröder 1959

Costa Rican record: Golfo de Nicoya, Puntarenas - Hartmann-Schröder 1959: 94-96.

Habitat: Intertidal, sand.

Distribution: Reported by Hartmann-Schröder from El Salvador and Costa Rica in the eastern Pacific (Hartmann-Schröder 1959).

Paleaquor nicoyensis Watson-Russel 1986

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335 (as Paleanotus heterosetae), Maurer et al. 1988: 46 ( as Paleonotus sp.), Watson Russel 1986: 170-172, Dean 1996a: 71.

Habitat: Subtidal, mud to sand, 11-44 m.

Distribution: Known only from the Golfo de Nicoya (Watson-Russel 1986).

Paleanotus heterosetae Hartman = Paleaquor nicoyensis

Paleonotus sp. = Paleaquor nicoyensis

Cirratulidae
Aphelochaeta longisetosa (Hartmann-Schröder 1965) = A. glandaria = Cirratulus sp.

Aphelochaeta glandaria Blake 1996

Material examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (50), Jan 1981 (2), Apr 1981 (2); Jun 1981 (1), Aug 1981 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jul 1980 (19); Oct 1980 (26); Jan 1981 (460); Apr 1981 (19); Jun 1981 (32); Apr 1982 (49). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Oct 1980 (2); Jan 1981 (4); Apr 1981 (5); Aug (1).

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335, Maurer et al. 1988: 46 (as Tharyx multifilis), Dean 1996a: 71 (as A. longisetosa). Golfo Dulce - Dean 1996b: 83 (as A. longisetosa), León-Morales & Vargas 1998: 85 (as A. longisetosa (in part).

Habitat: Subtidal muddy sand to sand, 11-18 m.

Distribution: Previously known only from central and southern California, USA (Blake 1996a).

Aphelochaeta marioni (Saint Joseph, 1894)

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291.

Habitat: Subtidal, muddy sand, 64 m.

Distribution: This species has been reported from numerous regions, however, this species may actually be a complex of morphologically similar species (see Blake 1996 for a review of this issue).

Aphelochaeta monilaris (Hartman 1960)

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291 (as Tharyx monilaris).

Habitat: Subtidal, muddy sand, 64 m.

Distribution: Widespread in the eastern Pacific (Blake 1996a)

Aphelochaeta multifilis 1909

Habitat: Subtidal, muddy sand and sandy mud, 64- 70 m.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291 (as Tharyx multifilis).

Distribution: Presently known only from California, USA (Blake 1996a).

Caulleriella alata (Southern 1914) = C. magnaoculata

Caulleriella bioculata (Keferstein 1862 ) = C. magnaoculata

Caulleriella magnaoculata Hartmann-Schröder 1962

Material examined: Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Jun 1981 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jul 1980, (2); Oct 1980 (25); Jan 1981 (22); Apr 1981 (20); June 1981 (15); Apr 1982, (2). Intertidal: Punta Morales, 10ºN, 84º58’ W, south side, intertidal sandy mud, Jun 1986 (2), Aug 1996 (1)

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335 (as Caulleriella bioculata & Caulleriella sp.), Maurer et al. 1988: 46 (as Caulleriella alata and Caulleriella bioculata), Dean 1996a: 71 (as Caulleriella alata).

Habitat: Subtidal mud and sand, 11-26 m, intertidal, sandy mud.

Distribution: C. magnaoculata is known from Peru (Blake 1996a).

Chaetozone corona Berkeley & Berkeley 1941

Costa Rican records: Golfo Dulce – Vargas et al. 1985: 335, Maurer et al. 1988: 46, Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal muds.

Distribution: Previously reported from southern California and western Mexico (Blake 1996a).

Chaetozone hedgepethi Blake 1996

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Aug 1981 (1).

Costa Rican records: Golfo de Nicoya –Dean 1996a: 72 (as Chaetozone sp. B).

Habitat: Subtidal, sand, 11 m.

Distribution: Previously known from northern California (Blake 1996a).

Chaetozone setosa Malmgren 1867 = Chaetozone sp. D = Chaetozone sp. E

Chaetozone sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 290.

Habitat: Subtidal, muddy sand, 64 m.

Chaetozone sp. B = Chaetozone hedgepethi (in part) = Chaetozone sp. X (in part)

Chaetozone sp. D

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980, (1). Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Jan 1981 (2); Apr 1981 (1); June 1981 (3); Aug 1981 (1). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Jun 1981, (1).

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72 (as Chaetozone setosa).

Habitat: Subtidal mud to sand, 11-26 m.

Chaetozone sp. E

Material Examined: Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Oct 1980, (1).

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335, Maurer et al. 1988: 46, Maurer & Vargas 1985: 464, Dean 1996a: 72 (all as Chaetozone setosa).

Habitat: Subtidal, muddy sand, 18 m.

Chaetozone sp. X

Material Examined: Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Oct 1980, (1).

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72 (as Chaetozone sp. B (in part).

Habitat: Subtidal, muddy sand, 18 m.

Cirratulus cirratulus (Müller 1766)

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291.

Habitat: Subtidal, sandy mud and muddy sand, 64-254 m.

Remarks: Blake (1996a) has indicated that many of the identifications of this species from the eastern Pacific are questionable. The Costa Rican material should be re-examined.

Cirratulus sp.

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Jan 1981 (1). Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Jul 1980 (1); Oct 1980 (3); Jan 1981 (1); Apr 1981 (2); June 1981 (3); Aug 1981 (4).

Costa Rican record: Golfo de Nicoya –Dean 1996a: 71 (as Aphelochaeta longisetosa (in part).

Habitat: Subtidal, mud and sand, 11-26 m.

Cirratulus sp. A

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291

Habitat: Subtidal, sandy mud, 70 m.

Cirratulus sp. B

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291.

Habitat: Subtidal, sandy mud, 70 m.

Cirriformia spirobrancha (Moore 1904)

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 291.

Habitat: Subtidal, sandy mud, 70 m.

Distribution: Previously reported from northern, central and southern California (Blake 1996a).

Monticellina cryptica Blake 1996

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (127); Jan 1981 (1); Jun 1981 (7). Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Oct 1980 (2). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jul 1980, (6); Oct 1980 (9); Jan 1981 (18); Apr 1981 (3); Jun 1981 (7); Aug 1981 (17); Apr 1982 (6). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Oct 1980 (1); Jan 1981 (3); Apr 1981 (4).

Costa Rican records: Golfo de Nicoya –Dean 1996a: 72 (as Tharyx sp. B).

Habitat: Subtidal, mud to sand, 11-26 m.

Distribution: Described from continental shelf depths off California by Blake (1996a).

Montecellina secunda (Banse & Hobson 1968)

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (1); Jun 1981 (2); Aug 1981 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jan 1981 (73); Apr 1981 (2). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Jan 1981 (2).

Costa Rican records: Golfo de Nicoya –Dean 1996a: 72 (as Monticellina sp. A).

Habitat: Subtidal, muddy sand to sand, 11-18 m.

Distribution: Previously reported from Washington State, USA (Blake 1996a).

Monticellina serratiseta (Banse & Hobson 1968)

Material Examined: Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Oct 1980 (1).

Costa Rican record: Golfo de Nicoya –Dean 1996a: 71 (as Monticellina tesselata (in part).

Habitat: subtidal muddy sand, 18 m.

Distribution: Known previously from shallow waters in Puget Sound, Washington, USA (Blake 1996a).

Monticellina tesselata (Hartman 1960)

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (89); Jan 1981 (1); Aug 1981 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jul 1980, (4); Oct 1980, (4); Jan 1981 (16); Apr 1981 (1); June 1981 (1); Aug 1981 (28). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Jan 1981 (1).

Costa Rican records: Golfo de Nicoya - Vargas et al. 1985: 335, Maurer et al. 1988: 46 ( both as Tharyx tesselata), Dean 1996a: 72 (in part).

Habitat: Subtidal mud to sand: 11-18 m.

Distribution: Central and southern California, USA (Blake 1996a).

Monticellina sp. A = Monticellina secunda

Monticellina sp. B

Material Examined: Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Oct 1980 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jan 1981 (1); Aug 1981 (2) Apr 1982 (1).

Costa Rican records: Golfo de Nicoya –Dean 1996a: 72 (as Monticellina tesselata (in part).

Habitat: Subtidal mud to muddy sand, 18- 26 m.

cf Tharyx epitoca Monro 1930

Costa Rican record: Golfo Dulce - Nichols Driscoll, 1976: 290.

Habitat: Subtidal, muddy sand, 64 m.

Distribution: Originally described from the Antarctic and Uschakov (1950) reported it from the Sea of Okothsk, Russia. The Costa Rican record is highly doubtful.

Tharyx marioni (Saint-Joseph 1894) = Aphelochaeta marioni

Tharyx monilaris Hartman 1960 = Aphelochaeta monilaris

Tharyx multifilis Moore 1909 = Aphelochaeta multifilis (sensu Nichols Driscoll 1976) = Aphelochaeta glandaria

Tharyx parvus Berkeley 1929

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 46, Vargas 1987: 304, Vargas 1989: 1909.

Habitat: intertidal muddy sand & subtidal mud.

Distribution: Previously reported from British Columbia, Canada, to northern California (USA) in the eastern Pacific (Blake 1996a).

Tharyx tesselata Hartman 1960 = Monticellina tesselata

Tharyx sp. A = Tharyx sp X

Tharyx sp. B = Monticellina cryptica

Tharyx sp. X

Material Examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (7); Jan 1981 (2); Apr 1981 (4); Jun 1981 (18) Aug 1981 (33). Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Jul 1980 (1); Oct 1980 (4); Apr 1981 (8); June 1981 (4); Aug 1981 (6). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jan 1981 (3); June 1981 (1); Aug 1981 (58). Sta. 30, 9º54’40" N, 84º45’50" W, 18 m, muddy sand, Oct 1980 (7); Jan 1981 (13); Apr 1981 (9); Aug 1981 (17). Sta. 33, 9º53’40" N, 84º53’20" W, 11 m, mud, Jul 1980 (1). Sta. 36, 9º55’32" N, 84º45’20" W, 18 m, sandy mud, Jul 1980 (2).

Costa Rican records: Golfo de Nicoya –Dean 1996a: 72 (as Tharyx sp. A (in part).

Habitat: Subtidal mud to sand, 11-26 m.

Cossuridae
Cossura brunnea Fauchald 1972

Costa Rican Records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47 (both as Cossura rostrata), Dean 1996a: 72. Golfo Dulce - Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal, mud to muddy sand, 11-18 m.

Remarks: This species has been recorded from Oregon to western Mexico in the eastern Pacific as well as in the western Atlantic but seems to be restricted to deeper (>1600 m) waters (Hilbig 1996). This identification needs to be substantiated.

Cossura laeviseta Hartmann-Schröder 1962

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72.

Habitat: Subtidal, mud to sand, 11-26 m.

Remarks: Reported only from its type locality in the shallow subtidal of Peru (Hartmann- Schröder 1962), thus the Costa Rican specimens need to be re-examined.

Cossura rostrata Fauchald 1972 = Cossura brunnea

Cossuridae sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 290.

Habitat: Subtidal, mud, 64-70 m.

Dorvilleidae

Anisoceras vittata Grube 1856 = Dorvillea vittata

Anisoceras bioculata Grube 1856 = Dorvillea bioculata

Dorvillea bioculata (Grube 1856)

Costa Rican record: Puntarenas – Grube 1856: 62.

Habitat: Subtidal?

Remarks: Hartman (1959) regarded this species as indeterminable

Dorvillea (Schistomeringos) annulata (Moore 1906)

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72 ( as Schistomeringos annulata).

Habitat: Subtidal, sand, 11 m.

Distribution: known from Washington (USA) to Western Mexico (Hilbig 1995a).

Dorvillea (Schistomeringos) pacifica (Westheide 1977)

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47 (both as Schistomeringos longicornis), Dean 1996a: 72.

Habitat: Subtidal, mud to sand, 11-26 m.

Distribution: Reported only from the Galapagos Islands (Westheide 1977).

Dorvillea vittata (Grube 1856)

Costa Rican record: Puntarenas – Grube 1856: 61 (as Anisoceras vittata)

Habitat: Subtidal?

Distribution: Known only from the single record from Costa Rica.

Dorvilleidae sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 83.

Habitat: Subtidal, mud, 70 m.

Meiodorvillea sp.

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47

Habitat: Subtidal, mud.

Pettibonea sp. A

Costa Rican record: Golfo de Nicoya; Dean 1996a: 72.

Habitat: Subtidal, muddy sand to sand, 11-18 m.

Pettibonea sp. B

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72.

Habitat: Subtidal, mud, 26 m.

Pettiboneia duofurca Wolf 1987

Costa Rican records: Golfo Dulce – Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal, mud.

Distribution: Known only from the Gulf of Mexico (Wolf, 1987) so this material should be re-examined.

Pettiboneia sp.

Costa Rican records: Golfo Dulce – Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal, mud.

Schistomeringos annulata (Moore 1906) = Dorvillea (Schistomeringos) annulata

Schistomeringos longicornis (Ehlers 1901) = Dorvillea (Schistomeringos) pacifica

Schistomeringos pacifica Westheide 1977 = Dorvillea (Schistomeringos) pacifica


Eulepethidae
Grubeulepis ecuadorensis Pettibone 1969

Costa Rican records: Golfo de Nicoya –Maurer et al. 1988: 47 (as Grubeulepis sp.), Dean 1996a: 72.

Habitat: Subtidal, muddy sand, 18 m.

Distribution: Previously reported subtidally in "Amuck" at 36.5 m. from the coast of Ecuador (Pettibone 1969).

Grubeulepis sp.= Grubeulepis ecuadorensis

Pareulepis sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 287.

Habitat: Subtidal, sandy mud (?), 254 m.

Eunicidae
Eunice violacea Grube 1856

Costa Rican record: Near Puntarenas –Grube 1856: 55-56

Habitat: Subtidal?

Distribution: Fauchald (1992a) regards this as an indeterminable species.

Eunice cedroensis Fauchald 1970

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72.

Habitat: Subtidal, mud to muddy sand, 20- 46 m.

Distribution: Described from Baja California, Mexico (Fauchald 1970).

Eunice lucei Grube 1856

Costa Rican record: near Puntarenas –Grube 1856: 57-58.

Habitat: Subtidal?

Remarks: Hartman (1959) listed this species as indeterminable.

Eunice mutilata Webster 1884

Costa Rican records: Golfo de Santa Elena, Parker Bay, Hartman 1944a: 113-114.

Habitat: Subtidal, sandy mud, 10-20 m.

Distribution: Common in the tropical Eastern Pacific and Western Atlantic Ocean (Fauchald 1977).

Eunice quadriocculata Grube 1856 = Marphysa quadriocculata

Eunice schemacephala Schmarda 1861

Costa Rican record: Golfo Dulce – Nichols Driscoll, 1976: 288 (as E. shemacephala (sic)

Habitat: Subtidal, mud, 70 m.

Remarks: Fauchald (1992a) regards this as an indeterminable species.

Eunice vittatopsis Fauchald 1970

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47. Golfo Dulce - Dean 1996b: 83, León-Morales & Vargas 1998: 85.

Habitat: Subtidal, mud.

Distribution: known in the Eastern Pacific from California (USA) to Panama (Fauchald 1977).

Marphysa conferta Moore 1911

Costa Rican record: Golfo Dulce – Dean 1996b: 83.

Habitat: Subtidal, mud.

Distribution: Previously reported from southern California (USA) to Baja California, Mexico (Hilbig 1995b).

Marphysa quadriocculata (Grube 1856)

Costa Rican record: Puntarenas – Grube 1856: 60.

Habitat: Subtidal?

Remarks: This species was characterized by Hartman (1944a) as incompletely known and has not been reported since its description.

Palola siciliensis (Grube 1840)

Costa Rican record: Golfo de Santa Elena, Parker Bay, Hartman 1944a: 131.

Habitat: Subtidal, in coral.

Distribution: Reported to be widespread in warmer waters but many records of this species could be suspect following Fauchald’s (1992a) review of type material.


Flabelligeridae
Pherusa sp.

Costa Rican record: Golfo Dulce – Nichols Driscoll 1976: 290.

Habitat: Subtidal, mud to sand, 70-254 m.

Pherusa inflata (Treadwell 1914)

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72.

Habitat: Subtidal, mud to sand, 11-26 m.

Distribution: Reported in the intertidal to shallow subtidal from Oregon to southern California in the eastern Pacific (Blake 2000b).

Pherusa capulata (Moore 1909)

Costa Rican record: Golfo Dulce – Dean 1996b: 83.

Habitat: Subtidal, sandy mud to sand, 75 –200 ma.

Distribution: Reported from intertidal to deep subtidal sandy sediments off the California coast (Blake 2000b).

Pherusa sp.

Costa Rican record: Golfo Dulce – Dean 1996b: 83

Habitat: Subtidal, mud.

Glyceridae
Glycera americana Leidy 1855

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47.

Habitat: Subtidal, mud.

Distribution: Reported in the Western Atlantic from Massachusetts (USA) to Argentina and in the eastern Pacific from the Straights of Magellan to Panama (Böggemann 2002).

Glycera brevicirris Grube 1870

Costa Rican records: Gulf of Nicoya –Böggemann 2002: 46

Habitat: Subtidal, muddy sand, 20-22 m.

Distribution: Widespread, reported from Costa Rica and Panama in the eastern Pacific (Böggemann 2002).

Glycera capitata Oersted, 1842 = Hemipodia californiensis

Glycera convoluta Keferstein 1862 = Glycera prosobranchia

Glycera dibranchiata Ehlers 1868 = Glycera guatemalensis

Glycera gigantea Quatrefages 1865

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47 (both as Glycera papillosa Grube).

Habitat: Subtidal, mud.

Distribution: Glycera gigantea, described from the French coast, has been synonomized with G. fallax which is known from the eastern North Atlantic and Mediterranean Sea (Böggemann 2002). The Costa Rica material needs to be re-examined.

Glycera guatemalensis Böggemann & Fiege 2001

Material examined: Golfo de Nicoya, red para camaron (shrimp trawl), Oct. 1975.

Costa Rican record: Golfo de Nicoya –Dean 1996a: 72.

Habitat: Collected by shrimp net at 36.7 m.

Distribution: Reported previously only from the type locality, Guatemala. (Böggemann & Fiege 2001).

Glycera longipinnis Grube 1878 = G. sphyrabranchia

Glycera oxycephala Ehlers 1887

Costa Rican records: Gulf of Nicoya - Vargas et al. 1985: 335, Maurer et al. 1988: 47. Caribbean, Manzanillo - Böggemann 2002: 40.

Habitat: Intertidal to subtidal sand and rubble.

Distribution: Widespread in temperate and tropical seas (Böggemann 2002).

Glycera papillosa Grube 1857 = Glycera gigantea

Glycera prosobranchia Boggemann & Fiege 2001

Material examined: Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Oct 1980 (28); Jan 1981 (4); Apr 1981 (2); Aug 1981 (7). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Oct 1980 (5); Jan 1981 (6); Apr 1981 (9); Jun 1981 (10); Aug, 1981 (6); Apr 1982 (13).

Habitat: Subtidal, sandy mud to sand, 10- 28 m.

Costa Rican records: Golfo de Nicoya –Vargas et al. 1985: 335, Maurer et al. 1988: 47, Dean 1996a: 72 (all as Glycera convoluta), Böggemann & Fiege 2001: 38-39.

Distribution: East Pacific, Mexico to Panama; Caribbean Panama (Böggemann & Fiege 2001).

Glycera robusta Ehlers 1868 = Glycera tesselata

Glycera sphyrabranchia Schmarda 1861

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47 (both as Glycera longipinnis).

Habitat: Subtidal, mud.

Distribution: This species is known from tropical seas including both the Atlantic and Caribbean coasts of Panama (Böggemann 2002).

Glycera tesselata Grube 1863

Material examined: Golfo de Nicoya, Sta. 24, 9º49’25" N, 84º41’20" W, 11 m, sand, Aug 1981 (1). Sta. 28, 9º52’16" N, 84º45’30" W, 26 m, mud, Oct 1980 (1). Sta. 29, 9º54’55" N, 84º45’15" W, 18 m, muddy sand, Jan 1981 (5); Apr 1982 (1). Sta. 34, 9º55’30" N, 84º50’05" W, 24 m, sand, Jul 1980, (1).

Habitat: Subtidal, in broken shell sediments and mud to sand, 18-46 m.

Costa Rican records: Golfo de Santa Elena, Playa Blanca - Hartman 1940: 247. Golfo de Nicoya - Dean 1996a: 72 ( as G. robusta).

Distribution: Böggeman (2002) states that this species is found on both sides of the north Atlantic, the Red Sea, the Mediterranean, both sides of southern Africa, the Caribbean Sea, and the Gulf of Mexico. He reported its distribution in the Pacific as from the Indo-Pacific and the northwestern Pacific but mentions that he has examined the type of Glycera fundicola Chamberlin 1919 from Peru and believes it to also be G. tesselata.

Hemipodus armatus Hartman 1950 = Hemipodia pustatula

Hemipodia pustatula (Friedrich 1956)

Material examined: Punta Morales, intertidal sandy mud (3).

Habitat: Intertidal, sandy beaches

Costa Rican records: Caribbean, Cahuita & Puerto Viejo - Fauchald 1973: 22 (as Hemipodus armatus Hartman 1950), Limon Cahuita, & Puerto Viejo - Dexter, 1974: 56 (as Hemipodus armatus Hartman 1950), Carretera, Cahuita, Puerto Vargas, Manzanillo & Gandoca - Böggemann 2002: 83. Pacific coast, Samara & Tamarindo - Fauchald 1973: 22 (as Hemipodus armatus Hartman 1950), Playa Matapalo, Guiones, Playa Sámara, Playa Carrillo, Punta Islita, Montezuma – Böggemann 2002: 83.

Distribution: Böggeman (2002) reports this species from the western Atlantic coast of South America and from Mexico to Chile in the eastern Pacific. Böggeman (2002) feels that Hemipodia armata (Hartman 1950) (= Hemipodus armatus) may be a variant of H. pustatula in which case the name Hemipodia armata would take precidence

Hemipodia californiensis (Hartman 1938)

Habitat: Intertidal and subtidal, mud to sand, 11-26 m.

Costa Rican records: Pacific coast, Golfo de Nicoya - Vargas et al. 1985: 335, Maurer et al. 1988: 47 (both as Glycera capitata), Dean 1996a: 72 (as Hemipodus simplex (Grube 1857), Playa Matapalo and Playa Flamingo –Böggemann 2002: 82. Golfo Dulce – Dean 1996b: 83, León-Morales & Vargas 1998: 85 (all as Glycera capitata).

Distribution: Reported in the eastern Pacific from California (USA) to Peru as well as southeastern South America (Böggemann 2002).

Hemipodus simplex (Grube 1857) = Hemipodia californiensis


Goniadidae
Glycinde armigera Moore 1911

Costa Rican records: Golfo de Nicoya -Vargas et al. 1985: 335, Maurer et al. 1988: 47, Vargas 1987: 304, 1988: 294, 1989: 1909. Golfo Dulce - Nichols Driscoll 1976: 288.

Habitat: Subtidal, mud, 64-254 m.

Вот она показалась опять, с нелепо скрюченными конечностями. В девяноста футах внизу, распростертый на острых лопастях главного генератора, лежал Фил Чатрукьян. Тело его обгорело и почернело.

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