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(Annelida: Polychaeta) from the Eastern Tropical Pacific Ocean

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(Annelida: Polychaeta) from the Eastern Tropical Pacific Ocean BULLETIN OF MARINE SCIENCE, 67(1): 45–53, 2000 TOMOPTERIDS (ANNELIDA: POLYCHAETA) FROM THE EASTERN TROPICAL PACIFIC OCEAN María Ana Fernández-Álamo ABSTRACT The distribution and composition of species of the holoplanktonic family Tomopteridae are presented. The material was taken from 579 zooplankton samples collected on 12 cruises made by the international oceanographic Program “EASTROPAC” during five of 2–3-mo periods between January–March 1967 to January–April 1968. A total of 4110 specimens were identified consisting of only one genus and eight species. Tomopteris elegans (61%) and T. planktonis (25%) were the dominant species during this period. T. euchaeta, and T. dunckeri are reported for the first time from the Pacific Ocean and, T. krampi is recorded for the first time in the southeastern region of the Pacific. Little has been published on holoplanktonic polychaetes from the eastern tropical Pa- cific Ocean. A few scattered records from this area can be found in the works of Chamberlin (1919), Treadwell (1943), Berkeley and Berkeley (1961, 1964) and Dales (1957). Phylo- genetic relationships of the famly Tomopteridae are not clear. Tomopterids have been considered as aberrant worms with problematic relationships to other polychaetes (Støp- Bowitz, 1948, Tebble, 1960). Due to their unique morphology and uncertain affinities, Mileikovsky (1977) erected the order Tomopterimorpha and Pleijel and Dales (1991) the superfamily Tomopteroidea. In a cladistic analysis of the Polychaeta Rouse and Fauchald (1997) excluded the Tomopteridae and other pelagic taxa from their restricted analyses, because their unusual morphology is associated with the pelagic existence and their level of study would have provided spurious results in their relationships with other polycha- etes. In their analyses, Tomopteridae is included in the Phyllodocida group. Most studies of tomopterids had been made on systematics and distribution, and al- most nothing is known about their biology and ecology. It is assumed that they are vora- cious predators, nevertheless nothing has been observed in their transparent guts (Ushakov, 1972). Pettibone (1963) also considers them as voracious predators in the plankton, and some species may at times be the dominant forms on the plankton community and must be of considerable importance as food of fishes. The eastern tropical Pacific Ocean is characterized by a very well-developed thermocline separating warm surface water from cooler subsurface water (Wyrtki, 1964). Brandhorst (1958) and Reid (1962) have shown that the divergences and convergences associated with the surface circulation result in the formation of ridges and troughs in the topogra- phy of the thermocline. These processes have a particular importance for fertilization of the surface layer and for the distribution of the standing crop of zooplankton in this re- gion. Wyrtki (1965) has described the general surface circulation of this region. To under- stand and to predict the nature of variation in the distribution of physical and biological properties in the eastern tropical Pacific, an international oceanographic program was developed named “EASTROPAC”. Field expeditions were carried out between January 1967–April 1968, and the study area was defined as latitude 20ºN to latitude 20ºS and from the coast of the American continents westward to longitude 126ºW (Love, 1972). 45 46 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000 MATERIALS AND METHODS The tomopterid polychaetes were sorted from zooplankton collections obtained on 12 expedi- tions during the “EASTROPAC” program. The physical and biological properties, nutrient chemis- try and meteorological data from the principal participating ships in this program are found in the EASTROPAC Atlas edited by Love (1972). This Atlas listed the stations data including the latitude, longitude, depth and date of collections. Because the description of the 579 stations examined would require so many pages, the reader may contact the author for station data. Zooplankton oblique hauls were taken with nets of 0.50 and 1.0 m mouth diameter, a length of 3.5 m in a paired-frame and constructed with of Nitex nylon with mesh apertures of 0.333 and 0.555 mm, respectively. Hauls were taken between about 200 m and the surface. The zooplankton samples were preserved at sea in buffered 10% formalin seawater (Laurs, 1972). The samples were split into 50% aliquots. One set was deposited in the Smithsonian Oceanographic Sorting Center where the worms were sorted. Generally the aliquot was sorted, but some times only a subsample of the aliquot was examined. A total of 579 samples were analyzed from five periods: January–April, April–May, June–July, and August–September 1967, and January–April 1968. Figure 1A,B,C,D show the stations localization of the 12 cruises. The geographical position of these stations is re- ported in detail in EASTROPAC Atlas (Love, 1972). Family Tomopteridae Grube, 1848 Tomopterids comprise polychaetes that lead a pelagic existence spending their entire life cycles in the water column. Principal adaptations to this mode of life are parapodial modification to biramous achaetous paddles with notopodia and neuropodia adapted for swimming into membranous structures named pinnules. There are never have more than two pairs of chaetigate processes which occur in the first pairs of parapodia. In the second pair, the acicula is prolonged and may extend as long as, or longer than the body. Day (1967) has considered Tomopteris as the only valid genus of this family, and this opinion is followed in this report. Tebble (1960) considers the division of Tomopteris in two sub- genera artificial and not practical. Genus Tomopteris Eschscholtz, 1825 Tomopteris euchaeta Chun, 1887 Material Examined.—207 spec. from 97 stations ranging from the coasts of Mexico (south of Cape Corrientes, 20°N) to Peru (south of Callao, 20°S), and westward to longi- tude 125°W, during five 2-mo cruise periods between January 1967 and April 1968. Type locality.—Mediterranean Sea. Remarks.—Rosa (1908) found this species in the Atlantic near the Caribbean Sea. He erected the genus Enapteris based on the structure of parapodia with rectangular rami and with the pinules only in the external part of the parapodial rami. Most authors have concurred; however, Day (1967) stated that Rosa’s specimens were juvenile stages. A study of the adults showed that the pinnules were divided into an outer frilly area bearing the glands, and an inner clear area which simulates a flattened extension of the ramus. I observed this character in all specimens studied in this survey which agrees with Day’s FERNÁNDEZ-ÁLAMO: TOMOPTERIDS FROM THE EASTERN TROPICAL PACIFIC 47 Figure 1. Map of the Eastern Tropical Pacific Ocean showing where the zooplankton samples were collected. A) from January to April, 1967; B) from April to August, 1967; C) from August to September, 1967; D) from January to April, 1968. opinion. The size of these specimens ranged from 4 to 19 mm with a mean of 8.4 mm, and 12–15 pairs of parapodia. Distribution.—Mediterranean Sea (Fauvel, 1923), Atlantic (Apstein, 1900) (Rosa, 1908; Støp-Bowitz, 1948; Day, 1967; Orensanz and Ramirez, 1973, as Enapteris euchaeta), Southwest region of Indian Ocean (Day, 1967). There are very few records in the Pacific Ocean, Day (1967) pointed out that T. euchaeta has been recorded in the North Pacific as T. nisseni, but he did not indicate the source for this information. This report validates the first record of T. euchaeta in the Pacific Ocean which was identified in an unpublished thesis (Fernández-Álamo, 1983). Rozbaczylo et al. (1987) recorded one specimen as Enapteris euchaeta collected off Coquimbo, Chile. Fernández-Álamo (1992, 1996) pro- vided the first records of it in the Gulf of California, and off the southwestern coast of Baja California, Mexico, respectively. Vicencio-Aguilar and Fernández-Álamo (1996) recorded it from the Costa Rica Dome, Central America. 48 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000 Tomopteris nationalis Apstein, 1900 Material Examined.—191 spec. from 93 stations ranging from the coasts of Mexico (south of Cape Corrientes, 20°N ) to Ecuador (south of Gulf of Guayaquil, 11°S), and westward to longitude 125°W, during five 2-mo cruise periods between January 1967 and April 1968. Type locality.—Naples, Italy. Remarks.—The systematic position of the species seems to be unclear since most au- thors have identified it as T. apsteini Rosa, 1908. However, Day (1967) mentioned that it is probably synonymous with T. nationalis Apstein, 1900. Fernández-Álamo (1983) agreed with this opinion because in Rosa’s paper the name “Tomopteris apsteini nom. nov” was used to redescribe specimens from Messina, Italy. His description is nearly identical with the description of T. nationalis in the same publication. In the survey region the size of the specimens ranged from 3 to 16 mm, with a mean of 6.5 mm, and 13–20 pairs of parapo- dia. Distribution.—Mediterranean Sea (Malaquin and Carin, 1911, as T. apsteini); Atlantic (Apstein, 1900; Rosa, 1908; Day, 1967), (Støp-Bowitz, 1948; Tebble, 1960, as T. apsteini). Previous records in the Pacific are scarce, probably T. eura Chamberlin (1919) found off Peru could be T. nationalis. Dales (1957) recorded it in the California Current. Berkeley and Berkeley (1964), Tebble (1962), and Sun and Wu (1979) reported it as T. apsteini off the coast of Peru in the subtropical region of the North Pacific, and China Sea, respec- tively. Fernández-Álamo (1991) made the first record from the Gulf of California, Fernández-Álamo (1996) found it off southwestern coast of Baja California, Mexico. Tomopteris dunckeri Rosa, 1908 Material Examined.—134 spec. from 83 stations ranging from the coasts of Mexico (south of Cape Corrientes, 20°N ) to Ecuador (north of Gulf of Guayaquil, 2°S), and westward to longitude 119°W, during five 2-mo cruise periods between January 1967 and April 1968. Type locality.—Ceylon (Sri Lanka), Indian Ocean.
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