Diversity and Distribution of Chilean Benthic Marine Polychaetes: State of the Art

BULLETIN OF MARINE SCIENCE, 67(1): 359–372, 2000

DIVERSITY AND DISTRIBUTION OF CHILEAN BENTHIC MARINE POLYCHAETES: STATE OF THE ART

Nicolás Rozbaczylo and Javier A. Simonetti

ABSTRACT Current knowledge of Chilean benthic marine polychaetes is reviewed. The history of the studies, researchers involved, the rate and localities of species descriptions are pre- sented, as they relate to the assessment of biogeographic units along the Chilean coast. Taxonomic richness along the Chilean coast is associated with differential sampling efforts weakening the assessment of biogeographic units.

Juan Ignacio Molina (1740–1829), in his pioneer work on the biological diversity of Chile, Saggio sulla storia naturale del Chili published in 1782, described numerous invertebrate species. However, he did not refer to polychaetes. The first descriptions of polychaetes from continental Chile were published in 1849. Based on specimens collected by the French naturalist Claudio Gay, all 15 species were described by Blanchard (1849) as new to science (Rozbaczylo, 1985). During the following years, the knowledge of Chilean polychaetes increased due principally to the activity of foreign expeditions and researchers who visited various places of Chile. In the last 30 yrs, the taxonomic knowledge of polychaetes has been increasing largely due to the work of Chilean scientists who have done studies on local faunas or reviewed specific groups of polychaetes. With almost 450 species known, this figure is considered an underestimate due to the overall scarcity of research carried out on Chilean polychaetes (Rozbaczylo and Carrasco, 1995). In this work we analyze the knowledge of benthic polychaetes along the Chilean continental coast and oceanic islands, in terms of the taxonomic richness, the main expeditions that have collected specimens in these areas, the researchers who have studied this material, and location of the main collections containing type specimens of Chilean species, as they relate to the assessment of faunal provinces. The recognition of biogeographic regions along the coast of Chile has been based on the distribution patterns of several invertebrate taxa including polychaetes. However, faunal studies along these geographical areas have been unequally carried out, being concentrated mainly in the central and southern regions while the north has been comparatively less studied which might bias the distinction of biogeographical regions based on invertebrate distributions.

TAXONOMIC RICHNESS AND ENDEMICITY OF MARINE BENTHIC POLYCHAETES IN CONTINENTAL CHILE AND OCEANIC ISLANDS

CONTINENTAL CHILE.—The coast of mainland Chile is about 4200 km long, extending from Arica (18°20'S) to Cape Horn in subantarctic waters (56°S) (Fig. 1). Some authors, such as Olsson (1961) and Marincovich (1973), based on the distribution of different groups of invertebrates, distinguished two biotic provinces along the temperate coast of Pacific South America, named Peruvian and Magellanic, and considered the Chiloé Is- land (ca 42°S) the boundary between them. Others, e.g., Dahl (1960) and Viviani (1979), have limited the southern extent of the Peruvian Province to ca 30°S, recognizing a tran-

359 360 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000

Figure 1. Map showing the main geographical regions considered for comparison of polychaete fauna along mainland Chile and the location of oceanic islands (drawn not to scale). ROZBACZYLO AND SIMONETTI: CHILEAN BENTHIC POLYCHAETES 361

Table 1. Taxonomic composition of marine benthic polychaetes in continental Chile compared with the World.

Teaxononmic level C*hil World O5rders 1 . (560%) 2 F6amilies 4 . (753%) 8 G4enera 21 . (021%) 1.00 S9pecies 44 . (5%) >8.000 *Pettibone (1982)

sition zone of cold temperate mixed waters between 30 and 42°S. Knox (1960) character- ized this zone as the Central Chilean Province, but others, Dahl (1960) and Dell (1971), have insisted on its transitional character. The different propositions about biogeographic provinces and provincial boundaries along Chile have been recently reviewed by Santelices (1991). World-wide, the taxonomic richness of polychaetes is estimated at more than 8000 species, grouped in 25 orders, 87 families and around 1000 genera (Fauchald, 1977, Pettibone, 1982). Along the coast of continental Chile, from Arica to Cape Horn, a total of 46 families belonging to 15 orders, with 214 genera and 449 species have been registered so far (Table 1). This region presents high endemicity at the species level estimated to be around 43% (Table 2). Similarly, high values of endemicity are also depicted by other groups of marine invertebrates, for instance, 55.6% of the littoral mollusks (Dall, 1909), 53.3% of the anomuran decapods (Haig, 1965), and 23.3% of the brachyuran crabs (Garth, 1957). The number of benthic polychaete species differs markedly among the three main geo- graphic regions along continental Chile (Fig. 1): northern Chile from Arica (18°28'S) to south of Coquimbo (32°S), central Chile from north of Valparaíso (32°S) to south of Chiloé Island (44°S), and southern Chile from Golfo de Corcovado (44°S) to Cape Horn (56°S). The number of species as well as genera and families are fewer in the northern region and greatest in the central region while the southern region presents intermediate values (Table 3). This tendency parallels the number of authors and publications about polychaetes in each of the regions (Table 4), strongly suggesting that the number of species both recorded and described in each of these regions are more an artifactual result than a biological phenomenon. Unpublished records of polychaetes from northern Chile collected in Antofagasta (23°29'S) and Carrizal Bajo (28°04'S) in 1996 and 1997 by one of us (NR), have indicated that at least three families and 20 species are new for these localities. Therefore, the comparatively low number of species known for the northern region of Chile could be explained by the low number of samples taken.

Table 2. Taxonomic richness and percentage of endemism of marine benthic polychaetes in continental Chile and oceanic islands*.

Asrea Faamilie Gsener Smpecie Endemis ()n) ()n ()n (% C6ontinental Chile 44291 434 4 . E5aster Island 205064.** Jáuan Fern n8dez Archipelag o 1633452 . (é*) no data for San F ló.ix and San Ambrosio Islands and Sala y Gmez Island (**) calculated from species identified to specific level 362 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000

Table 3. Taxonomic richness of marine benthic polychaetes occurring in northern, central and southern Chile and shared by the three regions.

Rsegion Faamilie Gsener Specie ()n) ()n (n N°'orthern Chile (18 28 −3°2 S ) 259 7611 C°entral Chile (32 −4°4 S ) 454 197 32 S°outhern Chile (44 −5°6 S ) 335 121 16 S1hared taxa 15102

CHILEAN OCEANIC ISLANDS.—Chile’s main oceanic islands, defined as those located in an ocean at a long distance from a continent, or those which emerge from the ocean floor and are isolated from the continent by water depths exceeding 200 m (Castilla and Oliva, 1987), are: (1) Easter Island, (2) the islands of the Juan Fernández Archipelago, (3) San Félix and San Ambrosio islands (formerly “Desventuradas”), and (4) Sala y Gómez Island. They are of volcanic origin and emerged in the Plio-Pleistocene (Gonzalez-Ferrán, 1987). They are of special interest to biologists given their geographical isolation and the possibility that through the study of their fauna and that of the continental shelf could be possible to unravel their zoogeographical relations as well as probable paths of colonization. The only information available about benthic polychaetes of these islands comes from Easter Island and the Juan Fernández Archipelago. There are neither records for San Félix and San Ambrosio islands nor for Sala y Gómez. The scientific knowledge for San Félix and San Ambrosio islands ( 26°17'S) is limited and recently emerging (Bahamonde, 1987, Rozbaczylo and Castilla, 1987). The tiny island of Sala y Gómez, located at 26°27'S, 3400 km west of the Chilean coast and 415 km west of Easter Island, has been rarely visited by expeditions in spite of its proximity to Easter Island (Rehder, 1980; Castilla and Rozbaczylo, 1987). EASTER ISLAND.—Easter Island (27°09'S, 109°3'W), named Rapa Nui (Great Island) or Te Pito o Te Henua (Navel of the world) in the native language, is located 3700 km off the South American continent and 4050 km from Tahiti, in Polynesia. Pitcairn Island, located 2200 km to the west, is the nearest inhabited island. At least 60 species of benthic polychaetes belonging to 25 families have been recorded from Easter Island, but only 47 of these have been identified to species, whereas the remaining 13 taxa are known only to genus or family level (Table 2). These values do not include the pelagic polynoid Podarmus ploa Chamberlin, 1919 described from off Easter Island. The polychaete fauna of Easter Island conforms to the pattern of tropical Indo-west Pacific affinity as in several other groups of marine invertebrates (Castilla and Rozbaczylo,

Table 4. Number of species of marine benthic polychaetes described for northern, central and southern Chile compared with the number of authors and publications in each geographical region.

Region Sdpecies dfescribe Number o Number of authors publications Northern Chile (18°'28 −3°2 S ) 289 1 0 Central Chile (32°−4°4 S ) 1268 204 Southern Chile (44°−5°6 S ) 648 122 T0otals for Chile 247 346 ROZBACZYLO AND SIMONETTI: CHILEAN BENTHIC POLYCHAETES 363

1987). Considering percentage of identified polychaetes, approximately 4% are endemic. However, according to Rehder (1980), the relatively high values of endemicity shown by other groups of marine animals of Easter Island justify Schilder’s (1965) placement of Easter Island and Sala y Gómez island in the Rapanui Province, rather than the Polynesian Province where they were previously located (Schilder, 1938–1939 fide Rehder, 1980). THE JUAN FERNÁNDEZ ARCHIPELAGO.—The Juan Fernández Archipelago (33°40'S, 79°00'W), located at 587 km from the South American continent, is composed of three islands: Robinson Crusoe, Santa Clara and Alejandro Selkirk. The number of polychaete species currently known in the Juan Fernández Archipelago is estimated at 43, belonging to 18 families, but only 37 have been determined to species; Spionidae with eight species, Syllidae with seven, and Nereididae with five species are the most diverse families. As with coastal northern Chile, the relatively low number of polychaete species recorded from the Juan Fernández Archipelago could be explained by the low frequency and ineffective studies carried out until now in this locality (Rozbaczylo and Castilla, 1987). From the 37 recognized species, 25% are considered endemic to the archipelago (Table 2).

HISTORICAL SUMMARY ABOUT MAIN CONTRIBUTIONS TO THE KNOWLEDGE OF CHILEAN POLYCHAETES ALONG CONTINENTAL CHILE AND CHILEAN OCEANIC ISLANDS

CONTINENTAL CHILE.—The knowledge of Chilean polychaetes began from the mid-19th century. The French zoologist Emil Blanchard studied the specimens collected by the French naturalist Claudio Gay, mainly from Chiloé (Ancud and Castro) and from Calbuco. It was published in Gay’s 1849 Historia Física y Política de Chile, with the description of 15 species. Of these, only three are currently regarded as valid, one was placed in syn- onymy while the remaining 11 are considered as indeterminable because of inadequate original descriptions (Hartman, 1959; Rozbaczylo, 1985). During the following 50 yrs, the number of described species increased rapidly and by the year 1901, 115 species had been described, which accounted for 43% of the currently known species in continental Chile. This work was carried out by European scientists such as Kinberg, Ehlers, McIn- tosh, Grube and Schmarda (Table 5). In a series of six works published between 1855 and 1867, Kinberg described 23 species from Chile, collected between January and February of 1852, mainly in the Strait of Magellan and Valparaíso by the Swedish expedition around the world (1851–1853) aboard the Frigate EUGENIE. The specimens deposited in the Swedish State Museum of Natural History at Stockholm were reviewed a century later by Hartman (1948). Similarly, Grube (1856, 1858), in his Annulata Örstediana described 14 new species from Valparaíso, Chile. The German annelidologist Ernst Ehlers published during 1897, 1900 and 1901 (a,b) four important works in which he describes 40 new species from Chile. The first of these works refers mainly to the species collected between the years 1892–1893, in the Magellan area, by the German zoologist W. Michaelsen. In his 1900s publication, Ehlers describes the specimens collected between the years 1895–1897 in the Magellan area and the Beagle Channel, by Ohlin and Ackerman, members of a Swedish expedition in charge of the geologist Otto Nordenskjöld. In 1901, he describes the species collected between the years 1893–1895 by the German zoologist Ludwig H. Plate, along the Chilean coast and the Juan Fernández Archipelago. His work of 1901(b) is a monograph of polychaetes from Peru to Cape Horn in which included complete characterizations and figures of the 364 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000

Table 5. Authors who have described polychaete species from continental Chile, years of descriptions and number of species described. (For most of these references see Rozbaczylo, 1985)

Authors and years of descriptions Species described Höartmann-Schr d5er (1962, 1963, 1965, 1991 ) 10 Ehlers (1897, 1900, 1901a, b) 40 Kinberg (1855, 1857, 1858, 1865, 1866, 1867) 23 McIntosh (1885) 13 Grube (1856, 1857, 1858, 1867, 1870) 12 Schmarda (1861) 11 Carrasco (1976,1977, 1981, 1983, 1987) 10 Hartman (1967) 7 Rozbaczylo et al. (1970, 1973, 1975, 1981, 1993, 1996) 6 Wesenberg-Lund (1962) 5 Caullery and Mesnil (1897) 4 Augener (1912, 1918, 1923) 3 Baird (1871) 3 Blanchard (1849) 3 Quatrefages (1865, 1866) 3 Blake (1979, 1983) 2 Fauchald (1974, 1980) 2 Monro (1936) 2 Arwidsson (1899) 1 Blake and Woodwick (1971) 1 Gallardo (1969) 1 Glasby and Hutchings (1987) 1 Hessle (1917) 1 Jeldes (1963) 1 Knight-Jones (1978) 1 Kucheruk (1979) 1 Kük1enthal (1887 ) Malard (1891) 1 Nilsson (1928) 1 Seidler (1922) 1 Södöerst r m1(1920 ) Treadwell (1924) 1 Wells (1963) 1 Westheide (1994) 1

majority of the species previously collected as well as descriptions of new ones. McIn- tosh (1885) was in charge of the polychaetes collected by the British expedition aboard H.M.S. CHALLENGER, 1873–1876. A total of 13 species were described from specimens obtained at great depths mainly at the Magellan area and off the coast of Valparaíso. The French expedition Mission du Cap Horn, aboard the LA ROMANCHE, explored the Strait of Magellan, the Beagle Channel and the archipelago of Cape Horn between the years 1882– 1883, however, the report on the polychaetes collected was published in 1941. A total of 43 species, including some obtained separately in Punta Arenas and in the Kerguelen islands, were studied by Fauvel (1941), but none were new species. Caullery and Mesnil ROZBACZYLO AND SIMONETTI: CHILEAN BENTHIC POLYCHAETES 365

Figure 2. Number of polychaete species from continental Chile described from 1849 to 1996.

(1897) studied the Spirorbidae collected during this expedition and described four new species. A period of low activity, expressed as number of species described, number of authors, and works published, occurred between 1902 and 1960 (Fig. 2). In that period, only seven authors in nine publications (Augener, 1912; Hessle, 1917; Augener, 1918; Söderström, 1920; Seidler, 1922; Augener, 1923; Treadwell, 1924; Nilsson, 1928; and Monro, 1936) described 10 new species from Chile. This figure contrasts with the 118 new species described between 1962 and 1969. This means that not until the beginning of the 1970s were 90% of species currently known from Chile described primarily by Wesenberg- Lund (1962), Hartmann-Schröder (1962a,b, 1963, 1965) and Hartman (1967) (Table 5). Wesenberg-Lund (1962) described the polychaeta (Errantia) collected during the Swedish expedition Lund University Chile Expedition, 1948–1949, under the direction of Brattström and Dahl. The sampling area of this expedition was concentrated mainly in Seno Reloncaví, Estero Reloncaví and Canal de Chacao (ca 41°S), extending southward to the Strait of Magellan and northward to Iquique (20°11'S). Most of the polychaetes come from shal- low waters or from beaches at low tide and rocky ponds, and a few samples from depths greater than 200 m. Taxa from 21 families were collected, with Chrysopetalidae and Sphaerodoridae being recorded for the first time. Five out the 87 species were new species and 33 were new to Chile. The Arenicolidae collected during this expedition were studied by Wells (1954, 1963) and Blake and Woodwick (1971) described Boccardia chilensis from this material. As a result of her stay in Chile, the German polychaetologist Hartmann-Schröder published a series of works representing a valuable contribution to the knowledge of the polychaete fauna of the southeast Pacific ocean and particularly of Chile (Hartmann- Schröder 1962a,b, 1963, 1965). In her 1962(b) paper, she reviewed the eulittoral species 366 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000

Table 6. Number of Chilean taxonomists** working with different taxonomic groups of animals. The number of taxonomists is not additive as some scientists devote time to more than one group (Simonetti, 1997).

Taxonomic group Number of taxonomists (%) I9nvertebrates 6 . (44) M0olluscs 1 . (6) C8rustaceans . (5) P4olychaetes . (3) E4chinoderms . (3) A1rachnids . (>1) I2nsects 3 . (21) O8thers* 2 . (18) V9ertebrates 4 . (31) F8ishes 1 . (12) A4mphibians 1 . (9) R2eptiles 1 . (8) B3irds . (2) M9ammals . (6) * Includes sponges, cnidarians, helminths, sipunculans, echiurans, priapulans, phoronids, brachiopods, bryozoans, chaetognaths, hemichordata, collembolans and proturans. ** In Chile, at least, 156 scientists are totally or partially devoted to taxonomic and systematic studies of biota. Approximately 24% (38) of these are botanists sensu lato (including algae to true vascular plants); the remaining 76% are zoologists. collected at various places along the coast of Chile and in subpatagonian coast of Argen- tina. Subsequently, sublittoral species from Chile especially those obtained between Coquimbo (30°S) and Chiloé Island (43°S) were published (Hartmann-Schröder, 1965). This work was conducted by the oceanographic expedition “Marchile I,” in the year 1960 aboard the corvette CHIPANA of the Chilean Navy. Hartmann-Schröder’s most recent work on Chilean polychaetes refers to species collected in Bahía Quillaipe, Seno de Reloncaví; 19 species were found, two of which were new to science (Hartmann-Schröder, 1991). In four publications, Hartmann-Schröder (1962b, 1963, 1965, 1991) described more than 100 species from Chile, currently accepted as valid. This figure doubles the contribution by Ehlers (1897, 1900, 1901a, 1901b), who with up to 40 species is considered the second most important author in terms of the number of polychaete species described from Chile, while Kinberg (1855, 1857, 1858, 1865, 1866, 1867) stands in a third place, with the description of 23 valid species. Hartman (1967), studied the species collected by the USNS ELTANIN expedition, 1962– 1966, mainly in Antarctic seas, from surface to depths greater than 5000 m. Seven species from western Chile (33°S), off Southwestern Chile (50°S), Strait of Magellan and Cape Horn area, were considered as new to science (Rozbaczylo, 1985). From the 1970s, the taxonomic and ecological knowledge of the polychaete fauna of continental Chile has increased with the contributions of Chilean investigators, namely Gallardo and Carrasco from Universidad de Concepción, Rozbaczylo and Castilla from Pontificia Universidad Católica de Chile, Santiago, and Cañete from Universidad de Magallanes, Punta Arenas. Current taxonomic studies on benthic polychaetes in Chile, are done primarily by four of these investigators, who represent around 3% of all Chil- ean scientists devoted totally or partially to taxonomic and systematic studies of biota (Table 6). ROZBACZYLO AND SIMONETTI: CHILEAN BENTHIC POLYCHAETES 367

Figure 3. Comparison between number of benthic polychaetes registered in northern, central and southern Chile and taxa shared by the three regions.

CHILEAN OCEANIC ISLANDS—Easter Island.—Six publications account for all work on the polychaetes of Easter Island (Chamberlin, 1919; Augener, 1922; Fauvel, 1936; Hartmann-Schröder, 1962b; Rozbaczylo and Castilla, 1988, and Cañete, 1997). The first specimens collected at Easter Island were obtained in 1904, during the third expedition (1904–1905) of the U.S. Fish Commission Steamer ALBATROSS to the Eastern Tropical Pacific, under the direction of Alexander Agassiz. The four species collected on the “beach of Easter Island”, Eurythoe complanata, Perinereis helleri, Cirriformia filigera nesophila and Arabella mutans, were studied by Chamberlin (1919). Augener (1922) mentioned Eurythoe complanata, found under rocks in the littoral region of Bahía Hanga Roa, as the only polychaete species collected on Easter Island, during the Swedish Pacific Expedi- tion, 1916–1917, under the direction of Carl Skottsberg. The material studied by Fauvel (1936) corresponded to a collection of eight species collected in Bahía Hanga Roa and Bahía Cave [sic], in 1934 by members of the Franco-Belgian Mission to Easter Island. Of the species collected, six were new records for the island: Pherecardia striata, Phyllodoce madeirensis, Platynereis dumerilii, Palola siciliensis, Lysidice nineta [= L. collaris], and Loimia medusa. In 1962, Hartmann-Schröder added another species, when she recorded Nereis callaona (Grube) from the island. Kohn and Lloyd (1973), reported at least 49 species from sand with associated algae, and a basalt boulder, from a tide pool between Hanga Roa and Hanga Piko, which was collected by John E. Randall, in 1969. Scolelepis anakenae, described by Rozbaczylo and Castilla (1988), from Anakena beach, is the only polychaete species recorded to date in a sandy beach at the island. The most recent work on the polychaetes of Easter Island is by Cañete (1997) who reviewed a small collection of Polynoidae collected from intertidal and subtidal shores which increased the number of known species of this family, from one to five. Most of the known polychaetes of Easter Island come from the intertidal zones between Bahía Hanga Roa and Hanga Piko, whereas the subtidal zone of the island remain practically unex- plored. THE JUAN FERNÁNDEZ A RCHIPELAGO.—The taxonomic knowledge on the polychaetes from the Juan Fernández Archipelago is due mainly to the works of Ehlers (1901a), Augener (1922) and Blake (1983). Ehlers identified a total of 18 species collected in Juan Fernández, between 1893–1895, by Dr. Ludwig H. Plate, eight of which were described as new. Augener (1922) studied the specimens collected during the Swedish Pacific Expedition of 1916–1917, and reviewed the material from the Plate collection previously studied by Ehlers (1901a). The monumental work The Natural History of Juan Fernández and Eas- ter Island, based on the Swedish Pacific Expedition constitutes one of the most important studies on the natural history of the Juan Fernández Archipelago. Seven new species and 368 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000

subspecies were described by Augener which almost doubling the number of known species. More recently, Blake (1983) studied the spionids collected in 1965-1966 by the RV ANTON BRUUN, as part of the “Southeastern Pacific Biological and Oceanographic Pro- gram”. LOCATION OF MAIN COLLECTIONS WITH TYPE MATERIAL OF CHILEAN POLYCHAETES.—Only two Chilean institutions hold documented collections of polychaetes in a well preserved state: the Museo de Zoología, Universidad de Concepción, Concepción and the Sala de Sistemática, Departamento de Ecología, Pontificia Universidad Católica de Chile, Santiago. Numerous collections of polychaetes collected by foreign expeditions and those of visit- ing investigators are deposited in foreign institutions without duplicates in Chilean museums. Similarly, practically all of the types and paratypes of the described species from the coasts of Chile and oceanic islands remain deposited in various institutions such as the British Museum (Natural History); Muséum National d’Histoire Naturelle, in Paris; Na- tional Museum of Natural History, Smithsonian Institution of Washington, DC; Naturhistoriska Riksmuseet Stockholm in Sweden; Uppsala Universitets Zoologiska Museum of Uppsala; Zoological Museum of the University of Copenhagen; Zoologisk Museum of Copenhagen, and the Zoologisches Institut und Zoologisches Museum der Universität Hamburg of Germany. Only about 4% of the type specimens (holotype and/or paratypes) of polychaetes from Chile are deposited in Chilean museums which represents a serious problem for the taxonomic and systematic study of the Chilean fauna. Accord- ing to Chilean legislation, type specimens should be deposited in the National Museum of Natural History in Santiago. If any new species is described, the request should be followed. Furthermore, and ideally, duplicate specimens of material collected in Chile should to be placed in a Chilean museum collection.

DISCUSSION

Chilean polychaetes have been studied for over 150 yrs. New species to science have been described at a rate of two species per year. This figure is due largely to the significant work by Hatmann-Schröder in the 1960s. If her monumental effort, is excluded from calculations, the historical rate of description is one species per year. During the last two decades, such rate is 0.9 species per year, suggesting that the apparent decrease in the addition of new species or records is more a consequence of a reduced taxonomic work than the fact that fewer species would remain to be described, particularly in oceanic islands. Therefore, current taxonomic richness might be an underestimation of the diversity of Chilean polychaetes. In fact, at least two new species have been described recently for northern Chile (Carrasco and Palma, 2000). This fact suggests that the current taxonomic richness might be an underestimation of the diversity of polychaetes. Furthermore, sampling efforts have been unevenly distributed along the Chilean coast and oceanic islands. As more works have been performed in a region, the more species-rich this regions turns to be. Additional polychaete taxonomic studies have significantly changed interpretation of biogeographic patterns and biotic provinces based on earlier, less complete data. Faunal affinities with other regions also should be cautiously interpreted as might be biased. This review suggests than, despite the significant work already carried out by foreign researchers, the true nature and magnitude of polychaete diversity in Chile is yet to be assessed. Unfortunately, only a small number of researchers are working on ROZBACZYLO AND SIMONETTI: CHILEAN BENTHIC POLYCHAETES 369 the taxonomy and ecology of polychaetes in Chile. As with systematists working on other taxa, human resources working on polychaetes are scarce and ill-supported to complete the survey of the Chilean fauna. The intensity and extent of Chilean coast being altered by anthropogenic action makes the need to increase the recruitment and training of young scientists to study polychaetes even more urgent.

ACKNOWLEDGMENTS

The authors are indebted to A. Angel and F. Labra for the kind help with preparation of English manuscript; M. Bobadilla helped with the preparation of final figures. This paper was prepared during a research project on taxonomy and ecology of marine benthic polychaetes from northern Chile, as part of the “Programa Sectorial Biomas” (1996–1997) supported by Fondecyt No.5960001. Very special thanks are due to J. Vásquez and P. Ojeda for inviting NR to participate in that scientific program and to J. Correa, Head of Department of Ecology, Facultad de Ciencias Biológicas and to the Vicerrectoría Académica, P. Universidad Católica de Chile, for the provision of funds that allowed NR to attend the 6th International Polychaete Conference at Curitiba, Brazil. We spe- cially thank the careful review from two anonymous reviewers and D. Reish that substantially im- proved this paper.

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ADDRESSES: (N.R.) Departamento de Ecología, Facultad de Ciencias Biológicas, Pontificia Universidad Católica de Chile, Casilla 114-D, Santiago, Chile. E-mail: ; (J.S.) Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile. E-mail: .