Polychaeta: Pectinariidae): Taxonomic Characterization, New Distributional Records and Ecological Notes from the Chilean Coast

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Pectinaria chilensis Nilsson, 1928 (polychaeta: Pectinariidae): Taxonomic characterization, new distributional records and ecological notes from the Chilean Coast

Article in Interciencia · October 2004

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The user has requested enhancement of the downloaded ﬁle. Pectinaria chilensis NILSSON, 1928 (POLYCHAETA: PECTINARIIDAE): TAXONOMIC CHARACTERIZATION, NEW DISTRIBUTIONAL RECORDS AND ECOLOGICAL NOTES FROM THE CHILEAN COAST

Rodrigo A. Moreno, Nicolás Rozbaczylo, Roger D. Sepúlveda, Franklin D. Carrasco and Raúl Soto

SUMMARY

The taxonomy, geographical distribution, and ecological as- characterization, illustrations and a list of new localities in the pects of Pectinaria chilensis are reviewed. Specimens were col- geographical distribution of this species are presented. Ecologi- lected in sublittoral soft bottoms, between 45 and 93m depth, at cal information (bathymetry, sediment type, total organic matter four localities in northern Chile. These specimens are compared content and dissolved O2 concentration of sea water above the with individuals collected from 17m depth at Coronel, Gulf of bottom) is included, as well as a key for differentiating the two Arauco, Chile, the type locality of the species. A taxonomic Pectinariidae species registered in Chile to date.

RESUMEN

Se revisa Pectinaria chilensis de la costa de Chile a partir caracterización taxonómica, ilustraciones, y nuevas localidades de ejemplares recolectados en fondos blandos sublitorales, entre en la distribución geográfica de la especie. Se provee infor- 45 y 93m de profundidad, en cuatro localidades del norte de mación del hábitat (batimetría, tipo de sedimento, contenido de

Chile. Estos especímenes fueron comparados con ejemplares materia orgánica total y O2 disuelto de fondo) y se incluye una recolectados a 17m de profundidad en Coronel, Golfo de clave para diferenciar las dos especies de Pectinariidae regis- Arauco, Chile, localidad tipo de la especie. Se incluye una tradas en Chile.

Introduction morphological characteristics gren and Pectinaria Savigny, accepted a total of 46 nomi- (Rouse and Pleijel, 2001). including within the latter the nal species as valid within the The family Pectinariidae Pectinariidae form a clade subgenera Lagis, Amphictene, two genera. The same author Quatrefages, 1865 includes with the Ampharetidae and and Pectinaria. Fauchald considered Lagis, Amphictene, the tubiculous polychaetes Alvinellidae and belong to (1977) considered the preced- and Cistenides to be subgen- known as “ice cream cone the Terebellida (Rouse and ing three subgenera, together era of Pectinaria. Hutchings worms” or “trumpet worms” Fauchald, 1997). Evidence of with Cistenides Malmgren, as (2000) recognized about 51 due to their characteristic monophyly in this group has separate genera, thus recog- described species and sug- cone-shaped tubes. The tubes been based on their conical nizing five genera in the fam- gested the need for a review are constructed of sand tubes of unique shape and ily (Petta, Lagis, Amphictene, of the family, with better defi- grains and/or shell fragments construction, the presence of Cistenides and Pectinaria). nitions of the genera, and an and foraminifera, in a one- a cephalic veil, and the de- Pettibone (1982) recognized adequate evaluation of their or two-layered arrangement velopment of a flattened only three genera (Amphicte- status. (Day, 1967; Wolf, 1984; scaphe, with spine-like cha- ne, Petta, and Cistena), con- Recently, Hutchings and Rouse and Pleijel, 2001). etae (Fauchald and Rouse, sidering Pectinaria, Cisteni- Peart (2002), recognized 53 These polychaetes are found 1997). des, and Lagis as synonyms species in 5 genera, previ- throughout the world, though According to Wolf (1984) of Cistena. Holthe (1986) in ously indicated by Fauchald they are rarely found in large the number of genera within his study of the family recog- (1977), based on a review of numbers. They form a group Pectinariidae has been confus- nized only two genera, Pecti- the Pectinariidae in Austra- that is anatomically complex, ing. Day (1967) recognized naria and Petta, and four sub- lian waters. For the genus yet relatively uniform in their only two genera: Petta Malm- genera within Pectinaria and Pectinaria there are 22 rec-

KEYWORDS / Benthos / Northern Chile / Pectinaria chilensis / Polychaeta / Southeastern Pacific / Received: 01/13/2004. Modified: 09/08/2004. Accepted: 09/13/2004.

Rodrigo A. Moreno. Marine Bi- Nicolás Rozbaczylo. Biology Pro- lica de la Santísima Con- UdeC, Chile. Professor, UdeC. ologist, Universidad Arturo fessor, Universidad de Chile cepción, Chile. M.S. in Zool- e-mail: [email protected] Prat, Chile. M.S. in Zoology, (UCh). Professor, Department of ogy, UdeC, Chile. Ph.D. Stu- Raúl Soto. Biology Professor, Universidad de Concepción Ecology, Pontificia Universidad dent, Institute of Ecology and Universidad de Chile (UCh). (UdeC), Chile. Researcher, Católica de Chile. Address: P.O. Evolution. Universidad Aus- M.S. in Zoology, UdeC, Chile. Chilean Biodiversity Program, Box 114-D, Santiago, Chile. e- tral de Chile. e-mail: Professor, Universidad Arturo Universidad Arturo Prat. e-mail: mail: [email protected] [email protected] Prat, Chile. e-mail: [email protected] Roger D. Sepúlveda. Marine Franklin D. Carrasco. Biologist, [email protected] Biologist, Universidad Cató- M.S. and Ph.D. in Zoology,

590 0378-1844/04/10/590-04 $ 3.00/0 OCT 2004, VOL. 29 Nº 10 RESUMO

Revisa-se Pectinaria chilensis da costa do Chile a partir caracterização taxonômica, ilustrações, e novas localidades de exemplares recolhidos em fundos brandos sub-litorais, en- na distribuição geográfica da espécie. Se fornece informação tre 45 e 93m de profundidade, em quatro localidades ao nor- do hábitat (batimetria, tipo de sedimento, conteúdo de maté- te do Chile. Estes espécimes foram comparados com exempla- ria orgânica total e O2 dissolvido de fundo) e se inclui uma res recolhidos a 17m de profundidade em Coronel, Golfo de clave para diferenciar as duas espécies de Pectinariidae re- Arauco, Chile, localidade padrão da espécie. Se inclui uma gistradas no Chile.

ognized species around the Universidad Católica de world, including 2 new spe- Chile, Santiago (SSUC), cata- cies (P. dodeka and P. logued under the following kanabinos) for the Australian numbers: SSUC 6980 to fauna (Hutchings and Peart, 6983. 2002). On the Southeastern Pacific Results coast of Chile, Pectinariidae is represented by two genera: Pectinaria chilensis Nilsson, Pectinaria and Cistenides, 1928: 37-40, fig. 11; Hart- with one species each: Pecti- man, 1941: 333 pl. 50 figs. naria chilensis Nilsson, 1928 12, 15, pl. 51, fig. 19. and Cistenides ehlersi Hessle, Material examined: 92 1917 (Rozbaczylo, 1985). The specimens. Arica (n=6), Con- aim of the present study was tinental shelf off Iquique Bay to review P. chilensis from (n=7); Patillos (n=2); Punta the Chilean coast, to conduct Coloso (n=61); Coronel (Gulf a taxonomic characterization, of Arauco) (n=16). to illustrate morphological as- Description: Individual pects, and to present new lo- polychaetes reached up to calities in the geographical 44mm in length and 6mm in distribution of the species. We width at the opercular level. present data on bathymetry, The operculum (Figure 1a) is sediment type, total organic a thick muscular plate which matter content, and dissolved forms the anterodorsal surface

O2 in the water above the of the (fused) prostomium bottom at the sample loca- and peristomium. There are tions. We also include a taxo- Figure 1. Pectinaria chilensis. a: anterior end in dorsal view; b: two anteroventral groups of nomic key to differentiate the anterior end in ventral view; c: scaphe in dorsal view; d: limbate stout, flattened golden paleae, two species of Pectinariidae setae from notopodium; e: neuropodial uncinus in lateral view. which are slightly dorsally recorded in Chile to date. (Scales: a-c= 1mm; d= 0.1mm; e= 0.01mm). curved and arranged in two oblique, overlapping rows, Materials and Methods with seven to eight paleae per 2002 in Coronel, Gulf of ment characteristics/type. To- row. The operculum is sur- Specimens of P. chilensis Arauco (36º42'S), at 17m tal organic matter (TOM) rounded dorsally and ventrally were collected as a part of depth. was estimated by weight loss by a raised, smooth, opercular three separate, larger studies: Four sediment subsamples after ignition of dried material border. Ventrally, below the 1) The Benthic Macrofauna were obtained at each locality in a furnace for 2h at 550ºC paleae is a thin membrane Diversity of Northern Chile using a 0.1m2 Van Veen grab. (Buchanan, 1971). Dissolved called the cephalic veil, which

Project, with samplings con- Three of these subsamples O2 content of seawater from is bounded by about 30 to 32 ducted between Sept 1998 and were used to obtain polycha- each sample station was de- thin papillae, which form a May 2003 at three localities, etes, and the fourth was used termined by the Winkler titra- hood in front of the mouth. including Arica (18º20'S), the to determine sediment charac- tion method (Strickland and Approximately 32 oral ten- continental shelf off Iquique teristics. Polychaetes were Parsons, 1972), for samples tacles of different size form Bay (20º11'S) and Patillos separated from the sediment collected in Niskin bottles two groups, arising at the (20º45'S), between 45 and using an 0.5mm mesh sieve. 0.5m over the bottom. point where the cephalic 93m depth; 2) The Minera Specimens were fixed in 10% Illustrations of the species membrane attaches at the Escondida Ltda. Environmental seawater-formalin solution. were prepared using a camera mouth (Figure 1b). Each indi- Project, with samplings con- Specimens of P. chilensis lucida mounted on a Wild M- vidual has two pairs of ten- ducted from 1990 to 2002 in were quantified, measured and 5 stereo microscope. Identi- tacular cirri: the anterior pair Punta Coloso, Antofagasta transferred to a 70% ethanol fied specimens were deposited is lateral and slightly ventral (23º45'S) at 60m depth; and 3) solution containing 8% glyc- in the Colección de Flora y to the paleae and the posterior a regional control program for erine for long-term preserva- Fauna Profesor Patricio Sán- pair is located laterally to the wastewater treatment, with tion. Folk (1974) was fol- chez, located in the Ecology mouth and connected by a samplings conducted in Nov lowed for evaluations of sedi- Department at the Pontificia long transverse ventral ridge.

OCT 2004, VOL. 29 Nº 10 591 Two achaetous segments fol- TABLE I low this ridge, each bearing a ENVIRONMENTAL PARAMETERS AT EACH SAMPLED LOCALITY ALONG lateral pair of lamellate THE CHILEAN COAST WHERE P. chilensis WAS PRESENT, BETWEEN 1998 AND 2003 branchiae similar in size (al- Locality Sediment Total organic Dissolved O Depth (m) though the first pair are 2 type matter (%) content (ml·l-1) slightly longer than the sec- ond pair). The number of se- Arica Fine sands 5.92 0.94 58 tigerous segments is always Continental shelf off Iquique Bay Mud 9.41 0.37 93 16; setigers 1-3 only have Patillos Fine sands 5.89 0.91 45 notosetae; setigers 4-15 have Punta Coloso Fine sands 5.24 0.5 60 uncini and notosetae, and Coronel (Gulf of Arauco) Mud * * 17 setiger 16 only notosetae; the * Not available. last segment bears a reduced parapodium which lacks setae. The notosetae (Figure 1d) are our samples compared with end projects slightly above uted from Aysen (43ºS) to the simple (limbate), and similar eight or nine cited in the the sediment surface so that Straits of Magellan (56ºS; along the entire length of the original description, and nine the opercular paleae can be Rozbaczylo, 1985), restricted body. The uncini (Figure 1e) or ten indicated by Hartman used to excavate within the to cold temperate waters of show a basal trough-like for the Peruvian specimens). sediment (Wolf, 1984). the Magellanic Province structure and two rows of Nilsson (1928) suggested that Specimens collected from (41ºS-56ºS; Ekman, 1953). both major and minor teeth. the cephalic membrane pos- the continental shelf off On the continental shelf off There are approximately 15 sesses about 60 papillae, Iquique Bay coexist with Iquique Bay, P. chilensis co- acicular scaphal setae on each while in our specimens the mats of the giant filamentous exists with Ampelisca arau- side of the anterodorsal mar- number of papillae is only sulfur bacteria Thioploca spp. cana and Thioploca spp. gin of the scaphe, of which around 30; the latter number and the deposit-feeding am- mats. This relationship may five or six of the inner setae is, however, comparable with phipod, Ampelisca araucana be the result of the sediment are smaller. There is an ap- the number observed by Hart- Gallardo. These species have and hydrographic characteris- proximately circular anal flap man (1941) for the Peruvian been observed at high densi- tics recorded for the bottom on the posterior-dorsal margin specimens, which had 30 to ties in environments of the of this locality, which has of the scaphe (Figure 1c); the 60 filiform papillae on the Peru-Chile Undercurrent muddy sediments with a high anal flap has a crenulate mar- free edge. These differences (PCU), i.e. the Gulf of Arau- content of total organic matter gin, and its upper side has an are considered to be intraspe- co, Chile, where sediments (>9.0%), and very low con- anal cirrus, which is con- cific variation. are often hypoxic (Carrasco centrations of dissolved O2 spicuous and located approxi- Distribution and bathym- and Gallardo, 1983). (0.37ml·l-1) at 93m depth, as- mately at the center. etry: Records of P. chilensis Observations: P. chilensis from new localities allow the Nilsson, 1928 was originally extension of its geographic Key to the species of Pectinariidae recorded from Chile described from a single speci- range in Chile, to over 18º of men collected at Coronel, latitude, reaching from 36º42' 1a Body with 16 setigerous of which 12 are uncinigerous; Gulf of Arauco (VIII Region, S (i.e. the type locality) to 7-10 pairs of opercular paleae; approximately 13-15 pairs Chile; Rozbaczylo, 1985). 18º20'S (Arica). Thus, the of scaphal hooks; uncini with two rows of teeth.... Hartman (1941) reported the geographic distribution of the ...... Pectinaria chilensis presence of the species at species ranges from Indepen- Independencia Bay, Peru from dencia Bay, Peru, to the Gulf 1b Body with 17 setigerous of which 13 are uncinigerous; 27 specimens collected be- of Arauco, Chile, reaching a 8-13 pairs of opercular paleae; about 10 pairs of scaphal tween the shore and 38m maximum depth of 93m. hooks; uncini with a single row of teeth.... depth. P. chilensis has been Physical and chemical pa- ...... Cistenides ehlersi recorded in the coast of Chile rameters: P. chilensis was by Oyarzún et al. (1987) at found associated principally Talcahuano (36º38'S), and by with fine sandy and muddy Discussion sociated with the Equatorial Carrasco et al. (1988) and sediments, with organic mat- Subsurface Waters mass Carrasco and Gallardo (1994), ter content greater than 5%, The results from our study (ESSW) of the Peru-Chile and Carrasco (1996) at and dissolved O2 concentra- suggest that Pectinaria chilen- Undercurrent. These environ- Concepcion Bay (36º40'S), to- tion of less than 1.0ml·l-1 in sis may have a continuous mental conditions are favor- gether with some ecological the overlying water above the distribution in warm temper- able for the development and data on the species. bottom (Table I). ate waters on the southeastern coexistence of populations of The samples reviewed in Ecological notes: Pectinari- Pacific coast, from 14°15’S A. araucana (see Carrasco the present study, from ids are sedentary free-living (Independencia Bay, Perú) to and Gallardo, 1983; Carrasco Coronel and northern Chile, tubiculous polychaetes which 36°42’S (Gulf of Arauco, and Arcos, 1984) and Thio- agree well with the original typically inhabit muddy, or Chile), within the Peruvian ploca spp. mats (Gallardo, description made by Nilsson fine sandy bottoms, and are Province (3-4ºS-41ºS; Ekman, 1977; Fossing et al., 1995), (1928), as well as with selective subsurface deposit- 1953), with a lack of records mainly due to the protection Hartman’s (1941) description, feeders. The anterior end of in intermediate areas between generated by this hypoxic except for the number of the tube is oriented with the Antofagasta and the Gulf of barrier against predators and opercular paleae (seven or mouth facing downwards, Arauco. On the other hand, competing species, which are eight paleae on each side in while the posterior (narrow) Cistenides ehlersi is distrib- unable to enter waters with

592 OCT 2004, VOL. 29 Nº 10 severe hypoxia (<0.5ml·l-1; strong enzymatic activity for Central Chile. Int. Revue ges. Hartman O (1941) Polychaetous an- Díaz and Rosenberg, 1995). a pyruvate oxidoreductase, the Hydrobiol. 73: 441-455. nelids. Pectinariidae. With a re- Carrasco FD, Gallardo VA (1994) view of all species from the On the other hand, the ab- alanopine dehydrogenase western hemisphere. Allan sence of A. araucana and (ALPDH). The presence of Diversidad, distribución y abun- dancia del macrobentos sublito- Hancock Pacific Exped., 7, Part Thioploca spp. at other locali- more than one pyruvate oxi- ral, y observaciones sobre la IV. pp. 325-345. ties analyzed could be ex- doreductase is typical for spe- dinámica temporal de corto Holthe T (1986) Evolution, systemat- plained by the presence of cies that have to cope with término de los sedimentos de la ics, and distribution of the Poly- moderate hypoxia (2-0.5ml·l-1; both functional and environ- Bahía Concepción, Chile. Gaya- chaeta Terebellomorpha, with a Díaz and Rosenberg, 1995), mental hypoxia (Gäde and na Oceanol. 2: 49-68. catalogue of the taxa and a bib- liography. Gunneria 55: 1-236. which does not provide an ef- Grieshaber, 1986). González Carrasco FD (1996) La macroinfauna bentónica de la Bahía Concep- Hutchings P (2000) Family Pectina- fective barrier against preda- and Quiñones (2000) con- ción, Chile: alta dominancia riidae. In Beesley PL, Ross tors and competitors, and clude that P. chilensis presents ecológica en el sublitoral some- GJB, Glasby CJ (Eds.) Polycha- could eliminate or notably re- a low capacity for coping ro frente a Lirquén. Gayana etes & Allies: The Southern Syn- duce populations of A. arau- with hypoxia. However, our Oceanol. 4: 1-12. thesis. Fauna of Australia. Vol. cana and Thioploca spp., as findings of P. chilensis in hy- Carrasco FD, Gallardo VA, Baltazar 4A. Polychaeta, Myzostomida, Pogonophora, Echiura, Sipun- well as the predominance of poxic zones do not agree with M (1999) The structure of the benthic macrofauna collected cula. CSIRO. Melbourne, Aus- fine sandy sediments and low the results of González and across a transect at the central tralia. pp. 219-222. organic matter content. These Quiñones (2000) for this spe- Chile shelf and relationships Hutchings P, Peart R (2002) A re- conditions do not affect P. cies. Therefore, it is necessary with giant sulfur bacteria view of the genera of Pectina- chilensis, since this species to analyze specimens from a Thioploca spp. mats. Cah. Biol. riidae (Polychaeta) together with inhabits sediments ranging greater number of localities Mar. 40: 195-202. a description of the Australian from fine sand to mud. Fur- along the Chilean coast to es- Day JH (1967) A monograph on the fauna. Rec. Austr. Mus. 54: 99- polychaeta of Southern Africa. 127. thermore, this species has an tablish whether P. chilensis Brit. Mus. Natur. Hist. Publ. Maier S, Gallardo V (1984) Nutri- opercular paleae, which can presents a low capacity to 656: 30-878. tional characteristics of two ma- effectively close the entrance adapt to the hypoxic condi- Díaz R, Rosenberg R (1995) Marine rine thioplocas determined by to the tube and may protect tions in the Humboldt Current benthic hypoxia: a review of its autoradiography. Arch. Micro- the worm from being eaten System. ecological effects and the biol. 139: 218-220. by predators (Hutchings, behavioural responses of benthic Nilsson D (1928) Neue und alte macrofauna. Ocean. Mar. Biol. Amphicteniden. Göteborgs Ve- 2000). ACKNOWLEDGEMENTS Ann. Rev. 33: 245-303. The mixotrophic condition tenskSamh. Handl. 33: 1-96. Ekman S (1953) Zoogeography of Oyarzún C, Carrasco FD, Gallardo of Thioploca spp. based on The authors thank Minera the sea. Sidgwick and Jackson. VA (1987) Some characteristics the oxidation of hydrogen sul- Escondida Ltda. for permit- London, UK. 417 pp. of macrobenthic fauna from the fide in the presence of low O2 ting the use and citation of Fauchald K (1977) The polychaete organic-enriched sediments at concentrations (Maier and biological material from the worms: Definitions and keys to Talcahuano, Chile. Cah. Biol. Gallardo, 1984; Fossing et al., Environmental Monitoring the orders, families and genera. Mar. 28: 429-446. Nat. Hist. Mus. Los Angeles 1995) plays an important role Program for Punta Coloso County. Sci. Ser. 28: 1-190. Pettibone MH (1982) Annelida. In in the biogeochemical control (Antofagasta, Chile), José Mi- Parker SP (Ed.) Synopsis and of the sediments (Schulz et guel Fariña, Guillermo Guz- Fauchald K, Rouse G (1997) Poly- Classification of Living Organ- chaete systematics: past and isms. McGraw-Hill. New York, al., 2000), detoxifying the up- mán, and Paula Neill for present. Zool. Scr. 26: 71-138. EEUU. Vol. 2. 1-43. per sediment strata of H S, valuable suggestions and criti- 2 Folk R (1974) Petrology of sedimen- Rouse GW, Fauchald K (1997) Cla- and thus generating favorable cal comments. Roger D. Se- tary rocks. Hemphill. Austin, distics and polychaetes. Zool. conditions for the colonization púlveda was supported by a TX, EEUU. 170 pp. Scr. 26: 139-204. of aerobic organisms (Gallar- CONICYT-Chile Doctoral Fossing H, Gallardo VA, Jørgensen Rouse GW, Pleijel F (2001) Polycha- do, quoted in Carrasco et al., Fellowship. BB, Hüttel M, Nielsen LP, etes. Oxford University Press. 1999), which are able to tol- Schulz H, Canfield DE, Foster New York, EEUU. 354 pp. S, Glud RN, Gundersen JK, erate low O2 concentrations REFERENCES Küver J, Ramsing NB, Teske A, Rozbaczylo N (1985) Los anélidos and are able to access the Thamdrup B, Ulloa O (1995) poliquetos de Chile. 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