Polychaetes of the Genus Ampharete (Polychaeta: Ampharetidae) Collected in Icelandic Waters During the BIOICE Project
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Helgol Mar Res (2012) 66:331–344 DOI 10.1007/s10152-011-0274-z ORIGINAL ARTICLE Polychaetes of the genus Ampharete (Polychaeta: Ampharetidae) collected in Icelandic waters during the BIOICE project Julio Parapar · Gudmundur V. Helgason · Igor Jirkov · Juan Moreira Received: 15 June 2010 / Revised: 14 August 2011 / Accepted: 2 September 2011 / Published online: 22 September 2011 © Springer-Verlag and AWI 2011 Abstract Based on material collected during the BIOICE the size, shape and disposition of the paleae, a special kind project oV Iceland, the taxonomy and distribution of eight of chaetae located on the Wrst body chaetiger. Several body species of polychaetous annelids belonging to the genus characters of high taxonomic relevance in Ampharetidae Ampharete (Polychaeta: Ampharetidae) are reviewed. Five are reviewed under the SEM. The distribution of each spe- of these species were previously reported in the area: cies oV the coast of Iceland and a key to all species Ampharete acutifrons, Ampharete borealis, Ampharete recorded in European Boreo-Arctic waters is provided. Wnmarchica, Ampharete octocirrata and Ampharete petersenae; two are new to Icelandic waters: Ampharete Keywords Polychaeta · Ampharetidae · Ampharete · baltica and Ampharete lindstroemi; and one is new to sci- Iceland · New species · BIOICE project ence: Ampharete villenai sp. nov. A. petersenae was found for the Wrst time after the original description and is rede- scribed. A. villenai sp. nov. is primarily characterized by Introduction The genus Ampharete Malmgren, 1866, with about 30 described species (Jirkov 1994) is the most speciose genus Communicated by Peter Funch. in the family Ampharetidae Malmgren, 1866. This number would exceed 40 species if Asabellides and Sabellides are J. Parapar (&) Departamento de Bioloxía Animal, Bioloxía Vexetal e Ecoloxía, considered synonymous. It is composed of surface deposit- Facultade de Ciencias, Universidade da Coruña, feeding polychaetes that live on soft bottoms from the inter- Alejandro da Sota 1, 15008 A Coruña, Spain tidal down to abyssal depths (Salazar-Vallejo 1996). Since e-mail: [email protected] its creation by Malmgren (1866), the genus has been well W G. V. Helgason characterized, not undergoing signi cant changes in its Institute of Biology, University of Iceland, diagnosis after the classical reviews by Hessle (1917), Day Sturlugata 7, 101 Reykjavík, Iceland (1964) and Holthe (1986a). Thus, the main diagnostic char- acters used to be included are the presence of papillose buc- I. Jirkov Department of Hydrobiology, Biological Faculty, cal tentacles, four pairs of branchiae, presence of paleae Moscow State University, Moscow, Russia (a special kind of forward-pointing chaetae, located on the third body segment), 14 thoracic chaetigers with notosetae J. Moreira (15 when considering paleae) and 12 with neurosetae and Estación de Bioloxía Mariña da Graña, Universidade de Santiago de Compostela, 12–26 abdominal segments (Reuscher et al. 2009). Never- Rúa da Ribeira 1, A Graña, 15590 Ferrol, Spain theless, Jirkov (1994, 2001, 2009) questions the taxonomic validity of the presence/absence of paleae. He showed that J. Moreira within some Ampharetinae genera traditionally considered Departamento de Biología (Zoología), Facultad de Ciencias, Universidad Autónoma de Madrid, Cantoblanco, as provided with paleae, the form and number of this kind 28049 Madrid, Spain of chaetae can vary signiWcantly, suggesting that their 123 332 Helgol Mar Res (2012) 66:331–344 presence/absence should not be used for generic deWnition. Hessle, 1917 are new for the studied area. Ampharete ville- Thus, in Ampharete, a variation from huge well-developed nai sp. nov. is new to science and described herein. paleae (A. longipaleolata Ushakov, 1950) to completely Ampharete goesi Malmgren, 1866, a mostly Arctic species absent (A. sibirica (Wiren, 1883) and A. petersenae Jirkov, (type locality Spitsbergen) but also reported from the Okhotsk 1997) is noted. Also, a species devoid of paleae was also Sea and Japan Sea (NW PaciWc), was the only previously recently described within the genus Anobothrus Levinsen, reported species that was not found in this study. Further- 1884, i.e. Anobothrus apaleatus Reuscher, Fiege & Wehe, more, two other species known from North Atlantic and the 2009. Moreover, Jirkov (1994, 2009) reported that the fac- Arctic, respectively, were not present in the BIOICE ultative absence of paleae in some specimens (ca. 5%) of a samples, i.e. Ampharete falcata Eliason, 1955 (low-boreal population of Ampharete saphronovae Jirkov 1994, a spe- species; type locality Swedish west coast) and Ampharete cies described with paleal chaetae of almost the same size vega (Wirén, 1883) (shallow-water Arctic euryhaline species, as the thoracic notochaetae, concluding that even for spe- type locality Bering Sea). cies diagnosis this character should be used with care. The use of the scanning electronic microscope (SEM) According to this, we consider the genus Asabellides has been proved a useful tool in the study of external struc- Annenkova, 1929 as a junior synonym of Ampharete tures of polychaetes in general and those of the clade Tere- because both genera were only distinguished by the bellomorpha in particular (Hutchings and Peart 2000; absence of paleae in the former. Furthermore, we also fol- Jouin-Toulmond and Hourdez 2006; Parapar and Moreira low Jirkov (2001, 2009) in considering the presence of 13 2008a, b). In this work, the SEM has been used to elucidate thoracic chaetigers (genus Sabellides Milne-Edwards, 1838 some anatomical characteristics, which are diYcult to in Malmgren 1866) instead of 14, as character states that do observe under the compound microscope, such as the pres- not justify the discrimination of genera. Consequently, the ence and distribution of body ciliature and the shape of genus Ampharete sensu Jirkov (1994, 1997, 2001, 2009) paleae, notochaetae and uncini. would include ampharetids with or without paleae, with The BIOICE sampling area covers a depth range from 20 four pairs of smooth branchiae, papillose buccal tentacles, to 3,500 m on both sides of the Greenland-Iceland-Faeroe 13–14 thoracic chaetigers (excluding paleae) provided with Ridge (GIF Ridge), which is generally less than 500 m deep notochetae and 11–12 thoracic uncinigers. and marks the boundary between the relatively warm North Since Holthe (1986a) reported 7 species of Ampharete in Atlantic Ocean and the much colder Nordic seas of the Arctic North Atlantic waters, only Jirkov (2001), within a larger Ocean. Other papers dealing with BIOICE polychaetes were study on Arctic Polychaeta, reviewed the presence of this those of Petersen (2000) on Fauveliopsidae, Kirkegaard genus in northern European waters. The scattered informa- (2001) on Glyceridae and Goniadidae, SanWlippo (2001) on tion on this genus in Icelandic waters was Wrst compiled by Serpulidae, Sigvaldadóttir (2002) on Spionidae, Chambers Wesenberg-Lund (1951), who mentions Ampharete acuti- and Woodham (2003) on Cirratulidae, Parapar (2003; 2006) frons (Grube, 1860), Ampharete goesi Malmgren, 1866 (as on Oweniidae, Parapar et al. (2011c) on Opheliidae, Parapar A. goësi), Ampharete borealis (Sars, 1856) (as Sabellides) et al. (2011b) on the Ampharetidae genus Amphicteis and and Ampharete octocirrata (Sars, 1835) (as Sabella). Later, Parapar et al. (2011a) on the genus Terebellides. Gardarsson (1973) reports the presence of Ampharete Wnmarchica (Sars, 1866) and recently Jirkov (1997) described a new species, Ampharete petersenae Jirkov, Materials and methods 1997, from the east coast. The BIOICE (Benthic Invertebrates of Icelandic Waters) This study is based on material collected in the framework of expeditions were part of an international collaborative pro- the BIOICE project. A total of 1371 specimens of the genus gramme that started in 1992 to conduct a thorough survey Ampharete were collected from 224 bottom samples (Fig. 1a) of the marine benthic fauna living in the 200 mile exclusive along the continental shelf and slope of Iceland. Several bot- economic zone (EEZ) of Iceland. Eight species belonging tom samplers (modiWed Rothlisberg–Pearcy epibenthic sledge, to the genus Ampharete sensu Jirkov (1994) were identiWed detritus sledge, triangle dredge, Agassiz trawl and Shipek grab) from material collected in those expeditions. Five species were used during the BIOICE cruises on diVerent research ves- have previously been reported in Icelandic waters, namely sels (Bjarni Sæmundsson, Håkon Mosby and Magnus Heina- Ampharete acutifrons, Ampharete borealis, Ampharete son). Sampling coordinates and abiotic features of the BIOICE Wnmarchica, Ampharete octocirrata and Ampharete pet- samples are available in the home page of the Icelandic Insti- ersenae; the latter is reported for the Wrst time after the tute of Natural History (http://www.utgafa.ni.is/greinar/ original description and is redescribed. Ampharete baltica BIOICE_station_list_91-04_Paper_A2.pdf) and also from the Eliason, 1955 and Ampharete lindstroemi Malmgren in authors (GVH) upon request. 123 Helgol Mar Res (2012) 66:331–344 333 Specimens were Wxed in 10% formalin buVered with borax and preserved in 70% ethanol. Animals were picked from samples by the staV of the Sandgerdi Marine Centre (SMC) and then identiWed to species level by the authors. Most of the material examined, including the type series of the new species, was deposited in the collections of the Ice- landic Museum of Natural History (IMNH, Reykjavik); some paratypes