Polychaetes of Greece: an updated and annotated checklist

Sarah Faulwetter; Nomiki Simboura; Nikolaos Katsiaras; Giorgos Chatzigeorgiou; Christos Arvanitidis

doi:10.3897/BDJ.5.e20997

Biodiversity Data Journal : Taxonomic Paper

Taxonomic Paper

Polychaetes of Greece: an updated and annotated checklist

Sarah Faulwetter^‡, Nomiki Simboura^§, Nikolaos Katsiaras^§,|, Giorgos Chatzigeorgiou^¶, Christos Arvanitidis^¶

‡ University of Patras, Department of Zoology, Section of Marine Biology, Patras, Greece

§ Institute of Oceanography, Hellenic Centre for Marine Research, Anavyssos, Greece

| Department of Marine Sciences, University of the Aegean, Mytilini, Greece

¶ Institute of Marine Biology, Biotechnology and Aquaculture, Hellenic Centre for Marine Research, Heraklion Crete, Greece

Corresponding author: Sarah Faulwetter (sarahfaulwetter@gmail.com)

Academic editor: Vasilis Gerovasileiou

Received: 14 Sep 2017 | Accepted: 13 Dec 2017 | Published: 22 Dec 2017

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Faulwetter S, Simboura N, Katsiaras N, Chatzigeorgiou G, Arvanitidis C (2017) Polychaetes of Greece: an updated and annotated checklist. Biodiversity Data Journal 5: e20997. https://doi.org/10.3897/BDJ.5.e20997

Abstract

Background

The last annotated checklist of marine polychaetes in Greece was published in 2001. Since then, global taxonomic progress, combined with many new species records for Greece, required a thorough review of the taxonomic, nomenclatural and biogeographic status of the national species list. This checklist revises the status of all extant polychaete species reported from the Greek Exclusive Economic Zone since 1832. The work was undertaken as part of the efforts on compiling a national species inventory (Greek Taxon Information System initiative) in the framework of the LifeWatchGreece Research Infrastructure.

New information

This checklist comprises an updated and annotated inventory of polychaete species in Greek waters, compiled from literature reports, online databases, museum collections and unpublished datasets. The list provides information on 836 species-level taxa from Greece, of which 142 are considered questionable. An additional 84 species reported in the past are currently considered absent from Greece; reasons for the exclusion of each species are given. Fourteen species are reported here for the first time from Greek waters. At least 52 species in the present list constitute in fact a complex of cryptic or pseudo-cryptic species. Forty-seven species are considered non-native to the area. In addition to the species-level taxa reported in this checklist, eleven genera have been recorded from Greece with no representatives identified to species level. One replacement name is introduced. For each species, a comprehensive bibliographic list of occurrence records in Greece and the synonyms used in these publications are provided as supplementary material. Where necessary, the taxonomic, nomenclatural or biogeographic status is discussed. Finally, the findings are discussed in the wider context of Mediterranean polychaete biogeography, taxonomic practice and worldwide research progress.

Keywords

Eastern Mediterranean, Polychaeta, Annelida, inventory, biodiversity, new records, replacement name

Introduction

Polychaete research in Greece

The history of polychaete studies in Greece can be traced back to classical antiquity. Aristotle, although he tended to neglect polychaetes in his works on natural history (Voultsiadou et al. 2017) gave the first description of a “scolopendra thalassia” (referring to Hermodice carunculata Pallas, 1776): “There are also marine scolopendras, similar to the terrestrial ones, but smaller in size; they live on the rocky substrates. Their colour is richer red and they bear more and much slender feet than the terrestrial scolopendras. They do not occur in great depths, as do the marine snakes” (History of Animals, 4^th century B.C.). Modern systematic polychaete research in the country started in the 19^th century (e.g. Brullé 1832, Forbes 1842, von Marenzeller 1893). Since then, most research has been directed towards the study of macrobenthic coastal soft bottom polychaete communities in an ecological context. To date, the only taxonomic work studying exclusively planktonic polychaetes in Greece is by Wesenberg-Lund (1939a), based on samples taken during the Danish Oceanographic Expedition of the research vessel “Thor” from a few stations in Greek waters (Mavraki et al. 2016). Hard substrates, meiobenthic communities and deep sea habitats are understudied in Greece (e.g. Tselepides et al. 2004) while no terrestrial or fresh water polychaetes are known from the country. Specialised taxonomic or faunistic works from Greece are rare and the large body of ecological studies comprises the most important source for species records and thus for the creation of national checklists.

Rationale for the checklist

The first polychaete checklists covering the Greek waters were those of Arvanitidis (2000a) for the Aegean Sea (including the Turkish Aegean and the Sea of Marmara) under the broader framework of the entire Mediterranean inventory and of Simboura and Nicolaidou (2001) for all Greek seas; both checklists included only benthic species. In the framework of the European Union project PESI (Pan-European Species directories Infrastructures), Faulwetter (2010) combined the above lists and additional sources into an updated checklist, but this was simply a computer-based compilation of existing sources without further verification or annotation of the records.

After 2001, marine research, including benthic surveys (carried out by the Hellenic Centre for Marine Research (HCMR) and several universities) was intensified. Many new species records – often not only new for Greece, but also for the Mediterranean Sea – resulting from these studies, as well as the worldwide progress of taxonomic knowledge, required a thorough review of the taxonomic, nomenclatural and biogeographic status of many polychaete species recorded from Greece.

The construction of the Greek Taxonomic Information System (Bailly et al. 2016), which forms the taxonomic backbone of the LifeWatchGreece Research Infrastructure, provided the framework for the present study: the compilation of a comprehensive, annotated and updated checklist of the Greek polychaete fauna, expanding and updating former works (i.e. Arvanitidis 2000a, Simboura and Nicolaidou 2001). In addition, this publication complements recent checklists from other areas of the Mediterranean and Black Sea (e.g. Çınar 2005, Castelli et al. 2008, Ayari et al. 2009, Zaâbi et al. 2009, Martín and Gil 2010, Zaâbi et al. 2012, Çınar et al. 2014, Mikac 2015) and thus serves as an information resource for future polychaete studies in this biogeographic region.

Taxonomic scope

Several recent studies indicate the paraphyletic status of polychaetes (e.g. Rouse and Pleijel 2001, de Oliveira Almeida et al. 2003, Weigert and Bleidorn 2016) and the higher classification of Annelida has been in a constant flux over the last years (e.g. Zrzavý et al. 2001, Helm et al. 2012, Weigert and Bleidorn 2016). As the phylogenetic position of many groups within the Annelida is not yet resolved, we chose to restrict the "polychaetes" to their traditional definition in this checklist and include those clades covered by Rouse and Pleijel (2001). We anticipate this approach to be more helpful to the majority of users of this checklist than following the most recent phylogeny which would exclude certain "traditional" polychaete families (see Weigert and Bleidorn 2016).

Geographic scope and characteristics of the region

This checklist includes all polychaete taxa reported from the waters of the Greek Exclusive Economic Zone (EEZ) (as delimited by the Marine Regions database, see also Bailly et al. 2016). It does not cover a biogeographic area but constitutes an inventory of the national polychaete fauna. Thus, species reported from neighbouring countries but not (yet) from Greece are not included in this list, despite the fact that they are highly likely to occur in Greek water too.

Greece is located centrally in the Eastern Mediterranean Sea and thus lies in one of the world's marine biodiversity hotspots (Bianchi and Morri 2000a, Coll et al. 2010). Its coastline stretches over 18,000 km, showing a variety of coastal landforms, from cliffs and rocky shores to beaches, lagoons and river deltas and encompassing around 9,800 islands and rocky islets (Issaris and Salomidi 2012). The Greek EEZ comprises parts of three biogeographically distinct water bodies: the Aegean Sea in the east, the Ionian Sea in the west and the Libyan Sea in the south. The Aegean Sea exchanges waters in the north with the Black Sea through the Bosphorus Strait, the Marmara Sea and the Strait of Dardanelles. Extensive faulting and vertical tectonic movements have resulted in the fracturing of the Hellenic mountain chains and the creation of deeply incised embayments and semi-enclosed basins, whereas active underwater volcanoes and shallow-water hydrothermal vents can be found in several places in the Aegean Sea (Dando et al. 1999; see also Sini et al. (2017) for a detailed overview of the ecology of the Aegean Sea). The island of Crete forms the southern border of the Aegean and separates it from the Libyan Sea. The Libyan Sea is characterised by deep sea features such as trenches, extensive deep sea plateaus, hydrothermal vents, cold seeps and mud volcanoes. It is part of the Levantine basin and linked to the Red Sea via the Suez canal which forms the point of entry for Lessepsian migrant species. In the west, the Ionian Sea communicates with the Western Mediterranean and the Adriatic Sea. The Greek part of the Ionian Sea (East Ionian Sea) is characterised by the existence of the deep Hellenic Trench with depths exceeding 5,000 m – the deepest part of the Mediterranean Sea. Primary production in the Mediterranean decreases towards the eastern basin and thus the Greek waters are highly oligotrophic (SoHelME 2005), characterised by a limited availability of nutrients, primarily phosphorus (Thingstad et al. 2005). However, within this oligotrophic environment relatively eutrophic areas do exist (e.g. the North-West Aegean is highly productive in terms of nutrients and biological production compared to the Ionian, the South Aegean and the Levantine basin (e.g. Lykousis et al. 2002)). Knowledge on the polychaete fauna in each of these regional seas is unequally distributed, as the overwhelming majority of benthic research (around 80% of studies) has been performed in the Aegean Sea, whereas the deep sea habitats characterising the Libyan Sea are almost completely unexplored.

Materials and methods

Literature resources

Existing checklists on Greek polychaetes were combined, complemented, verified and annotated with information on species occurrences in Greece, using over 260 bibliographic and online resources, covering almost 200 years of research (1832–2017). A comprehensive list of all references from Greece for each species is contained in Suppl. material 2. These sources include publications from scientific journals, book chapters, theses, conference proceedings and biogeographic and museum collection databases. In addition, unpublished records generated in the framework of research projects conducted within the Hellenic Centre of Marine Research (HCMR) after 2001 were used to assess the occurrence status of many controversial and non-native species (Suppl. material 1). We deliberately chose to include grey literature and database records, as many of these have been used in secondary works and we prefer to draw attention to doubtful records rather than ignoring them. The additions to the previous checklists comprise polychaete records from 486 sampling stations which were compiled from a) literature and data sources from after 2001 and b) literature and data sources from before 2001 if these had not been covered by previous checklists (Fig. 1). The majority of these stations were monitored regularly or occasionally. Samples were mainly collected from shallow soft-bottom habitats, though a few samples from pelagic, deep-sea and hard substrate habitats are also included.

Figure 1.

Sampling stations from which polychaetes were recorded, compiled from literature sources and databases which where not included in previous checklists.

The final checklist was submitted to the World Register of Marine Species and the Pan-European Species directories Infrastructure to replace the latest version by Faulwetter (2010).

How to read this checklist

The nomenclature used here follows the World Register of Marine Species (WoRMS) and the most recent taxonomic literature. Taxa are arranged alphabetically in hierarchical order (genera and species within families); suprafamiliar classifications are omitted. The inclusion of species follows a conservative approach: species with a questionable presence in Greek waters are included in the list and clearly indicated as such; we excluded only species that are either considered taxonomically indeterminable according to the most recent taxonomic works or where Greek records were shown to be misidentifications. These are presented separately at the end of the manuscript.

For each species, one or more of the following types of information may be provided:

1. Nomenclature

If no information is provided in the "Nomenclature" field: Species has only been recorded under its currently accepted name.

If information is provided in the "Nomenclature" field: Not a comprehensive list of synonyms (for this, the reader is advised to consult WoRMS), but all names under which the species has been reported from Greece (respective literature resources for each name in Suppl. material 2). Subjective and objective synonyms (subjective: two different type specimens exist and original descriptions exist, synonymy is based on expert opinion; objective: different names exist as a result of assigning a species to a different genus or rank but are based on the same type specimen) as well as different rank assignments (e.g. subgeneric vs. generic level) are listed. Misspellings and spelling variations (e.g. species epithets ending in -a vs. -us) are excluded. Misidentifications (i.e. specific specimens which were in the past identified as belonging to another species but then re-examined) are also omitted to avoid confusion; instead, a remark is given in the Notes field and the record is annotated in Suppl. material 2.

2. Native Status

If no information is provided in the "Native status" field: Species is considered native to the Mediterranean.

If information is provided in the "Native status" field: Contains information on the native status of the species in Greece, following the terminology and definitions by Zenetos et al. (2011):

non-native (established): Species with at least two independent published records from Greece, separated in time and space;
non-native (casual): Reported only once in the scientific or grey literature from Greece;
non-native (questionable): Non-native species reported from Greece whose presence in the country is questionable and needs to be confirmed by an expert, or which have an unresolved taxonomic status. In this paper, doubtful records of species with an alien origin are not indicated as non-native if they have never been reported before as alien from the Mediterranean, whereas species previously reported as alien from the Mediterranean Sea but shown here to be questionable are retained as "non-native (questionable)" to highlight their status for further assessment by experts;
cryptogenic: Species of unknown origin that could be either native or introduced, clear evidence for either origin being absent.

3. Notes

If no information is provided in the "Notes" field: Species has an uncontroversial taxonomic and biogeographic status and regular records from Greece and the Mediterranean exist. A full reference list of all reports from Greece for each species can be found in Suppl. material 2.

If information is provided in the "Notes" field: May contain information on:

the taxonomic history, nomenclatural or occurrence status where these are deemed useful for assessing the taxonomic or occurrence status of the species in Greece.
literature references reporting the species from Greece and additional information on the species' distribution: only in cases where the species has been reported fewer than three times from Greece (excluding checklists and other secondary works).
whether the species has originally been described from the Mediterranean.

Distribution information:

Where statements are made on the occurrence (or absence) of the species in other areas of the Mediterranean, this information was derived from a combined assessment of:

the checklist of Mediterranean polychaetes by Martín and Gil (2010);
recent regional checklists of polychaetes in the Mediterranean (e.g. Çınar 2005, Castelli et al. 2008, Ayari et al. 2009, Zaâbi et al. 2009, Çınar et al. 2014, Mikac 2015);
extensive unpublished notes by C. Arvanitidis covering the literature on Mediterranean polychaetes until ca. 1994;
literature research.

Information on species distributions outside the Mediterranean are derived from a combined assessment of literature sources and distribution records in the Ocean Biogeographic Information System (OBIS) (unless a specific source is cited). Distribution information is not intended to list all known distributions of the species but rather to provide an estimate of the species’ likelihood to occur in the Mediterranean.

Questionable species:

Species are indicated as "questionable" if one or more of the following criteria are met:

the species concept is not clearly defined or outdated and in need of revision, rendering literature records highly uncertain;
species is indicated as possibly absent from the Mediterranean in a recent taxonomic work (preferably a review) based on examination of Mediterranean specimens;
species is in the Mediterranean reported exclusively (or almost exclusively) from Greece and has otherwise a distinctly different distribution range;
species has been reported only once in grey literature before 1993 (chosen as a cut-off date, as thorough investigations of the Greek polychaete fauna followed afterwards)

Species reported as "cf." in primary records are not indicated as questionable, as the modifier "cf." already implies uncertainties in the identification and often a possible deviation from the nominal species.

Species complexes:

Species known to constitute a complex of cryptic (morphologically indistinguishable but genetically different) or pseudo-cryptic (morphologically very similar) species are here treated as sensu lato, following Nygren and Pleijel (2011). Thus, clarifications pending through molecular analyses, we assume the presence of at least one species of the complex in Greece. The keyword "Species complex" in the Notes field indicates that the name has been applied to a species complex as defined above.

Data resources

Taxon treatment

Acoetidae Kinberg, 1856

Eupanthalis kinbergi McIntosh, 1876

Distribution:

Acrocirrus frontifilis (Grube, 1860)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Croatia).

Flabelligena mediterranea (Kolmer, 1985)

Nomenclature:

Flabelligella mediterranea Kolmer, 1985

Distribution:

Horizon:

Notes:

Originally described from the Hellenic Trench (Matapan Deep; 35°49'48''N, 22°20'42''E; 4690 m depth), no other records from Greece.

Macrochaeta clavicornis (M. Sars, 1835)

Ampharete grubei had long been considered a junior synonym of Ampharete acutifrons (Grube, 1860) and was reinstated by Jirkov (2001). Primary literature records from Greece under the name Ampharete grubei exist but were subsequently reported by Arvanitidis (2000a) and Simboura and Nicolaidou (2001) as Ampharete acutifrons.

Ampharete octocirrata (Sars, 1835)

Nomenclature:

Sabellides octocirrata (M. Sars, 1835)

Amphicteis gunneri (M. Sars, 1835)

Anobothrus gracilis (Malmgren, 1866)

Distribution:

Horizon:

Notes:

Reported from worldwide locations but probably restricted to circumpolar areas; specimens from other regions may belong to other species (Jirkov 2008).

Auchenoplax crinita Ehlers, 1887

Lysippe labiata Malmgren, 1866

Horizon:

Notes:

Reported from Greece by Arvanitidis (1994). In the Mediterranean also reported from the western basin (Fauvel and Rullier 1959), otherwise distributed in the Caribbean and tropical Atlantic. Chloeia viridis is the name prevailing in literature, with Chloeia candida as a synonym (referred to synonymy by Hartman (1948), although Barroso and Paiva (2011) stress that this synonymy should be re-evaluated). The publication year of Kinberg's species is sometimes cited as 1910, but the species was in fact described in 1857 (Kinberg 1857) and thus the name Chloeia candida takes priority.

Chloeia venusta Quatrefages, 1866

Distribution:

Horizon:

Notes:

Distribution:

Horizon:

Notes:

Questionable status. The Greek records of Notopygos megalops probably belong to a different species. Notopygos megalops was originally described based on a juvenile specimen from the Caribbean Sea and used to be considered a synonym of Notopygos crinita (Grube, 1855) before it was re-described by Yáñez-Rivera and Carrera-Parra (2012) from the Caribbean Sea based on adult characters. This re-description of Notopygos megalops differs from the description by Fauvel (1923) (which was probably used for the identification of the Greek specimens) in the following characters: Branchiae from chaetiger 6 (Yáñez-Rivera and Carrera-Parra 2012) instead of chaetiger 7 (Fauvel 1923); anterior branchiae with main stem and seven branchial filaments, in median chaetigers branching into four stems with five to seven filaments each (Yáñez-Rivera and Carrera-Parra 2012) vs. small tufts of 2–4 filaments anteriorly and only a single papilliform filament anteriorly Fauvel (1923). Yáñez-Rivera and Carrera-Parra (2012) define the start of the branchiae on chaetiger 6 as characteristic for the species; this seems to be invariable in juvenile individuals, too (McIntosh 1895). Thus, the species described by Fauvel (1923) and any specimens identified based on his description probably belong to a different species.

Pareurythoe borealis (M. Sars, 1862)

Distribution:

Nomenclature:

Hermione hystrix (Savigny in Lamarck, 1818) | Laetmonice hystrix (Savigny in Lamarck, 1818)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean.

Pontogenia chrysocoma (Baird, 1865)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean.

Arenicolidae Johnston, 1835

Abarenicola claparedi (Levinsen, 1884)

Nomenclature:

Abarenicola claparedi (Levinsen, 1884) | Arenicola claparedei Levinsen, 1884

Distribution:

Horizon:

Notes:

Type locality: Mediterranean.

Arenicola marina (Linnaeus, 1758)

Distribution:

Horizon:

Notes:

Questionable status. Reported from Greece by Antoniadou et al. (2006) from rocky shores. The record is considered questionable on the basis of habitat and geographic distribution (the species occurs in tidal mudflats of the European Atlantic coast and North Sea).

Branchiomaldane vincenti Langerhans, 1881

Distribution:

Horizon:

Notes:

Reported from Greece by Arvanitidis (1994) and Chatzigeorgiou et al. (2016). In the Mediterranean also known from Israel (Ben-Eliahu 1976b), Tunisia (Cantone et al. 1978), Spain (Alós 1990), Italy (Castelli et al. 2008), Egypt (Abd-Elnaby 2009), Turkey (Çınar et al. 2014) and the Adriatic (Mikac 2015), otherwise distributed in the East Atlantic (Gravina 1991).

Asteriomyzostomidae Jägersten, 1940

Asteriomyzostomum asteriae (Marenzeller, 1895)

Nomenclature:

Myzostoma asteriae Marenzeller, 1895

Distribution:

Horizon:

Notes:

Originally described from Greece as parasitic on the starfish species Sclerasterias richardi (Perrier in Milne-Edwards 1882) and Sclerasterias neglecta (Perrier, 1891) collected near Santorini, Kythira and Samos; no other records from Greece. Almost nothing is known of the species' distribution; the extensive description of its anatomy by Stummer-Traunfels (1903) is based on specimens provided by Marenzeller, presumably from the same collection as the type material.

Capitellidae Grube, 1862

Capitella capitata (Fabricius, 1780)

Nomenclature:

Capitella capitata (Fabricius, 1780) | Capitella capitata capitata (Fabricius, 1780)

Distribution:

Horizon:

Notes:

Questionable status. Species complex. Several sibling species have been separated from Capitella capitata sensu lato worldwide since Grassle and Grassle (1976) showed it to constitute a complex of species. Blake (2009) restricts Capitella capitata sensu stricto to Arctic areas and considers Mediterranean specimens, including the description by Fauvel (1927), to show significant differences to Capitella capitata and thus to belong to other species. In Greece, there is evidence so far of at least another sibling species population (Gamenick et al. 1998). However, because of the high morphological similarity of the various sibling species, they cannot be reliably distinguished in routine investigations (Blake 2000). See also notes under Capitella teleta Blake, Grassle & Eckelbarger, 2009.

Capitella giardi (Mesnil, 1897)

Nomenclature:

Nomenclature:

Leiocapitella dollfusi (Fauvel, 1936) | Leiocapitella glabra Hartman, 1947

Distribution:

Horizon:

Notes:

Bellan (1964a) proposed the synonymy of Leiocapitella glabra (type locality California) with Leiocapitella dollfusi (originally described from the Moroccan coasts) based on the overlapping variability in the chaetal formula of the two species. Ben-Eliahu and Fiege (1995), probably unaware of Bellan (1964a), proposed a possible synonymy of Leiocapitella dollfusi with Leiocapitella glabra on the basis of the identical hook dentition, in addition to the chaetal formula (Capaccioni-Azzati and El-Haddad 2015). However, it is noteworthy that the presence of branchiae in posterior segments, a character reported by Fauvel (1936a) for Leiocapitella dollfusi, has not been confirmed in material previously identified as Leiocapitella glabra (Hartman 1947, Ben-Eliahu and Fiege 1995), due to lack of posterior parts. In the present study, in agreement with Bellan (1964a), and Capaccioni-Azzati and El-Haddad (2015), we consider the older name Leiocapitella dollfusi as having priority over Leiocapitella glabra.

Leiocapitellides analis Hartmann-Schröder, 1960

Native status:

Non-native (casual)

Distribution:

Horizon:

Notes:

Reported from Greece by Akoumianaki (2004) and Neofitou et al. (2014). In the Mediterranean also known from Egypt (Abd-Elnaby 2009). Originally described from the Red Sea.

Leiochrus alutaceus Ehlers, 1908

Distribution:

Horizon:

Notes:

Questionable status. In the Mediterranean only reported from Greece (Akoumianaki 2004). Commonly distributed in the Pacific. Worldwide, records of this genus are rare and uncertain (e.g. Hilbig and Blake 2006, Probert et al. 2009). However, as species of Leiochrides can key out as Leiochrus in Fauchald (1977), the presence of the former genus in Greece is possible if the key by Fauchald has been used.

Mastobranchus trinchesii Eisig, 1887

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Gulf of Naples).

Mediomastus capensis Day, 1961

Native status:

Non-native (casual)

Native status:

Non-native (questionable)

Distribution:

Horizon:

Notes:

Questionable status. Reported from Greece by NCMR (1992), Simboura (1996) and Katsiaras and Simboura (2015). In the Mediterranean also known from Italy (Gravina and Somaschini 1990), Spain (Capaccioni-Azzati et al. 1992), Cyprus (Çınar 2005), Turkey (Çınar et al. 2014) and the Adriatic (Mikac 2015). Originally described from Brazil. However, Capaccioni-Azzati and El-Haddad (2015) list some noteworthy inconsistencies amongst Mediterranean records and the original description regarding the presence of chaetae on the peristomium and the last setiger with capillaries present. Therefore, it is likely that the Mediterranean records belong to an undescribed species.

Notomastus aberans Day, 1957

Native status:

Non-native (established)

Nomenclature:

Spiochaetopterus costarum (Claparède, 1869) | Telepsavus costarum Claparède 1869

Distribution:

Horizon:

Notes:

Species complex. Spiochaetopterus costarum was long believed to be a cosmopolitan species but in fact the name has been applied to a number of pseudo-cryptic species (e.g. Bhaud 1998, Bhaud et al. 2003). Mediterranean specimens are currently considered to belong to Spiochaetopterus costarum sensu stricto, as the type locality of the species is in the Gulf of Naples.

Spiochaetopterus solitarius (Rioja, 1917)

Nomenclature:

Phyllochaetopterus solitarius Rioja, 1917| Spiochaetopterus solitarius (Rioja, 1917)

Spiochaetopterus typicus M. Sars, 1856

Aphelochaeta filiformis (Keferstein, 1862) | Cirratulus filiformis Keferstein, 1862

Aphelochaeta marioni (Saint-Joseph, 1894)

Nomenclature:

Nomenclature:

Caulleriella bioculata (Keferstein, 1862) | Heterocirrus bioculatus (Keferstein, 1862)

Caulleriella viridis (Langerhans, 1881)

Distribution:

Horizon:

Notes:

Reported from Greece by Maidanou et al. (2017). The only other Mediterranean record is an unconfirmed record from Corsica (Ifremer 2016) and neither of the two Mediterranean reports is accompanied by a taxonomic description. Originally described from Madeira, little is known on its distribution range. If its presence is confirmed, the species should probably be considered non-native to the Mediterranean.

Chaetozone caputesocis (Saint-Joseph, 1894)

Nomenclature:

Caulleriella caputesocis (Saint-Joseph, 1894) | Chaetozone caputesocis (Saint-Joseph, 1894)

Chaetozone corona Berkeley & Berkeley, 1941

Nomenclature:

Chaetozone corona Berkeley & Berkeley, 1941

Native status:

Non-native (established)

Distribution:

Horizon:

Notes:

Reported from Greece by Simboura et al. (2010), found several times afterwards (HCMR, unpublished data). In the Mediterranean also known from Turkey (Çınar et al. 2014) and the Adriatic Sea (Munari et al. 2017). Chambers et al. (2011) doubt the validity of the Mediterranean records and the presence of the species in the Mediterranean (mainly on the basis of its Pacific distribution) but Le Garrec et al. (2016) present evidence of a wide distribution of the species in the Bay of Biscay, indicating that Chaetozone corona is an established alien species along the coasts of Europe and was probably introduced via shipping from its native distribution.

Chaetozone gibber Woodham & Chambers, 1994

Distribution:

Horizon:

Notes:

Reported from Greece by Simboura et al. (2010) and Katsiaras and Simboura (2015). In the Mediterranean also known from Tunisia (Zaâbi et al. 2012), Turkey (Çınar et al. 2014) and the Adriatic (Mikac 2015); otherwise distributed around the British Isles and along the European Atlantic coasts. Chambers et al. (2011) remark that records of Chaetozone gibber from the Mediterranean could in fact belong to other species, as the Mediterranean environment differs significantly from conditions in the species' native distribution range.

Chaetozone setosa Malmgren, 1867

Distribution:

Horizon:

Notes:

Questionable status. Species complex. Chaetozone setosa is a complex of pseudo-cryptic species, with at least five species in northern Europe alone (Chambers and Woodham 2003). Several authors (e.g. Simboura 1996, Çınar and Ergen 2007, Mikac 2015) doubt the validity of Mediterranean records, as the species has likely been confused with Chaetozone gibber Woodham & Chambers, 1994, Chaetozone corona Berkeley & Berkeley, 1941 or Chaetozone carpenteri McIntosh, 1911 and probably does not occur in the Mediterranean.

Chaetozone zetlandica McIntosh, 1911

Nomenclature:

Caulleriella zetlandica (McIntosh, 1911)

Distribution:

Horizon:

Notes:

Chambers et al. (2011) remark that records of Chaetozone zetlandica might be misidentifications, as the Mediterranean environment differs significantly from conditions in the species' native distribution range in the North Atlantic.

Cirratulus cirratus (O. F. Müller, 1776)

Distribution:

Horizon:

Notes:

Species complex. Often considered cosmopolitan, but ecological, morphological and genetic differences have been found in specimens from different geographical locations (Petersen 1999, Carr et al. 2011, Weidhase et al. 2014).

Cirriformia tentaculata (Montagu, 1808)

Nomenclature:

Audouinia tentaculata (Montagu, 1808) | Cirriformia tentaculata (Montagu, 1808)

Dodecaceria capensis Day, 1961

Native status:

Non-native (established)

Distribution:

Horizon:

Notes:

In the Mediterranean only reported from Greece (e.g. Bogdanos and Satsmadjis 1983, Bogdanos and Satsmadjis 1987, Simboura 1996; full reference list in Suppl. material 2). Known from South Africa. Identification of several Greek specimens confirmed by P. H. Gibson (pers. comm. in Simboura 1996).

Dodecaceria concharum Ørsted, 1843

Distribution:

Horizon:

Notes:

Nomenclature:

Caulleriella killariensis (Southern, 1914)| Heterocirrus killariensis (Southern, 1914) | Tharyx killariensis (Southern, 1914)

Timarete filigera (Delle Chiaje, 1828)

Nomenclature:

Cirriformia filigera (Delle Chiaje, 1828) | Timarete filigera (Delle Chiaje, 1828)

Distribution:

Horizon:

Notes:

Species complex. Timarete filigera has been recorded from worldwide locations, but many of these records probably belong to different species. Specimens from West Atlantic localities identified as Timarete filigera were re-examined morphologically and genetically by Magalhães et al. (2014) and shown to belong to undescribed species or to species previously treated as synonyms of Timarete filigera. Mediterranean specimens are currently considered to belong to Timarete filigera sensu stricto, as the type locality of the species is in the Gulf of Naples.

Timarete punctata (Grube, 1859)

Native status:

Non-native (casual)

Distribution:

Horizon:

Notes:

Species complex. Timarete punctata, originally described from the Caribbean, constitutes a complex of cryptic species. Molecular analyses of specimens morphologically identified as Timarete punctata by Seixas et al. (2017) revealed that samples from the West Atlantic and the Pacific comprise two cryptic species, one of them widely distributed and exhibiting a low genetic diversity, the other restricted to a single location. The authors hypothesise that the low genetic diversity of the widely distributed cryptic species could be attributed to a recent human-mediated introduction at these sites. In the Mediterranean reported from Greece (Maidanou et al. 2017) and Turkey (Çınar et al. 2014).

Cossuridae Day, 1963

Cossura coasta Kitamori, 1960

Native status:

Non-native (questionable)

Distribution:

Horizon:

Notes:

Questionable status. Arvanitidis (1994) questions the presence of the species in the area, as the description of the specimen found by Bogdanos and Fredj (1983) shows differences both to the species described by Kitamori (1960) and to Cossura soyeri Laubier, 1964. Without re-examination of the Mediterranean specimens referred to Cossura coasta, the status of the species in the area remains doubtful.

Type locality: Mediterranean (Adriatic). Considered cosmopolitan. However, Jumars (1974) notes differences between populations and suggests the possibility of more than one species existing under the name of Dorvillea rubrovittata.

Dorvillea similis (Crossland, 1924)

Native status:

Non-native (casual)

Distribution:

Horizon:

Notes:

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Águilas, Spain).

Protodorvillea atlantica (McIntosh, 1885)

Nomenclature:

Protodorvillea atlantica (McIntosh, 1885) | Schistomeringos atlanticus (McIntosh, 1885)

Distribution:

Horizon:

Notes:

Questionable status. Reported from Greece by Vamvakopoulou (1991). In the Mediterranean also known from Tunisia (Zaâbi et al. 2009) and the Adriatic (Mikac 2015). Originally described from deep waters off the Azores (Atlantic); as the Greek record is from a lagoonal habitat, it must be considered questionable.

Protodorvillea biarticulata Day, 1963

Native status:

Non-native (established)

Distribution:

Horizon:

Notes:

In the Mediterranean reported from Greece (Bogdanos and Satsmadjis 1983, Bogdanos and Diapoulis 1984, Simboura et al. 1995b) and Egypt (Abd-Elnaby and Gab-Alla 2007), otherwise distributed in the Indo-Pacific, South Africa and Brazil.

Protodorvillea kefersteini (McIntosh, 1869)

Nomenclature:

Protodorvillea kefersteini (McIntosh, 1869) | Schistomeringos kefersteini (McIntosh, 1869) | Staurocephalus kefersteini McIntosh, 1896

Schistomeringos neglecta (Fauvel, 1923)

Nomenclature:

Dorvillea neglecta (Fauvel, 1932) | Schistomeringos neglecta (Fauvel, 1923)

Schistomeringos rudolphi (Delle Chiaje, 1828)

Nomenclature:

Dorvillea rudolphi (Delle Chiaje, 1828) | Schistomeringos rudolphi (Delle Chiaje, 1828) | Staurocephalus rudolphi (Delle Chiaje, 1828)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Gulf of Naples).

Eulepethidae Chamberlin, 1919

Grubeulepis augeneri Pettibone, 1969

Distribution:

Horizon:

Notes:

One specimen from Greece in the collections of the Senckenberg Museum (SMF 15256, 35°52'58.2"N, 27°42'49.2"E, 115 m depth, coll. date 2005-09-05, det. D. Fiege). Type locality: Mediterranean (Adriatic).

Grubeulepis katzmanni Pettibone, 1986

Nomenclature:

Eunice harassii Audouin & Milne Edwards, 1833

Distribution:

Horizon:

Notes:

Records of Leodice harassii could include specimens of Eunice rubrocincta Ehlers, 1868, a Mediterranean species which Fauvel (1923) considered synonymous with Leodice harassii. Leodice rubrocincta was resurrected by Fauchald (1992) and is included as a valid species in the key by Carrera-Parra and Salazar-Vallejo (1998).

Leodice torquata (Quatrefages, 1866)

Nomenclature:

Eunice torquata Quatrefages, 1866 | Leodice torquata (Quatrefages, 1866)

Distribution:

Horizon:

Notes:

The species resembles Leodice laurillardi (Quatrefages, 1866) in a number of diagnostic characters. The latter was synonymised with Leodice torquata by Pruvot and Racovitza (1895); this was accepted by Fauvel (1923) and later authors. However, Fauchald (1992) resurrected Leodice laurillardi as a valid species and Arias et al. (2015) re-described the species based on type material and new specimens from the Mediterranean. They suggest that Leodice laurillardi could be much more widespread than previously thought but was possibly being overlooked in the Mediterranean due to the confusion with Leodice torquata.

Lysidice collaris Grube, 1870

Native status:

Non-native (questionable)

Distribution:

Horizon:

Notes:

Marphysa sanguinea (Montagu, 1813)

Distribution:

Horizon:

Notes:

Species complex. Considered cosmopolitan in the past, but Marphysa sanguinea actually comprises a complex of numerous pseudo-cryptic species (Zanol et al. 2016). Hutchings and Karageorgopoulos (2003) designated a neotype based on pectinate chaetae distribution instead of branchiae and Hutchings et al. (2012) encourage re-examination of all material outside the English Channel and North Sea. Recent studies have described several new species similar to Marphysa sanguinea or resurrected species from synonymy with the latter (Lewis and Karageorgopoulos 2008, Molina-Acevedo and Carrera-Parra 2015, Zanol et al. 2016, Liu et al. 2017). Wijnhoven and Dekker (2010) question the presence of Marphysa sanguinea in the Mediterranean Sea and encourage comparisons of specimens with the neotype.

Palola siciliensis (Grube, 1840)

Nomenclature:

Eunice siciliensis Grube, 1840 | Palola siciliensis (Grube, 1840)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Palermo, Sicily, Italy).

Euphrosinidae Williams, 1852

Euphrosine foliosa Audouin & Milne Edwards, 1833

Euphrosine myrtosa Savigny in Lamarck, 1818

Distribution:

Horizon:

Notes:

Questionable status. Reported from Greece by Bogdanos and Satsmadjis (1983). In the Mediterranean also reported from the Levantine Basin (Ben-Eliahu 1995) and questionably from the Adriatic (Mikac 2015), otherwise distributed in the Indo-Pacific. It might either be an overlooked alien species in the area or a misidentification.

Fabriciidae Rioja, 1923

Horizon:

Notes:

Originally described from Greece by Fitzhugh et al. (1994). See remarks under Pseudofabriciola analis Fitzhugh, Giangrande & Simboura, 1994.

Flabelligeridae de Saint-Joseph, 1894

Bradabyssa villosa (Rathke, 1843)

Nomenclature:

Brada villosa (Rathke, 1843)

Distribution:

Glycera alba (O.F. Müller, 1776)

Nomenclature:

Glycera alba (O.F. Müller, 1776) | Glycera minuta (Bobretzky, 1870)

Distribution:

Horizon:

Notes:

Böggemann (2002) distinguishes the species from Glycera tridactyla Schmarda, 1861, mainly on the basis of the shape of the proboscidial papillae and ailerons; although the separation was judged problematic by Worsfold (2007). Parapar and Moreira (2015) agree in separating the two species on the basis of the above two characters.

Glycera capitata Ørsted, 1843

Distribution:

Horizon:

Notes:

Frequently reported from the Mediterranean (e.g. Castelli et al. 2008, Ayari et al. 2009, Çınar et al. 2014, Mikac 2015), but Böggemann (2002) and Böggemann (2015) restricts its distribution to Arctic, Antarctic, cold temperate areas and deep sea down to 5655m (Böggemann 2015). The Mediterranean material could possibly be Glycera lapidum Quatrefages, 1866 or Glycera noelae Böggemann, Bienhold & Gaudron, 2012 (Böggemann, pers. comm.)

Glycera celtica O'Connor, 1987

Distribution:

Horizon:

Notes:

Böggemann (2002) synonymised Glycera dayi O' Connor, 1987 with Glycera celtica but this synonymy has not been accepted unanimously (Worsfold 2007, Parapar and Moreira 2015).

Glycera fallax Quatrefages, 1850

Nomenclature:

Glycera gigantea Quatrefages, 1866

Glycera lapidum Quatrefages, 1866

Distribution:

Horizon:

Notes:

O'Connor (1987) reports at least four varieties amongst the examined material from the North-East Atlantic, although these correspond in fact to different species (Böggemann, pers. comm.).

Glycera rouxii Audouin & Milne Edwards, 1833

Distribution:

Horizon:

Notes:

Böggemann (2002) considers Glycera rouxii a synonym of Glycera unicornis Lamarck, 1818 (based on comparisons with original description as the holotype is lost). He considers the shape of the retractable branchiae (reported as bifid in Glycera unicornis) not to be a diagnostic character, since variations were found in the same populations (Böggeman, pers. comm.). However, Worsfold (2007) and Parapar and Moreira (2015) debate this synonymy and highlight differences in parapodial lobes besides those of the branchiae, although parapodial lobes were found to change between juvenile and adult specimens by Böggemann (2002). They propose that the branchial shape remains a diagnostic character, which should however be used with caution due to retractability. Despite the available evidence for the synonymy, two recent works consider both species as valid. Therefore, the past records of Glycera rouxii in Greece (see Supplementary material) are currently retained as separate, until a response to the arguments of Parapar and Moreira (2015) is available. Type locality: Mediterranean (Marseille).

Glycera tesselata Grube, 1863

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Croatia).

Glycera tridactyla Schmarda, 1861

Nomenclature:

Glycera convoluta Keferstein, 1862 | Glycera tridactyla Schmarda, 1861

Distribution:

Horizon:

Notes:

Böggemann (2002) considers Glycera convoluta a synonym of Glycera tridactyla based on comparisons with the original description and illustrations. A number of authors support this synonymy (e.g. Worsfold 2007, Parapar and Moreira 2015).

Glycera unicornis Lamarck, 1818

Distribution:

Horizon:

Notes:

See notes under Glycera rouxii Audouin & Milne Edwards, 1833.

Goniadidae Kinberg, 1865

Glycinde bonhourei Gravier, 1904

Native status:

Non-native (casual)

Distribution:

Horizon:

Notes:

Reported from Greece by Simboura (2008). In the Mediterranean also known from Israel (Böggemann 2005), Egypt (Ben-Eliahu 1972a) and Turkey (Çınar et al. 2014). Originally described from Bonhoure (Djibuti).

Glycinde nordmanni (Malmgren, 1866)

Nomenclature:

Eone nordmanni Malmgren, 1866 | Glycinde nordmanni (Malmgren, 1866)

Goniada emerita Audouin & Milne Edwards, 1833

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Nice, France).

Goniada gigantea (Verrill, 1885)

Distribution:

Horizon:

Notes:

Reported from Greece by Böggemann (2005) from material collected by E. Marenzeller in 1894 on the Cyclades plateau (as Goniada emerita Audouin & Milne Edwards, 1833). In the Mediterranean also known from Spain and France (Böggemann 2005). Otherwise reported from the West and East Atlantic and North-East Pacific (Böggemann 2005).

Goniada hexadentes Böggemann & Eibye-Jacobsen, 2002

Distribution:

Horizon:

Notes:

Reported from Greece by Böggemann (2005) from the abyssal zone (3848 m) off the Peloponnese. In the Mediterranean also known from Spain, Italy and Israel (Böggemann 2005), otherwise reported from the Eastern Atlantic and the Indian Ocean (Böggemann 2005).

Goniada maculata Ørsted, 1843

Goniada norvegica Ørsted, 1845

Goniadella bobrezkii (Annenkova, 1929)

Distribution:

Horizon:

Notes:

In the Mediterranean only reported from Greece (Arvanitidis 2000a). Commonly distributed in the Black Sea (Annenkova 1929) and around the North-East Atlantic (Wolff and Stegenga 1975, Kirkegaard 2001).

Goniadella gracilis (Verrill, 1873)

Horizon:

Notes:

Schmidt and Westheide (2000) assess the genetic differentiation of worldwide Hesionides arenaria populations. Specimens from Crete group with those from near the type locality in the North Sea, indicating the presence of Hesionides arenaria sensu stricto in Greece.

Hesionides gohari Hartmann-Schröder, 1960

Distribution:

Horizon:

Notes:

Species complex. Schmidt and Westheide (1999) recover three genetic lineages from worldwide localities which could correspond to cryptic species. Specimens from Crete group with those from near the type locality in the Red Sea, indicating the presence of Hesionides gohari sensu stricto in Greece.

Hesiospina aurantiaca (M. Sars, 1862)

Nomenclature:

Hesiospina aurantiaca (M. Sars, 1862) | Hesiospina similis (Hessle, 1925)

Nomenclature:

Gyptis mediterranea Pleijel, 1993

Distribution:

Horizon:

Notes:

Distribution:

Horizon:

Notes:

Reported from Greece by Arvanitidis (1994) based on a single specimen. Type locality: Mediterranean (Banyuls-sur-Mer, France). Pleijel (1998) considers the species a junior synonym of Podarkeopsis arenicolus (La Greca, 1946) but does not provide details and does not formalise the synonymy. As Rizzo and Salazar-Vallejo (2014) list both species in their key and recognise difference in the size of the eyes, length of the median antenna and chaetal details, the species is therefore treated as valid here, pending further information.

Psamathe fusca Johnston, 1836

Nomenclature:

Kefersteinia cirrhata (Keferstein, 1862)

Syllidia armata Quatrefages, 1866

Nomenclature:

Magalia perarmata Marion & Bobretzky, 1874 | Syllidia armata Quatrefages, 1866

Iospilidae Bergström, 1914

Phalacrophorus pictus Greeff, 1879

Distribution:

Heterospio mediterranea Laubier, Picard & Ramos, 1973

Distribution:

Horizon:

Notes:

In the Mediterranean known from Greece, Turkey (Çınar et al. 2014) and Cyprus (Çınar 2005), otherwise distributed along the Atlantic coast of Europe.

Lumbrinerides carpinei (Ramos, 1976)

Distribution:

Horizon:

Notes:

New record for Greece. Two specimens, Pachia Ammos, Crete, 35°06'39.6"N, 25°48'32.4"E, 1–5 m depth, fine to coarse sand, collected in the framework of the MEDCORE project (unpublished data from Papageorgiou et al. 2006). Literature used for identification: Ramos (1976). Type locality: Western Mediterranean.

Lumbrinerides laubieri Miura, 1980

Distribution:

Horizon:

Notes:

In the Mediterranean only reported from Greece (Simboura and Zenetos 2005). Known from deep waters off the Atlantic coast of Europe.

Lumbrinerides neogesae Miura, 1981

Native status:

Non-native (casual)

Distribution:

Horizon:

Notes:

New record for Greece. One specimen, Elafonisi, Crete, 35°16'20.7"N, 23°32'15.9"E, 1 m depth, fine sand, collected in the framework of the MEDCORE project (unpublished data from Papageorgiou et al. 2006). Literature used for identification: Miura (1980), Gravina and Cantone (1991). In the Mediterranean also known from Italy (Gravina and Cantone 1991). Originally described from South Africa.

Lumbrineriopsis paradoxa (Saint-Joseph, 1888)

Nomenclature:

Lumbriconereis paradoxa Saint-Joseph, 1888 | Lumbrineriopsis paradoxa (Saint-Joseph, 1888) | Lumbrineris paradoxa Saint-Joseph, 1888

Lumbrineris coccinea (Renier, 1804)

Nomenclature:

Lumbriconereis coccinea (Renier, 1804)| Lumbrineris coccinea (Renier, 1804)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean. Carrera-Parra (2006) considers records from outside the Mediterranean Sea doubtful.

Lumbrineris inflata Moore, 1911

Distribution:

Horizon:

Notes:

Questionable status. Reported from Greece by Arvanitidis (2000a) and Kitsos (2003); the first Mediterranean record was by Giangrande et al. (1981). Arvanitidis (2000a) argues that the species may have been underreported in the Mediterranean due to its similarity with Lumbrineris coccinea (Renier, 1804). Carrera-Parra (2006) and D’Alessandro et al. (2016) consider Mediterranean records of the species doubtful based on its otherwise East Pacific distribution. Çınar (2009) argues that the description and illustrations of Lumbrineris inflata by Giangrande et al. (1981) fit the characters of Lumbrineris perkinsi Carrera-Parra, 2001. Lumbrineris inflata differs from Lumbrineris perkinsi mainly in the number of teeth of maxilla III (M III). Giangrande et al. (1981), as well as Arvanitidis (2000a) for Greece, report a maxillary formula which could correspond to Lumbrineris perkinsi ("M III with 3–4 teeth"). Although published descriptions and records from the Mediterranean indicate the presence of Lumbrineris perkinsi, re-examination of all the material is needed to draw conclusions about the identity of the Greek records.

Lumbrineris latreilli Audouin & Milne Edwards, 1834

Nomenclature:

Lumbriconereis latreilli Audouin & Milne Edwards, 1834 | Lumbrineris latreilli Audouin & Milne Edwards, 1834

Distribution:

Horizon:

Notes:

Frequently reported from Greece, however, mainly on the basis of external and unreliable diagnostic characters. Species descriptions focusing on the maxillary apparatus and other unambiguous and size-independent characters have recently become available (Carrera-Parra 2006, Martins et al. 2012, D’Alessandro et al. 2016). Therefore, re-examination of material could reduce the frequency of records and even reveal the presence of overlooked species. In the Mediterranean, the species is widely recorded (e.g. Campoy 1982, Çınar 2005, Castelli et al. 2008, Ayari et al. 2009, Çınar et al. 2014, Mikac 2015). Otherwise distributed in the North-East Atlantic; its presumed cosmopolitan distribution is questionable (Carrera-Parra 2006).

Lumbrineris luciliae Martins, Carrera-Parra, Quintino & Rodrigues, 2012

Distribution:

Horizon:

Notes:

New record for Greece. Specimens previously reported as as Lumbrineris cingulata Ehlers, 1897 by Chatzigeorgiou et al. (2016) from rocky shores in northern Crete were re-examined and found to belong to Lumbrineris luciliae as described by Martins et al. (2012), although the reported slight distal curvature of the aciculae could not be observed. In the Mediterranean also known from Italy (D’Alessandro et al. 2016). Otherwise known from the Atlantic coast of the Iberian Peninsula (Martins et al. 2012).

Lumbrineris nonatoi Ramos, 1976

Lumbrineris tetraura (Schmarda, 1861) | Scoletoma tetraura (Schmarda, 1861)

Distribution:

One specimen from Greece in the collections of the Senckenberg Museum (SMF 11286, 39°15'0.6"N, 23°42'32.4"E, 1243 m, coll. date 1998-01-01, det. D. Fiege). In the Mediterranean also known from the coasts of France and Spain (Labrune et al. 2007, Serrano-Samaniego 2012) and the Adriatic (Mikac 2015).

Maldanidae Malmgren, 1867

Axiothella constricta (Claparède, 1869)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Gulf of Naples).

Axiothella rubrocincta (Johnson, 1901)

Distribution:

Horizon:

Notes:

Questionable status. Species complex. In the Mediterranean only reported from Greece (Dando et al. 1995). Distributed along the Pacific coast of North America where at least two morphologically indistinguishable forms with different life histories occur (Wilson 1983).

Chirimia biceps (M. Sars, 1861)

Nomenclature:

Asychis biceps (M. Sars, 1861) | Chirimia biceps (M. Sars, 1861) | Chirimia biceps biceps (Sars, 1861)

Clymenella cf. koellikeri (McIntosh, 1885)

Distribution:

Horizon:

Notes:

In the Mediterranean only reported from Greece (Simboura 1996). The specimens differ from Clymenella torquata (Leidy 1855) (originally from the East coast of North America but occurring non-natively on the Atlantic coast of Europe) in the shape and number of acicular chaetae and from Clymenella cincta (Saint-Joseph, 1894) (occurring in the Mediterranean) in the shape of the collar and of the nuchal organs. The Greek specimens are most similar to the Pacific species Clymenella koellikeri, from which they differ in the following characters: Nuchal slits of Clymenella koellikeri reach up to the middle of the cephalic plate (until the lateral notches in the Greek material) and the rim or the collar of the 4^th chaetiger is smooth or slightly wavy in Clymenella koellikeri, whereas one Greek specimen shows a small lateral recess at the rim of the collar (Simboura 1996). The species is found regularly in Greece but is usually reported at genus level due to its uncertain identity. It may be an overlooked or cryptic species.

Euclymene collaris (Claparède, 1869)

Nomenclature:

Clymene collaris (Claparède, 1869) | Euclymene collaris (Claparède, 1869)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Gulf of Naples).

Euclymene lombricoides (Quatrefages, 1866)

Nomenclature:

Clymene lombricoides Quatrefages, 1866 | Euclymene lombricoides (Quatrefages, 1866)

Euclymene oerstedii (Claparède, 1863)

Nomenclature:

Clymene oerstedii Claparède, 1863 | Euclymene oerstedii (Claparède, 1863)

Euclymene palermitana (Grube, 1840)

Nomenclature:

Clymene palermitana Grube, 1840 | Euclymene palermitana (Grube, 1840) | Praxillella palermitana (Grube, 1840)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Palermo, Sicily, Italy).

Heteroclymene robusta Arwidsson, 1906

Nomenclature:

Clymene robusta (Arwidsson, 1906) | Euclymene robusta (Arwidsson, 1906) | Heteroclymene robusta Arwidsson, 1906

Johnstonia clymenoides Quatrefages, 1866

Distribution:

Horizon:

Notes:

Reported from Greece by Bogdanos and Satsmadjis (1983) and Xenopoulou (1987). In the Mediterranean also known from France, Spain and Israel (Mackie and Gobin 1993), Egypt (Abd-Elnaby 2008), Turkey (Çınar et al. 2014) and the Adriatic (Mikac 2015), otherwise distributed along the European and North African Atlantic coasts (Mackie and Gobin 1993).

Leiochone leiopygos (Grube, 1860)

Nomenclature:

Clymenura clypeata (Saint-Joseph, 1894) | Leiochone clypeata Saint-Joseph, 1894 | Leiochone leiopygos (Grube, 1860)

Lumbriclymene minor Arwidsson, 1906

Macroclymene santandarensis (Rioja, 1917)

Nomenclature:

Clymene santandarensis Rioja, 1917 | Euclymene santandarensis (Rioja, 1917) | Macroclymene santandarensis (Rioja, 1917)

Maldane capensis (Day, 1961)

Nomenclature:

Asychis capensis Day, 1961 | Maldane capensis (Day, 1961)

Distribution:

Horizon:

Notes:

Questionable status. In the Mediterranean only reported from Greece (Papadopoulos 1986). Known from South Africa.

Maldane glebifex Grube, 1860

Maldane sarsi Malmgren, 1865

Maldanella harai (Izuka, 1902)

Distribution:

Horizon:

Notes:

Questionable status. In the Mediterranean only reported from Greece (Bogdanos and Nicolaidou 1985, Bogdanos and Satsmadjis 1987, Nicolaidou and Papadopoulou 1989, Koulouri et al. 2015). Outside the Mediterranean reported from worldwide locations, mostly from bathyal and abyssal depths.

Metasychis gotoi (Izuka, 1902)

Nomenclature:

Asychis gotoi (Izuka, 1902) | Metasychis gotoi (Izuka, 1902)

Native status:

Non-native (established)

Distribution:

Horizon:

Notes:

Originally distributed in the Red Sea, the Indian Ocean and the North Pacific; commonly reported from locations throughout the Mediterranean (Simboura and Zenetos 2005). The first Mediterranean record of Metasychis gotoi is from the Northern Adriatic (Fauvel 1940), constituting one of the earliest records of non-native species in the basin. As Metasychis gotoi is reported from worldwide distributions, it is likely that the name refers to a complex of cryptic species. The Mediterranean material could belong to an overlooked native species (J. Langeneck, pers. comm.).

Micromaldane ornithochaeta Mesnil, 1897

Distribution:

Horizon:

Notes:

Reported from Greece by Eleftheriou et al. (1990) and NCMR (2000a). In the Mediterranean also known from Israel (Ben-Eliahu 1976b), Spain (Alós 1990), Italy (Gherardi et al. 2002), Egypt (Abd-Elnaby 2008) and the Adriatic (Mikac 2015). Present in the adjacent Sea of Marmara (Çınar et al. 2014). Otherwise distributed along the European Atlantic coasts (Gherardi et al. 2002).

Nicomache lumbricalis (Fabricius, 1780)

Distribution:

Horizon:

Notes:

Questionable status. Frequently reported from Greece and other Eastern Mediterranean locations; however, de Assis et al. (2007) in their review of the genus restrict its distribution to boreal regions.

Nicomache maculata Arwidsson, 1911

Distribution:

Horizon:

Notes:

Questionable status. In the Mediterranean only reported from Greece (e.g. Bogdanos and Satsmadjis 1983, Eleftheriou et al. 1990, Tselepides 1992, Simboura 1996; full reference list in Suppl. material 2), de Assis et al. (2007) restrict its distribution to the Shetland Islands.

Nicomache trispinata Arwidsson, 1906

Distribution:

Horizon:

Notes:

Reported from Greece by Tselepides (1992). In the Mediterranean also known from France (Clausade 1969), Israel (Ben-Eliahu 1995) and Italy (Castelli et al. 2008), otherwise distributed along the Atlantic coasts of Europe and in the North Sea.

Petaloproctus terricolus Quatrefages, 1866

Praxillella affinis (M. Sars in G.O. Sars, 1872)

Nomenclature:

Clymene affinis M. Sars in G.O. Sars, 1872 | Praxillella affinis (M. Sars in G.O. Sars, 1872)

Praxillella gracilis (M. Sars, 1861)

Nomenclature:

Clymene gracilis Sars, 1861| Praxillella gracilis (M. Sars, 1861)

Praxillella lophoseta (Orlandi, 1898)

Nomenclature:

Clymene lophosetosa Orlandi, 1898 | Praxillella lophoseta (Orlandi, 1898)

Distribution:

Horizon:

Notes:

Type locality: Mediterranean (Gulf of Naples).

Praxillella praetermissa (Malmgren, 1865)

Distribution:

Horizon:

Notes:

Species complex. At least three putative cryptic species exist, one each in the Arctic and West Atlantic (Carr et al. 2011) and another in Portugal (Lobo et al. 2016).

Praxillura longissima Arwidsson, 1906

Distribution:

Horizon:

Notes:

Questionable status. In the Mediterranean only reported from Greece (Eleftheriou et al. 1990). Commonly distributed in the North-East Atlantic.

Rhodine gracilior Tauber, 1879

Rhodine loveni Malmgren, 1865

Myzostomatidae Benham, 1896

Myzostoma cirriferum Leuckart, 1836

Distribution:

Horizon:

Notes:

Reported from Greece by Chatzigeorgiou et al. (2016) based on a single specimen. In the Mediterranean also known from Tunisia (Wesenberg-Lund 1939b), France (Laubier and Paris 1962), Italy (Castelli et al. 2008) and the Adriatic (Mikac 2015), otherwise distributed along the Atlantic coasts of Europe (Nygren and Pleijel 2015).

Nephtyidae Grube, 1850

Rarely reported from the Mediterranean, but was considered a synonym of Nephtys hombergii Savigny in Lamarck, 1818 until its resurrection and redescription by Rainer (1989) and could therefore be underreported.

Nephtys caeca (Fabricius, 1780)

Distribution:

Horizon:

Notes:

Ravara et al. (2010) re-examined material from the Mediterranean and identified it as Nephtys caeca, concluding that, although the species is commonly distributed in Arctic waters, it occasionally occurs in warmer waters of the Mediterranean as far as the Black Sea.

Nephtys ciliata (Müller, 1788)

Distribution:

Alitta succinea (Leuckart, 1847)

Nomenclature:

Neanthes succinea (Leuckart, 1847) | Nereis succinea Leuckart, 1847

Distribution:

Horizon:

Notes:

Fauvel (1923) considered Nereis lamellosa Ehlers 1868 a synonym of Nereis succinea but revoked this synonymy in a subsequent publication, describing the differentiating characters (Fauvel 1936b). Later authors (e.g.