FORUM

In polychaete taxonomy, as in many other scientific areas, there are some very influential scientists; these scientists usually were extremely productive and their publications included several faunistic studies or revisions. At least among polychaete taxonomists, these efforts often made them believe cosmopolitan species were common in the group. This conclusion usually overlooked basic differences in mean water temperature, sediment type, water depth, and salinity, what has been regarded as the ecological horizon (Salazar-Vallejo, Carrera-Parra, González, & Salazar-González, 2014). Foremost among these scientists are Olga Hartman and Pierre Fauvel and despite their efforts, they did not always include the study of type material for their nomenclatural acts, such that their conclusions were mostly based upon publications. This was a difficult task because descriptions and illustrations, if any available together with the descriptions, were not standardized until recently, and at the same time, diagnostic features have been continually refined. The result of these comparisons of very heterogeneous precedent works is that there are many taxonomic problems in polychaete species names, and current taxonomists must devote some time for solving earlier confusion, or inadequate taxonomic procedures because this is a very relevant activity.

Our objective with this contribution was to analyse what is the situation for Nereis falsa de Quatrefages, 1866. For this, we have made a careful study of previous publications and with our understanding of the systematics of nereidid polychaetes, we hope to clarify the current situation, to propose some means for promoting discussion and actions, and suggested some basic research for solving the issue.

For this purpose, all relevant publications were collected and the corresponding sections for N. falsa were translated and, whenever relevant, were fully quoted in order to show earlier ideas or conclusions. Documents available in our personal libraries or downloadable from Biodiversity Heritage Library, Internet Archive, and la Fédération Française des Sociétés de Sciences Naturelles were carefully studied.

Historical account

As a part of the Fauna Ibérica series, Núñez (2004) made an examination of the genus Nereis Linnaeus, 1758 in the Mediterranean region. He listed N. pulsatoria (Savigny, 1822) and N. splendida Grube, 1840, but N. falsa de Quatrefages, 1866 was not recognized. Characters for N. pulsatoria include antennae about as long as the palps, a smooth anterior peristomial margin which is not projected, and homogomph falcigers with short, slightly denticulated blades. Additionally, area VI has 4-10 paragnaths in two transverse series, and areas VII-VIII have paragnaths in 3-4 series. In N. splendida antennae are shorter than palps, peristomial anterior mid-dorsal margin projects anteriorly, pharyngeal area VI has 3 paragnaths in an inverted triangle or 4(-5) paragnaths in a rhombus, areas VII-VIII have paragnaths in several series, and the homogomph falcigers have long, markedly denticulated blades. This variation has been indicated before by Amoureux (1976: 340), although he was referring to N. falsa, and Gravina, Lezzi, Bonifazi, and Giangrande (2015: 159) illustrated the morph having 3 paragnaths in area VI for N. falsa.

It is interesting that for N. splendida two patterns are recognized for area VI despite the fact there is usually a very low variation in paragnath number in this area (González-Escalante & Salazar-Vallejo, 2003; Conde-Vela & Salazar-Vallejo, 2015). Further, Alós et al. (2004) listed the junior synonyms of N. pulsatoria as N. zonata Malmgren, 1867 described from Spitsbergen and N. cylindrata Ehlers, 1868 from the Adriatic Sea, whereas for N. splendida they included N. falsa de Quatrefages, 1866 originally described from the Black Sea and N. parallelogramma Claparède, 1868 from the Mediterranean Sea. How did we arrive to this point?

Rathke (1837) made a large report on the fauna of Crimea and its shores on the Black Sea, including mammals, amphibians, reptiles, fishes (8 new species), crustaceans (15 n. spp), worms (4 n. spp), and cnidarians (1 n. sp). In the section devoted to worms, Rathke (1837: 412-415, Pl. 7, Fig. 1, Fig. 4-8) made a thorough description of some of his nereidids from Balaklava Bay (44°30’ N, 33°36’ E) as N. pulsatoria? This was because he noted some differences from the description of N. pulsatoria (Savigny, 1822) such as body and eye pigmentation, and the size of the upper parapodial lobes, with dorsal cirri clearly longer than upper notopodial lobes in median and posterior chaetigers. However, the arrangement of paragnaths or fine details of the chaetal blades were not clarified or illustrated. Rathke compared his specimens with N. pulsatoria (Savigny, 1822: 33), and not after Lycoris pulsatoria Savigny in Lamarck (1818: 313) as listed in WoRMS (2016), because the latter is a nomen nudum.

For parapodial features, Savigny (1822: 33) indicated: “Cirres courts; le cirre supérieur n’atteint pas même le sommet de la branchie.” [Transl.: Cirri short; the dorsal one does not reach branchial (upper notopodial ligules) tips]. Rathke also had at hand the series by Audouin and Milne-Edwards (1832: Pl. 13, Figs. 8-13, 1833: 216-217), who provided a description and high quality illustrations for N. pulsatoria, now regarded as a junior synonym of N. zonata Malmgren, 1867 in WoRMS.

Grube (1840: 73-74) studied Rathke’s material and added some features to the earlier description, especially regarding the chaetal blades and paragnath arrangement on the pharynx, but made no further comment about its identity. For chaetal blades, Grube (1840: 73) indicated: “diesen die Anhängsel der langen Borsten gesägt erschienen, indessen treten die Zähnehen erst bei einer bedeutenderen Vergrösserung hervor.” [Transl.: the blades of the long chaetae are denticulate, but their number is only evident with a more powerful enlargement]. The paragnath pattern details are very difficult to understand, indeed. In the same contribution he described N. cultrifera (now in Perinereis), N. imbecillis (incertae sedis fide Fauvel, 1923: 362), and N. splendida. For the latter Grube (1840: 75-76) stated that the body was golden with dorsal transverse lines on each segment, the antennae were smaller than the palpophore, the longest tentacular cirri reach segment 3, and some chaetae had an additional tooth. Grube also indicated the pharyngeal paragnath patterns resembled those present in N. pulsatoria but the pharynx was not everted. This latter feature led Fauvel (1916: 81) to consider N. splendida as indeterminable.

de Quatrefages (1866: 505) proposed N. falsa for what Rathke had regarded as N. pulsatoria? However, it must be emphasized that he made no formal description or illustration for N. falsa, nor had he any specimens. Consequently, based upon Rathke’s description and illustration, and in comparison with Savigny or Audouin and Milne-Edwards, de Quatrefages indicated that: “La tête est plus courte; les antennes latérales plus grosses …; les tentacules sont plus longs; l’anneau buccal égale en longuer les deux suivants … Aux pieds, les sois sont plus nombreuses et les languettes à peu près égales” [Transl.: The head is shorter; the antennae thicker …; the tentacular cirri longer; the peristomium as long as the following two segments … In parapodia, chaetae are more abundant and lobes almost equal (to each other)]. No type material exists because Rathke’s collection was not preserved in the Kantiana University in Tartu (olim Dorpat), Estonia.

Some accounts include Nereis clava Leach in de Blainville, 1825 (p. 439), despite the fact that Hartman (1959: 256) regarded it as a nephtyid or indeterminable. It must be rejected from this discussion because de Blainville (1825: 439-440) compared it to what we now regard as Nephtys ciliata (Müller, 1778), Audouin and Milne-Edwards (1833: 257) regarded it as a junior synonym of N. hombergi Savigny in Lamarck, 1818, and de Quatrefages (1866: 434) confirmed its similarities with Nephtyidae.

Claparède (1868: 477, Pl. 9, Fig. 7, 10, Fig. 2) described N. (Nereilepas) parallelogramma by emphasizing that his species was “évidemment distincte de la N. pulsatoria Mont. (Sav.) avec laquelle M. Grube l’a confondue. La seule proportion des cirres suffirat déjà à la différencier, car ils dépassent notablement la languette supérieure chez notre espèce, tandis qu’ils sont plus courts qu’elle chez la N. pulsatoria” [Transl.: evidently distinct from N. pulsatoria Mont. (Sav.) with which it was confused by Mr. Grube. The cirri proportion will be enough to differentiate them, because they markedly surpass the dorsal lobe in our species, whereas they are shorter in N. pulsatoria]. In the following page, Claparède (1868: 478) referred to the paragnath pattern in the pharynx and explained the etymology: “L’article basilaire de la trompe est renflé sur le dos en deux éminences portant quelques paragnathes plus gros que ceux des autres groups. Ils sont en général au nombre de quatre, disposes en parallélogramme” [Transl. The basal pharynx ring is bulged dorsally into two projections larger than those present in other groups. They are generally four in number, arranged in a parallelogram].

Synonymy: Advances and Retreats

Grube (1873: 70, p. 15 in preprint) included many known nereidid species into several genera and groups and concluded that Nereis: “parallelogramma Clap., welche mit der von mir beschriebenen N. splendida zusammenfällt” [Transl.: parallelogramma Clap., coincides with the N. splendida described by me].

Von Marenzeller (1874: 466-470, Pl. 7, Fig. 3) first recorded Hediste diversicolor Müller, 1776 for the Mediterranean Sea with specimens from Muggia Bay, Gulf of Trieste, which had been found living in fresh water. He made an extensive description and in the footnotes (p. 469) he confirmed the previous synonymy by Grube, and concluded: “N. falsa Quatrefages, eine N. pulsatoria Audouin & Milne-Edwards, ja selbst eine N. diversicolor gewesen sein.” [Transl.: N. falsa Quatrefages, a N. pulsatoria Audouin and Milne-Edwards, can be the same as N. diversicolor].

Bobretzky (1881: 192-193) indicated that by (Transl.) “recognizing that Lycoris pulsatoria Rathke is categorically different from a nereidid described under this name either by Savigny (1822: 33) or by Audouin and Milne-Edwards (t. XXVII, 1832: pl. XIII, fig. 8-13 and t. XXIX, 1833: 216), Quatrefages (1866) established for Black Sea specimens a distinct species, N. falsa. Marenzeller (1874) recently supposed N. falsa to be the same as N. parallelogramma Clprd., which Claparède (1868: 477 and 1870: 84) recognized as the same species as that identified by Grube (1840: 73), in comparison with the authentic specimens of Rathke, identified as Nereis pulsatoria. On this basis Marenzeller proposed to forget the name provided by Claparède to the above mentioned species and to replace it by the older one, N. falsa.” Further, Bobretzky (1881), after a thorough study of L. pulsatoria? Rathke, synonymyzed it with Hediste diversicolor. This conclusion was followed by Hartman (1959: 259) as she regarded N. falsa as a junior synonym of H. diversicolor.

Fauvel (1923: 335-336), apparently overlooked Bobretzky (1881), and keyed-out N. falsa as closely allied to N. pelagica Linnaeus, 1758. He distinguished these two species by their chaetal blades: short, blunt, rather smooth in N. pelagica, while they are long, denticulate, with a distal tooth fused to blade’s tip in N. falsa. Fauvel (1923: 337), then listed, under N. falsa his early ideas (Fauvel 1913: 63, 1916: 81) about junior synonyms: N. splendida Grube, 1840 (indeterminable), N. parallelogramma Claparède, 1868, N. pervisceralis Claparède, 1868, N. lucipeta Ehlers, 1908, and N. splendida sensu Ehlers, 1913. These species deserve some comments.

For N. splendida, Fauvel (1916: 83) noted that Grube (1873: 70, p. 15 in preprint) had regarded his species as a synonym of N. parallelogramma, and that “Il est fort possible qu’il s’agisse de la même espèce, mais la description de Grube, sans figures, bien qu’assez détaillée, ne precise pas suffisamment certains points pour que cette identité puisse être reconnue d’une façon indubitable.” [Transl.: It is highly likely that it is the same species, but Grube’s description, without figures, and despite its many details, does not sufficiently specify several points for regarding this identity without any doubt]. Actually, Grube (1873: 70) grouped N. diversicolor, N. pelagica, N. parallelogramma and his N. splendida due to the paragnath patterns of their pharynx: area VI with 4(-5) paragnaths in a group, and he indicated that his species was coincident with N. parallelogramma. This conclusion has been indicated by Claparède (1868: 478) but he referred to what Grube indicated for N. pulsatoria, not for N. splendida. Claparède did not deposit any of his specimens because he wanted other naturalists to study living organisms rather than preserved ones.

Including N. perivisceralis in the list of synonyms was incorrect, despite the small size of the specimens (1 cm), because Claparède (1868: 471, Pl. 12, Fig. 1) stated that its pharynx has a single series of paragnaths in areas VII-VIII, whereas N. parallelogramma has 2(-3) transverse series in areas VII-VIII, although they were not illustrated. There is often a great deal of size-dependent variation but this refers to the number of paragnaths per series, not to the number of series.

Ehlers (1908: 69, Pl. 8, Figs. 7-13) described N. lucipeta based upon some atokous and epitokous specimens collected in Southern Angola. It is very similar to Mediterranean epitokous specimens illustrated by Fauvel (1916, Pl. 5, Figs. 4-7), but there are some subtle differences in parapodial features in anterior chaetigers, especially in the dorsal cirri. In Mediterranean specimens the dorsal cirri in chaetiger 7 are swollen along 2/3 of its length and it is twice longer than wide, whereas in N. lucipeta it is swollen along ¾ of its length and it is three times longer than wide. Further, in subsequent segments the dorsal cirri are barely longer than the upper notopodial ligules, whereas in N. lucipeta they are twice longer. Consequently, N. lucipeta must be regarded as a distinct species. The syntype series of N. lucipeta is in Berlin (ZMB 4440). A later record by Ehlers (1913: 496) as N. splendida from Simonstown, South Africa, must correspond to the atokous form of N. lucipeta and as such, could be the one characterized and illustrated by Day for South Africa (1962: 639, 1967: 317, Fig. 14.7k-o). This specimen might also be in Berlin, but it was not listed by Hartwich (1993) since it was not a species newly described by Ehlers. It is interesting to note that Ramsey (1914: 212) with some South African specimens, regarded N. lucipeta as a junior synonym of N. pelagica, and that Augener (1918: 184) listed N. lucipeta as a junior synonym of N. callaoana (Grube, 1857) and recorded this Eastern Pacific species from Togo and Namibia. These specimens must be studied to clarify this supposed affinity.

In fact, Day (1962: 639) followed Hartman (1959: 271) by pointing out another important issue: N. splendida Grube, 1840 is a junior homonym of N. splendida de Blainville, 1825. Audouin and Milne-Edwards (1833: 257) indicated that de Blainville introduced the name for N. clava (see above), and that it is a junior synonym of N. hombergi. Anyway, N. splendida Grube, 1840 must be replaced (ICZN, 1999: Ch. 12, Art. 60), as has been recently indicated by Gravina et al. (2015: 162).

Distribution

As indicated above, N. falsa was based upon Nereis pulsatoria? Rathke, 1837 from the Black Sea, and it is currently regarded as having a very wide distribution. This species has been frequently recorded from the Mediterranean Sea: in the harbour of Monaco (Fauvel, 1913) and included in the Fauna of France (Fauvel, 1923), in Alexandria, Egypt (Fauvel, 1937), in the Corsica Channel (Aliani & Meloni, 1999), in the Algeciras Bay (Sánchez-Moyano, García-Adiego, Estacio, & García-Gómez, 2001), in the National Park of Circeo in Italia (Andrea & Giancarlo, 2003), in the Bay of Izmir (Çinar et al., 2008), in Greece (Faulwetter, 2010), and in the region of Ceuta, Strait of Gibraltar (Guerra-García, Gonzáles-Vila, & García-Gómez, 2003). Several other records have been made for Western Africa: Morocco (Fauvel, 1936; Gillet, 1986, 1988), Western Sahara (Rullier & Amoureux, 1969), Mauritania (Gillet, 1990), Senegal (Fauvel, 1950; Sourie, 1954; Fauvel & Rullier, 1957, 1959) and Guinea (Amoureux, 1973).

According to WoRMS (2016), N. falsa has also been recorded for the Gulf of Mexico (Taylor, 1984; Salazar-Vallejo & Jiménez-Cueto, 1997; Felder & Camp, 2009), Caribbean Sea (de León-González, Solís-Weiss, & Ochoa-Rivera, 1999; Miloslavich et al., 2010) and Brazil (Amaral, Nallin, Steiner, Forroni, & Filho, 2013). The same species has been recorded for the Indian Ocean in Madagascar (Fauvel, 1919), and from Durban Bay, South Africa (Day & Morgans, 1956; Day, 1962, 1967). It is remarkable that despite the fact N. falsa was proposed for a species from the Black Sea, it has not been recorded there (Surugiu, 2005; Şahin & Çinar, 2012).

Biology and ecology

The reproduction and oogenesis of N. falsa were studied by Daas, Younsi, Daas-Maamcha, and Scaps (2010) and Daas, Younsi, Daas-Maamcha, Gillet, and Scaps (2011). In Algeria, sexual reproduction does not include epitoky, and spawning occurred in August/September. The lack of epitoky contradicts what was regarded as typical by Fauvel (1923), and provides additional indications that more than a single species is present in the Mediterranean.

There are a few data concerning their abundance in Algeria: mean density was 11.27 ind.m-2 with a minimum of 7.83 ind.m-2 and a maximum of 14.50 ind.m-2 (Daas et al., 2011). Regarding their living substrate, N. falsa was found on chains by Fauvel (1913), rocks (Fauvel, 1937; Gillet, 1986, 1988), and hard substrates with red algae (Daas et al., 2010, 2011) but also found on soft substrates such as mud (Fauvel, 1937) and sand (Rullier & Amoureux, 1969). This species also lives on floating debris (Aliani & Molcard, 2003), and algae (Thiel & Guttov, 2005) or as an epibiont on the carapace of loggerhead turtles (Pfaller, Bjorndal, Reich, Williams, & Frick, 2006) and in mussel beds (Çinar et al., 2008). These latter findings could explain a large distribution and if all records really belong to the same biological species, there would be a low genetic variability.

Means for solution?

Despite the fact that N. splendida might have priority over N. falsa, because the former was not well defined, the type specimen originally deposited in Berlin is lost (Hartwich, 1993: 139), and there are no other specimens in Wroclaw (Wiktor, 1980), we must follow Fauvel and regard it as indeterminable. A recent proposal for reinstating N. splendida by Núñez (2004: 375) and Alós et al. (2004: 513), probably by strict priority, should not be followed due to the lack of type material, the incomplete original description, and because it is a junior homonym.

Nevertheless, as indicated by the above analysis, there are three distributional alternatives: 1) N. falsa is an euryhaline species that can be present in the Black Sea and other Mediterranean localities and elsewhere; 2) N. falsa is restricted to the Black Sea, its type locality, and it could be regarded as resembling H. diversicolor; or 3) Two species are involved under the same name with two combinations: 3a) None live in the Black Sea and the two are common in the Mediterranean and adjacent NE Atlantic; and 3b) Non NE-Atlantic records belong to other, perhaps undescribed, species.

The first two alternatives, involving N. falsa living in the Black Sea, must be rejected. As indicated above, recent studies on Black Sea polychaetes have failed to confirm the presence of N. falsa for the type locality, and despite ecological changes since its original description, a local extinction is unlikely. Further, at least von Marenzeller (1874) and Bobretzky (1881) regarded N. falsa as a junior synonym of H. diversicolor which would somehow support the second alternative. However, two different species currently regarded as H. diversicolor were recognized in the Baltic Sea (Audzijonyte, Ovcarenko, Bastrop, & Väinölä, 2008), and three different clades that can be regarded as different species were found in European seas, with a distinct form present in the Black Sea (Virgilio, Fauvelot Constantini, Abbiati, & Backeljau, 2009).

For the third option, and its combinations, it must be reminded that two paragnath patterns in area VI are included under N. splendida - one with 4-5 paragnaths in a diamond, and one with 3 paragnaths in an inverted triangle. The next question would be, because the species name N. splendida must be rejected, how should we call the species living outside the Black Sea, and apparently present throughout at least the Mediterranean Sea? Take N. parallelogramma, which despite the fact no type was deposited, it was described and illustrated in full detail and deserves to be reinstated.

A further issue is what should we call other species resembling N. falsa from other localities? Not an easy answer, either. Regional records should be addressed in each ocean basin and by clarifying the status of type materials and track the sequence of proposed names for each region.

Finally, we concluded that, since our study was based upon publications, not on specimens, we considered that there are four desirable future steps to fully solve the N. falsa problem, and all must be based on the study of topotype or type specimens:

1. Delineate H. diversicolor in the Baltic Sea after which other similar Mediterranean species can be delineated and described.

2. Find a Neapolitan specimen, or series of specimens, to clarify the variation in area VI (3 vs 4 paragnaths), and then proceed to redescribe them as N. parallelogramma Claparède, 1868 (area VI with 4 paragnaths), and propose a neotype.

3. Find out other specimens from the Black Sea, Sevastopol or nearby areas, and in Trieste or Naples for what was regarded by Grube as N. splendida, because its type locality was not indicated. If they consistently have area VI with 3 paragnaths, redescribe it as N. falsa and propose a neotype, because N. splendida is unavailable.

4. Other non-Mediterranean records of N. falsa deserve a similar approach; some regional species names must be reinstated, as N. lucipeta for South Africa, after making a redescription and probably including the proposal for a lectotype, or if different, then full descriptions will be needed for several specimens now included as records for N. falsa.

These steps will be relevant to avoid any future confusion and will certainly depend on collective efforts by several interested scientists. Hope this note will encourage our colleagues to move towards this direction.

ACKNOWLEDGMENTS

Biodiversity Heritage Library, Internet Archive, and la Fédération Française des Sociétés de Sciences Naturelles made available many difficult to find documents. Julia Dunaeva, from the Library of the Zoological Institute and Museum, Russian Academy of Sciences, Saint-Petersburg, provided a fine scan of Rathke’s plates, and Tulio Villalobos one of Ehlers’ plates. The careful reading and recommendations of an anonymous referee and of Daisy Cristina Arroyo Mora helped to improve this contribution.

RESUMEN

¿Qué tan falsa es Nereis falsa (Annelida, Phyllodocida, Nereididae)? Entre los nombres de especies de poliquetos hay muchos problemas taxonómicos y resolver los procedimientos taxonómicos confusos o inadecuados consume mucho tiempo y es muy importante. Nuestro objetivo en esta contribución es analizar cuál es la situación para Nereis falsa de Quatrefages, 1866; fue basada en Nereis pulsatoria? Rathke, 1837 del Mar Negro, y se considera como una especie de amplia distribución. La especie se ha recolectado en diferentes sustratos bénticos e incluso puede hallarse en objetos flotantes o sobre tortugas marinas. Nereis falsa se ha registrado del Mar Mediterráneo, en el Atlántico oriental a lo largo del África, en el Atlántico occidental (Golfo de México, Mar Caribe, Brasil), y en el Índico. Sin embargo, a pesar de haber sido propuesta para una especie del Mar Negro no se ha vuelto a encontrar en el mismo. ¿Cómo conjugar que una especie pueda alcanzar una vasta distribución y faltar en su localidad tipo? Después de un estudio cuidadoso de las publicaciones sobre el tema y con nuestra comprensión de la sistemática de los poliquetos neréididos, clarificamos la situación prevalente al examinar varias especies relacionadas e indicamos algunas cuestiones nomenclaturales. Nuestro análisis indica que hay más de una especie bajo el mismo nombre y en esta contribución, nos enfocamos al problema, proponemos algunas formas para promover la discusión y la acción, y sugerimos algunas actividades de investigación para resolver el problema.

Palabras clave: taxonomía, especies cosmopolitas, especies exóticas, áreas faríngeas, localidades tipo.

REFERENCES

Aliani, S., & Meloni, R. (1999). Dispersal strategies of benthic species and water current variability in the Corsica Channel. Scientia Marina, 63, 137-145. Retrieved from http://scimar.icm.csic.es/scimar/pdf/63/sm63n2137.pdf

Aliani, S., & Molcard, A. (2003). Hitch-hiking on floating marine debris: macrobenthic species in the Western Mediterranean Sea. Hydrobiologia, 503, 59-67. Retrieved from http://link.springer.com/article/10.1023/B:HYDR.0000008480.95045.26

Alós, C., Laborda, J., Núñez, J., Parapar, J., Besteiro, C., Moreira, J., San Martín, G., & Alonso-Zarazaga, M. A. (2004). Nomenclatura: Lista de sinónimos y combinaciones. Fauna Ibérica, 25, 495-521.

Amaral, A. C. Z., Nallin, S. A. H., Steiner, T. M., Forroni, T. O., & Filho, D. G. (2013). Catálogo das Espécies de Annelida Polychaeta do Brasil. Retrieved from http://www.ib.unicamp.br/museu_zoologia/sites/www.ib.unicamp.br.museu_zoologia/files/Cat%C3%A1logo_Polychaeta_Brasil_Amaral_et_al_2013_1a.pdf

Amoureux, L. (1973). Quelques annélides polychètes d’Afrique occidentale et équatoriale. Cahiers ORSTOM, 11, 41-65.

Amoureux, L. (1976). Une intéressante collection de Néréidiens (annélides polychètes) des côtes marocaines du détroit de Gibraltar. Nereis moroccensis, espèce nouvelle pour la Science. Bulletin du Muséum National d’Histoire Naturelle, Paris, 3e série, Zoologie, 258, 370, 337-349.

Andrea, P., & Giancarlo, M. J. (2003). La Fauna del Parco Nazionale del Circeo. Progetto “Parchi in qualità” ovvero “applicazione pilota del Sistema di Gestione Ambientale nelle aree naturali protette”. ENEA, 1-81.

Audouin, J. V., & Milne-Edwards, H. (1832). Classification des Annélides et description de celles qui habitent les côtes de la France. Annales des Sciences Naturelles, Paris, 27, 337-447. Retrieved from http://www.biodiversitylibrary.org/item/29379#page/341/mode/thumb

Audouin, J. V., & Milne-Edwards, H. (1833). Classification des Annélides, et description de celles qui habitent les côtes de la France. Quatrième famille: Néréidiens. Annales des Sciences Naturelles, Paris, 29, 195-269 (issued also as a book, 1834, volume 2, part 1). Retrieved from www.biodiversitylibrary.org/item/29342#page/199/mode/1up

Audzijonyte, A., Ovcarenko, I., Bastrop, R., & Väinölä, R. (2008). Two cryptic species of the Hediste diversicolor group (Polychaeta, Nereididae) in the Baltic Sea, with mithocondrial signatures of different population histories. Marine Biology, 155, 599-612. Retrieved from http://dx.doi.org/10.1007/s00227-008-1055-3

Augener, H. (1918). Polychaeta. Beitrage zur Kenntnis des Meeresfauna West-Afrikas. Herausgegeben von W. Michaelsen. Z.L. Friedrerichsen & Co., Hamburg 2, 67-625, 6 Pls. Retrieved from http://www.biodiversitylibrary.org/page/7172395#page/78/mode/1up

Bobretzky, N. (1881). [Additions to the annelid fauna of the Black Sea]. Zapiski Kievskogo Obshchestva Estestvoispytatelej, 6, 183-212, 2 Pls.

Çinar, M. E., Katağan, T., Koçak, F., Öztürk, B., Ergen, Z., Kocatas, A., Önen, M., Kirkim, F., Bakir, K., Kurt, G., Dağli, E., Açik, S., Doğan, A., & Özcan, T. (2008). Faunal assemblages of the mussel Mytilus galloprovincialis in and around Alsancak Harbour (Izmir Bay, eastern Mediterranean) with special emphasis on alien species. Journal of Marine Systems, 71, 1-17. Retrieved from http://dx.doi.org/10.1016/j.jmarsys.2007.05.004

Claparède, E. (1868). Les Annélides Chétopodes du Golfe de Naples. Mémoires de la Société de Physique et d’Histoire naturelle de Genève, 19, 313-584. Retrieved from www.biodiversitylibrary.org/item/18568#page/11/mode/1up

Claparède, É. (1870). Les annélides chétopodes du Golfe de Naples. Supplément. Mémoires de la Société de physique et d’histoire naturelle de Genève, 20, 365-542. Retrieved from http://www.biodiversitylibrary.org/item/18576#page/13/mode/1up

Conde-Vela, V. M., & Salazar-Vallejo, S. I. (2015). Redescriptions of Nereis oligohalina (Rioja, 1946) and N. garwoodi González-Escalante & Salazar-Vallejo, 2003, and description of N. confusa sp. n. (Annelida, Nereididae). ZooKeys, 518, 15-49. Retrieved from http://dx.doi.org/10.3897/zookeys.518.9564

Daas, T., Younsi, M., Daas-Maamcha, O., Gillet, P., & Scaps, P. (2011). Reproduction, population dynamics and production of Nereis falsa (Nereididae Polychaeta) of the rocky coast of El Kala, National Park, Algeria. Helgoland Marine Research, 65, 165-173. Retrieved from http://dx.doi.org/10.1007/s10152-010-0212-5

Daas, T., Younsi, M., Daas-Maamcha, O., & Scaps, P. (2010). Ovogénèse de Nereis falsa Quatrefages, 1866 de la région de El-Kala, Algérie. Bulletin de la Societé Zoologique de France, 135, 131-141. Retrieved from http://www.snv.jussieu.fr/zoologie/Bulletin/Documents/SZF135(1-2)/Daas135(131-141).pdf

Day, J. H. (1962). Polychaeta from several localities in the western Indian Ocean. Proceedings of the Zoological Society, London, 139, 627-656. Retrieved from http://dx.doi.org/10.1111/j.1469-7998.1962.tb01597.x

Day, J. H. (1967). A Monograph on the Polychaeta of Southern Africa. British Museum (Natural History) Publications, 656, 38+878 pp. Retrieved from www.biodiversitylibrary.org/item/35415#page/7/mode/1up

Day, J. H., & Morgans, J. F. C. (1956). The Ecology of South African estuaries, 7. The Biology of Durban Bay. Annals of the Natal Museum, 13, 259-312.

de Blainville, H. (1825). Néréide, Nereis. Dictionnaire des Sciences Naturelles, 34, 408-455. Retrieved from http://www.biodiversitylibrary.org/item/81569#page/3/mode/1up

de León-González, J. A., Solís-Weiss, V., & Ochoa-Rivera, V. (1999). Nereidids (Polychaeta) from the Caribbean Sea and adjacent Coral Islands of the Southern Gulf of Mexico. Proceedings of the Biological Society of Washington, 112, 667-681. Retrieved from http://www.biodiversitylibrary.org/item/107571

de Quatrefages, A. (1866). Histoire naturelle des Annelés marins et d’eau douce. Annélides et Géphyriens. Librarie Encyclopédique de Roret. Paris, Volume 1, 588 pp. Retrieved from http://books.googleusercontent.com/books/content?req=AKW5QaeDSqZaosjLopsQxZOr4FZ3VA3ZkXKm-TYo9GWOMvtywq0SsiPrazgof9p__Fvl6W1IqQDGCVHzxTlwf8hF6MFUoYU3LlYNyImKWZJBa-SYTwEfOOe2UuRTvL6QrcIOQHJ9-zBES7MoMYz4gRYlfGgU5hEkscPM3LH_sks9zdHvxUxDM3gGPxUxmpzOpQkL4NCRJy58FIGq_-Hd8VEfuW-bubOCUyHMZUG_575XJqb5Wk27fehoEP8Ubs68TsgzJOpmNTDZAbk_4xTsteIDqQLZ1gLy8hCNUH6aT_ZZ1Rg9My6AOAA)

Ehlers, E. (1908). Die Bodensässigen Anneliden aus den Sammlungen der deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer ‚Valdivia‘ 1898-1899, 16, 1-168. Retrieved from www.biodiversitylibrary.org/item/18687

Ehlers, E. (1913). Die Polychaeten-Sammlungen der Deutschen Südpolar-Expedition 1901-1903. Deutsche Südpolar-Expedition, 1901-1903, 13, 397-598, Pls 26-46. Retrieved from www.biodiversitylibrary.org/item/18731#page/461/mode/1up

Fauvel, P. (1913). Quatrième note préliminaire sur les Polychètes provenant des campagnes de l’Hirondelle et de la Princesse-Alice, ou deposées dans le Musée Océanographique de Monaco. Bulletin de l’Institut Océanographique de Monaco, 270, 1-80. Retrieved from www.biodiversitylibrary.org/item/173088#page/256/mode/1up

Fauvel, P. (1916). Annélides polychètes pélagiques provenant des Campagnes de l’Hirondelle et de la Princesse-Alice (1885-1910). Résultats des Campagnes Scientifiques accomplies sur son Yacht par Albert 1er, Prince Souverain de Monaco, 48, 1-152, 9 Pls. Retrieved from www.biodiversitylibrary.org/item/103595#page/11/mode/1up

Fauvel, P. (1919). Annélides polychètes de Madagascar, de Djibouti et du Golfe Persique. Archives de Zoologie Expérimentale et Générale, 58, 315-473.

Fauvel, P. (1923). Polychètes errantes. Faune de France, 5, 1-488. Retrieved from www.faunedefrance.org/bibliotheque/docs/P.FAUVEL%28FdeFr05%29Polychetes-errantes.pdf

Fauvel, P. (1936). Contribution a la faune des annélides polychètes du Maroc. Mémoires de la Société des Sciences Naturelles du Maroc, 43, 1-143. Retrieved from http://www.annelida.net/docs/Fauvel1936_ContrFaunaAnnPolychetesMarocOCR.pdf

Fauvel, P., (1937). Les fonds de pèche près d’Alexandrie, 11. Annélides polychètes. Direction des Recherches des Pêcheries, Notes et Mémoires, Cairo, 19, 1-60. Retrieved from https://www.lifewatchgreece.eu/sites/default/files//article_files/19.%20Annelides%20Polychetes.pdf

Fauvel, P. (1950). Contribution à la faune des annélides polychètes du Sénégal. Bulletin de l’Institut Français d’Afrique Noire, 12, 335-394.

Fauvel, P., & Rullier, F. (1957). Nouvelle contribution à la faune des annélides polychètes du Sénégal. Bulletin de l’Institut Français d’Afrique Noire, 19, 24-97.

Fauvel, P., & Rullier, F. (1959). Contribution à la faune des annélides polychètes du Sénégal et de la Mauritanie. Bulletin de l’Institut Français d’Afrique Noire, 21, 477-533. Retrieved from http://www.mr.refer.org/numweb/IMG/pdf/n02_Flauvel_Rullier.pdf

Faulwetter, S. (2010). Check-list of marine Polychaeta from Greece. Aristotle University of Thessaloniki. Assembled in the framework of the EU FP7 PESI project.

Felder, D. L., & Camp, D. K. (2009). Gulf of Mexico: Origins, Waters, and Biota. Biodiversity. Texas: Texas A&M Press, College Station.

Gillet, P. (1986). Contribution à l’étude écologique des annélides polychètes de l’Estuaire du Bou Regreg (Maroc) (Thèse de doctorat). Université d’Aix Marseille, France.

Gillet, P. (1988). Structure des peuplements intertidaux d’annélides polychètes de l’Estuaire du Bou Regreg (Maroc). Bulletin d’Ecologie, 19, 33-42.

Gillet, P. (1990). Note sur les annélides polychètes du Banc d’Arguin (Mauritanie) et description de Marphysa mauritanica n. sp. Beaufortia, 40(4), 73-84. Retrieved from http://repository.naturalis.nl/document/548721

González-Escalante, L. E., & Salazar-Vallejo, S. I. (2003). A new estuarine species, Nereis garwoodi (Polychaeta: Nereididae) from Bahía Chetumal, Mexican Caribbean coast. Revista de Biología Tropical, 51, 155-164. Retrieved from http://www.scielo.sa.cr/scielo.php?script=sci_arttext&pid=S0034-77442003000100012

Gravina, M. F., Lezzi, M., Bonifazi, A., & Giangrande, A. (2015). The genus Nereis L., 1758 (Polychaeta, Nereididae); state of the art for identification of Mediterranean species. Atti della Società Toscana di Scienze Naturali, Memorie, serie B, 122, 147-164. Retrieved from http://www.stsn.it/AttiB2015/GRAVINA.pdf

Grube, A. E. (1840). Actinien, Echinodermen und Würmer des Adriatischen- und Mittelmeers, nach eigenen Sammlungen beschrieben. J.H. Bon, Königsberg, 92 pp. Retrieved from www.biodiversitylibrary.org/item/40483#page/5/mode/1up

Grube, A. E. (1873). Die Familie der Lycorideen und die Aufstellung fon Gruppen in der Gattung Nereis. Einundfünfzigster Jahres-Bericht der Schlesischen Gesellschaft für vaterländische Cultur Breslau, 51, 56-73. Retrieved from www.biodiversitylibrary.org/item/118508#page/76/mode/1up

Guerra-García, J. M., Gonzáles-Vila, F. J., & García-Gómez, J. C. (2003). Aliphatic hydrocarbon pollution and macrobenthic assemblages in Ceuta harbour: a multivariate approach. Marine Ecology Progress Series, 263, 127-138. Retrieved from http://dx.doi.org/10.3354/meps263127

Hartman, O. (1959). Catalogue of the Polychaetous Annelids of the World, Pt. 1. Allan. Hancock Occasional Papers, 23, 1-353.

Hartwich, G. (1993). Die Polychaeten-Typen des Zoologisches Museums in Berlin. Mitteilungen der Zoologischen Museum, Berlin, 69, 73-154. Retrieved from http://dx.doi.org/10.1002/mmnz.19930690106

ICZN (1999). International Code of Zoological Nomenclature (4th edn). International Trust for Zoological Nomenclature (The Natural History Museum), London, 306 pp. Retrieved from http:// www.iczn.org/iczn/index.jsp

Lamarck, J. B. (1818). Histoire naturelle des Animaux sans Vertèbres, préséntant les caractères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation des principales espèces qui s’y rapportent; precedes d’une Introduction offrant la determination des caracteres essentiels de l`Animal, sa distinction du vegetal et des autres corps naturels, enfin, l’Exposition des Principes fondamentaux de la Zoologie. Paris, Deterville, volume 5, 612 pp. Retrieved from www.biodiversitylibrary.org/bibliography/12712

Miloslavich, P., Díaz, J. M., Klein, E., Alvarado, J. J., Díaz, C., Gobin, J., Escobar-Briones, E., Cruz-Motta, J. J, Weil, E., Cortés, J., Bastidas, A. C., Robertson, R., Zapata, F., Martín, A., Castillo, J., Kazandjian, A., & Ortiz, M. (2010). Marine Biodiversity in the Caribbean: Regional Estimates and Distribution Patterns. PLoS ONE 5(8), e11916. Retrieved from http://dx.doi.org/10.1371/journal.pone.0011916

Núñez, J. (2004). Familia Nereididae Savigny, 1822. Fauna Ibérica, 25, 293-390.

Pfaller, J. B., Bjorndal, K. A., Reich, K. J., Williams, K. L., & Frick, M. G. (2006). Distribution patterns of epibionts on the carapace of loggerhead turtles, Caretta caretta. Journal of the Marine Biological Association of the United Kingdom. Marine Biodiversity Records, 1(e36), 1-4. Retrieved from http://dx.doi.org/10.1017/S1755267206003812

Ramsay, L.N.G. (1914). Report on the scientific results of the voyage of S.Y. “Scotia” during the years 1902, 1903 and 1904, under the leadership of William S. Bruce, 8. Polychaeta of the family Nereidae, collected by the Scottish National Antarctic Expedition. Transactions of the Royal Society of Edinburgh, 50, 41-48. Retrieved from http://www.biodiversitylibrary.org/item/15641

Rathke, H. (1837). Zur Fauna der Krym. Mémoires de l’Academie Impériale des Sciences de St.Pétersbour (Mémoirs des Savants Étrangers), 3, 290-454, Pls 1-10. Retrieved from archive.org/details/zurfaunaderkryme00rath

Rullier, F., & Amoureux, L. (1969). Nouvelle contribution à l’étude de la faune des annélides polychètes du Maroc. Bulletin de la Société des Sciences Naturelles et Physiques du Maroc, 49, 109-142.

Şahin, G. K., & Çinar, M. E. (2012). A check-list of polychaete species (Annelida: Polychaeta) from the Black Sea. Journal of the Black Sea/Mediterranean Environment, 18, 10-48. Retrieved from http://www.blackmeditjournal.org/pdf/vol18no1pdf2.pdf

Salazar-Vallejo, S. I., Carrera-Parra, L. F., González, N. E., & Salazar-González, S. A. (2014). Biota portuaria y taxonomía. In A. Low-Pfeng, P. A. Quijón, & E. M. Peters-Recagno (Eds.), Especies Invasoras Acuática: Casos de Estudio en Ecosistemas de México. México, Semarnat, INECC, & Univ. Prince Edward Island. Retrieved from http://www2.inecc.gob.mx/publicaciones/consultaPublicacion.html?id_pub=713

Salazar-Vallejo, S. I., & Jiménez-Cueto, M. S. (1997). Neréididos (Polychaeta) del Caribe mexicano con una clave para las especies del Gran Caribe. Revista de Biología Tropical, 44, 361-377. Retrieved from http://www.ots.ac.cr/rbt/attachments/volumes/vol44-3B/07-Salazar-Nereididos.pdf

Sánchez-Moyano, J. E., García-Adiego, E. M., Estacio, F. J., & García-Gómez, J. C. (2001). Influence of the density of Caulerpa prolifera (Chlorophyta) on the composition of the macrofauna in a meadow in Algeciras Bay (Southern Spain). Ciencias Marinas, 27, 47-71. Retrieved from http://www.redalyc.org/pdf/480/48027104.pdf

Savigny, J. C. (1822). Système des annélides, principalement de celles des côtes de l’Égypte et de la Syrie, offrant les caractères tant distinctifs que naturels des Ordres, Familles et Genres, avec la description des Espèces. Description de l’Égypte, Histoire Naturelle, 1(3), 1-128. Retrieved from biodiversitylibrary.org/page/41329897#page/425/mode/1up

Sourie, R. (1954). Contribution à l’étude écologique des côtes rocheuses du Sénégal. Mémoire de l’Institut Français d’Afrique Noire, 38, 1-342 p.

Surugiu, V. (2005). Inventory of inshore polychaetes from the Romanian coast (Black Sea). Mediterranean Marine Science, 6, 51-73. Retrieved from http://dx.doi.org/10.12681/mms.193

Taylor, J. L. (1984). Family Nereididae Johnston, 1845. In J. M. Uebelacker & P. G. Johnson (Eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico (pp. 31.1-31.42). Final report to the Minerals Management Service, contract 14-12-001-29091. Barry A. Vittor & Associates, Mobile, Alabama.

Thiel, M., & Guttov, L. (2005). The ecology of rafting in the marine environment, 2. The rafting organisms and community. Oceanography and Marine Biology, Annual Review, 43, 279-418. Retrieved from https://epic.awi.de/11613/1/Thi2005a.pdf

Virgilio, M., Fauvelot, C., Constantini, F., Abbiati, M., & Backeljau, T. (2009). Phylogeography of the common ragworm Hediste diversicolor (Polychaeta: Nereididae) reveals cryptic diversity and multiple colonization events across its distribution. Molecular Ecology, 18, 1980-1194. Retrieved from http://dx.doi.org/10.1111/j.1365-294X.2009.04170.x

Von Marenzeller, E. (1874). Zur Kenntnis der adriatischen Anneliden. Sitzungsberichte der Kaiserliche Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Klasse, 69, 407-482. Retrieved from archive.org/details/cbarchive_109423_zurkenntnisderadriatischenanne1874

Wiktor, J. (1980). Type-specimens of Annelida Polychaeta in the Museum of Natural History of the Wroclaw University. Annales Zoologici, 35, 267-283.

WoRMS (2016). Nereis falsa. In G. Read, & K. Fauchald (Ed.), World Polychaeta Database. marinespecies.org/aphia.php?p=taxdetails&id=130397 on 2016-01-22