Six species of Pectinariidae are known from Northern Europe, but their classification is controversial. In the literature, these species are regarded either as the genus PettaMalmgren, 1866 or the genus PectinariaLamarck, 1818 with subgenera: AmphicteneSavigny, 1822, PectinariaMalmgren, 1866, CistenidesMalmgren, 1866 and LagisMalmgren, 1866, or each subgenus is considered a valid genus. Both alternatives are currently correct, but for the purpose of this paper each is considered a separate genus, as recommended by Hutchings & Peart (2002) and Hutchings et al. (2020). Pectinariidae include five genera with 66 species worldwide (Read & Fauchald 2020) and six species in Northeastern Europe: Petta pusillaMalmgren, 1866, Amphictene auricoma (Müller, 1776), Pectinaria belgica (Pallas, 1766), Cistenides granulata (L., 1767), C. hyperboreaMalmgren, 1866 and Lagis koreniMalmgren, 1866. Petta pusilla and C. hyperborea are not discussed in detail in this paper as there is no problem with the diagnosis or the species has a northern distribution, respectively. However, the diagnosis of the remaining species was inconsistent throughout the literature. Characteristics of genera and species were quite confusing and resulted in misidentifications. This problem was likely started by Fauchald (1977) and has been consequently replicated in a number of more recent studies (e.g. Hutchings & Peart 2002; Gil 2011). Malmgren (1866) described the above-mentioned species in detail and gave generic diagnosis of the five genera. As more species were described, it became apparent that some characters within the generic diagnosis provided by Malmgren (1866) were more variable. When Fauchald (1977) gave the new generic diagnosis, he may have overlooked those described by Malmgren (1866) and Annenkova (1929), for in the latter both Cistenides species did not fit within Cistenides described by Fauchald (1977). Until recently, only two species of Pectinariidae were known to occur in the Netherlands: L. koreni and A. auricoma, both easily distinguishable. However, the discovery of two large specimens on the Dutch Oyster Grounds, whose characters do not resemble those of the two known species, revealed that another species occurs in the Dutch North Sea. For these specimens, it proved impossible to determine whether they were P. belgica or Cistenides when using Fauchald (1977), Hutchings & Peart (2002) or Gil (2011). With the use of other keys, such as Hartman-Schröder (1996), Holthe (1986), Jirkov & Leontovich (2013), Jirkov (2001) and Malmgren (1866), it became clear that the Dutch specimens must belong to P. belgica (Pallas). However, many mistakes have been made in the past with respect to P. belgica and L. koreni, resulting in mixing of characteristics and a number of false records. This paper focuses on the historical and current state of knowledge about Dutch Pectinariidae.
Materials and methods
The records of L. koreni, A. auricoma, and the 2018 record of P. belgica from the Dutch North Sea are based on the material collected in the period of 1991–2019 by Rijkswaterstaat, Eurofins AquaSense (formerly Grontmij and AquaSense), Ecosub, Bureau Waardenburg and NIOZ. During the first years (1991–1994), 25 sites located along five transects perpendicular to the Dutch coast were sampled. This method was revised and, starting from 1995, it has been decided to collect single samples at 100 sites scattered across the DCS (Dutch Continental Shelf). Each site was sampled for benthic fauna and sediment using a Reineck box corer (0.078 m2). Benthic samples were stained with rose bengal, sorted and, whenever possible, the collected specimens were identified to the species level. P. belgica collected by NIOZ was sampled with a Triple-D dredge (Bergman & van Santbrink 1994) as part of a sampling campaign in 2019, when the Frisian Front was monitored. Dredge samples (20 cm × 20 cm) were collected over a stretch of exactly 100 m, resulting in a sample of 20 m2. A sample was sieved first in a net behind the dredge (mesh size 8 mm), and then with a sieve on board (mesh size 5 mm). Everything that was collected was sorted and identified immediately on board. Species that were questionable or, for example, rare and sought for a reference collection (like Pectinaria belgica) were preserved in 4% formaldehyde and brought to the laboratory for additional examination.
Basic description of Pectinariidae
Pectinariidae, “trumpet worms” or “ice-cream-cone worms” (in Dutch: Goudkammetjes), are easily recognizable by their distinctive anterior golden paleae (Fig. 1). They live in a fragile and thin cone-shaped tube consisting of sand, small shells, or other small particles. The diagnosis of the family is presented in Rouse & Pleijel (2002), Zhang & Hutchings (2019) and Hutchings et al. (2020), and some relevant characters of the European species are reproduced below. The anterior-dorsal part of the head (operculum) is marked by two bundles of large golden paleae that extend forward. The dorsal edge of the operculum, the opercular rim, can be either serrated (Amphictene) or smooth (other genera). The thoracic region bears two pairs of branchiae and three segments with notopodial hairs only. The abdominal region has 12 or more segments with both notopodial hairs and transverse rows of neuropodial uncini (Fig. 2). The total number of segments with notochaetae and neurochaetae has been used by Malmgren (1866) as a generic character (Table 1), but with the description of more recent species this number proved to vary within a genus and to be stable within a species. At least in Dutch specimens of A. auricoma and L. koreni, the number of segments with notochaetae and neurochaetae remains the same as individuals of the species grow (T. van Haaren, unpublished observations). The scaphe, the posterior part of the body, is usually well demarcated from the abdomen, except in Petta. The scaphe segments are achaetous, though there may be up to 20 or more pairs of stout spine-like chaetae, called scaphal hooks, around the area where the scaphe joins the abdomen. In most 19th-century taxonomic papers, only the number of paleae, notochaetae and neurochaetae, the shape of the opercular rim (smooth or serrated) or cephalic veil (with or without cirri) are reported (e.g. Savigny 1822; Grube 1851; Johnston 1865; Levinsen 1884). Sometimes the shape of the sand tube is also reported (curved or straight) (Savigny 1822; Johnston 1865; Malmgren 1866 and others), but in the author's opinion it has no diagnostic value. The shape of the tube depends on the size, and often the thinner, fragile end section breaks off, “changing” the shape from a curved to a straight tube. Other research indicates that one has to pay attention to several other characteristics, such as the number of vertical rows of major teeth on the uncinus (e.g. Annenkova 1929; Fauvel 1936; McIntosh 1922; Hutchings & Peart 2002), which was mostly neglected in the 18th and 19th century papers. Nowadays, it is used in the generic diagnosis and regarded as a diagnostic feature to distinguish Cistenides from Pectinaria (Fauchald 1977; Hartmann-Schroder 1996; Hutchings & Peart 2002; Hutchings et al. 2020).
Figure 1
Basic morphology of Pectinariidae (L. koreni); outside and inside the tube (photo courtesy of Hans Hillewaert). The white line represents 1 cm.
Figure 2
Details of the abdominal segment in Lagis koreni showing notopodial hairs and neuropodial uncini (photo courtesy of T. van Haaren)
Historical investigation is necessary to determine whether P. belgica has ever been found in the Netherlands before. It is not unlikely that mistakes have been made in the past. After Pallas (1766) described the first two species, including P. belgica, the number of Pectinariidae species increased. A literature review for descriptions of Pectinariidae species in Europe and a comparison with the original species of Pallas (1766) is important to determine the actual distribution of this species.
Peter Simon Pallas (1741–1811) was born in Berlin and studied at the universities of Berlin, Halle, Göttingen, Leiden and London. From 1762 to 1766 he stayed in ‘s-Gravenhage (The Hague), after which he completed Miscellanea zoologica (1766) where he described several vertebrate and invertebrate species new to science, including Nereis cylindraria var. belgica. Several of his works, including Miscellanea zoologica, have been translated by P. Boddaert from Latin into Dutch (Boddaert 1770). In 1767, Pallas became a professor at the St. Petersburg Academy of Science (Russian Kunstkamera at the Academy of Sciences), where he led several expeditions in Russia. He finished his work in Crimea and moved to Berlin, where he died in 1811. His material on Nereis cylindraria var. belgica is untraceable, or at least not to be found in the St. Petersburg Museum (N. Slepkova, personal communications) or the Naturalis Biodiversity Center (Leiden).
In Miscellanea zoologica, Pallas describes two varieties of Nereis cylindraria, N. cylindraria var. capensis (from the Cape of Good Hope, South Africa) and N. cylindraria var. belgica, based on specimens washed up on Dutch beaches. As Pallas was staying in The Hague at that time, his material of var. belgica was most likely from the beaches of Scheveningen. In that period, there was only one main road from Den Haag to Scheveningen (Scheveningse weg), leading to the Oude Kerk (English: old church), so it can be assumed that his material originates from the beach near the Oude Kerk, Scheveningen (52°6′28″N; 4°16′12″E).
Both varieties can be attributed to Pectinariidae. N. cylindraria var. capensis (as “Kaapsche Verscheidenheid”, the corresponding Dutch vernacular name of this species in Boddaert 1770) differs from N. cylindraria var. belgica (“Hollandsche verscheidenheid” in Boddaert 1770) in a crenulated dorsal edge behind the head, while in belgica this edge is smooth (Pallas 1766; Boddaert 1770). Furthermore, it appears from the original description of N. cylindraria var. capensis that N. cylindraria var. capensis has 13 neurochaetae. In the following description of N. cylindraria var. belgica, Pallas (1766) writes that this species is the same as the previous one, except for the size (three times smaller; the tube is up to 2.5 duim in N. cylindraria var. belgica and up to 5 duim in N. cylindraria var. capensis; 1 duim = 1 inch), the dorsal edge (crenulated in N. cylindraria var. capensis and smooth in N. cylindraria var. belgica), and the scaphe (long in N. cylindraria var. capensis and short in N. cylindraria var. belgica). Several other details can be derived from the plates. It is clear from Figure 5, Plate IX in Pallas (or Plate IV in Boddaert 1770) that var. belgica has 14 or 15 notochaetal bundles posterior to the gills (Fig. 3). The number of neurochaetae is impossible to determine and one can only guess whether Pallas ever saw a difference in the number of neurochaetae between N. cylindraria var. capensis and N. cylindraria var. belgica. It cannot be inferred from that paper whether var. belgica also has 13 neurochaetae, as does var. capensis.
Figure 3
Nereis cylindraria var. belgica, Fig. 3 (tube) and Fig. 5 (animal) in Pallas 1766 (source: Iconographia Zoologica)
There is also the question of the etymology of the species epithet “belgica”. The Netherlands was formerly divided into two regions, the northern region and the southern region. The northern region, known as the “united provinces of the Netherlands” or in Latin “Belgium Foederatum”, was a federal republic from 1588 until the Batavian revolution in 1795. It largely covered the territory of the present-day Netherlands. The southern region covered the territory of today's Belgium and was occupied by Spain (1585–1713) and Austria (1714–1795). The latter period is also known as the “Austrian Netherlands” or in Latin “Belgium austriacum”. Thus, when Pallas stayed in The Hague (1762–1766), he actually was in the northern region or Belgium Foederatum, which explains the name belgica”. Papers from before 1795 in which Belgium or Belgii is reported refer either to the present-day Netherlands or to Belgium, e.g. Linné (1788, p. 3749) says “habitat ad Belgii littora”. Later reports of P. belgica from Belgium (e.g. Southern 1910, 1914; McIntosh 1922) are either due to this geographical misunderstanding of the name Belgium/Belgii, or to the assumption that Malmgren described the same species as that described by Pallas.
In 1767, Linné describes Sabella granulata, but his description of the species [“Lives in a curved dark colored sandy tube (…) in deeper parts of the North Atlantic Ocean”] is very scanty.
Houttuyn (1771) provides a Dutch translation of Linnaeus’ S. granulata (species 809 on pages 611–612) saying that the tube is composed of brown sandy grains and has dark color. Houttuyn (1771) also writes that such tubes of sand and shell debris can also be found on Dutch beaches. According to the current state of knowledge, however, it appears that these tubes, containing shell remains, may represent tubes of Lanice conchilega (Pallas, 1766), another common species on the Dutch coast. Furthermore, species with features of the current concept of C. granulata have never been found in the (Dutch) North Sea. It is therefore most likely a common mistake in 18th and 19th century papers to name each species with a sandy tube S. granulata, P. belgica, or perhaps L. conchilega. Malmgren (1866) wrote a more detailed description, also adding that the paleae of P. granulata lack a thin tip (“apice haud tenui attenuatto”), a character now considered diagnostic.
The description of Amphitrite auricoma in Müller (1776) is very scanty. He describes the species as having golden fans of stiff bristles on both sides of the head and living in fragile sand tubes. However, in 1779 and 1788, Müller gave a more detailed description and a very nice color plate (Plate XXVI, Fig. 5) clearly showing the serrated edge of the opercular rim, a characteristic feature of this species at that time. Nowadays, this serrated edge is regarded as a generic feature of the genus Amphictene.
It is also interesting that Müller creates a new name for Sabella granulata L., as he lists it under the synonymy of A. auricoma. Since Fabricius was Müller's student and they were in contact, Müller published some data from Fabricius before the famous book of this author on the Greenland fauna was published (Fabricius 1780). It seems that Fabricius did indeed find the species in Greenland, and it is possible that Müller is referring to the findings of Fabricius in Greenland. However, Müller (1766) does not make any direct reference to the work of Fabricius, except maybe the use of the Greenlandic (vernacular) name (Imab Polia). As Fabricius was a linguist, he used to name each species with a Greenlandic vernacular name as well, and most likely Müller copied that in his 1776 book. The extensive description by Fabricius (1780) in his book “Fauna groenlandica” on A. auricoma is therefore interesting. Fabricius writes “Segmenta 3 sequentia utrinque papillo conico setis paucis aureis armato, sursum retroque tendente; 12 sequentia similia, fed fetis pluribus et longioribus (…)”; translated by the author as: “3 segments papillose with golden hairs, 12 others more or less similar, but with more and longer hairs”. The presence of 15 pairs of notochaetae suggests that he actually referred to L. koreni, the only NW European species with this character. Grube (1871) suggests that Fabricius’ A. auricoma is synonymous with P. granulata.
In 1866, a revision on Scandinavian polychaetes was published by Anders Johan Malmgren. In his chapter on the family Amphictenea, he gives a redescription of P. belgica (Pallas) and A. auricoma (Müller), describes the genus Petta and the type species Petta pusilla, the genus Lagis with the type species L. koreni and the genus Cistenides with the type species Sabella granulata L. and the new species C. hyperborea. The diagnostic features of these species are presented in Table 1.
These features of the European (read: Scandinavian) species were generally followed by subsequent authors, but the number of paleae is more variable. Malmgren's material of P. belgica from Bohuslän (Swedish west coast along the Kattegat) is still available for study and has been assigned as the neotype of P. belgica (Pallas) by Nielsen et al. (1977). It is present in the Swedish Museum of Natural History, Invertebrate Zoology, type no. 3138. The nomenclatural problem of these European Pectinariidae, and especially L. koreni vs P. belgica, is extremely complex and results from the accumulation of successive errors and misinterpretations. The main problem was that Malmgren assigned the name belgica to an actually new species, and created a new species (L. koreni) for what actually was belgica Pallas. Morphology, ecology and distribution data clearly show that Nereis cylindraria var. belgica Pallas must be identical to L. koreni Malmgren. It is most likely that Pallas was collecting Pectinariidae specimens with 15 segments with notochaetae and 12 segments with neurochaetae, as this is the only species on the west coast of the Netherlands (see the next paragraph). On the other hand, faunistic changes over the last 250 years cannot be entirely ruled out. Consequently, Malmgren's material of P. belgica (17 notochaetae, 13 neurochaetae) cannot be identical with Pallas’ material of Nereis cylindraria var. belgica. This matter was addressed by Lucas & Holthuis (1975). However, returning to the correct meaning of belgica would mean that many contemporary papers would have to change the meaning and create a third name, while suppressing koreni (as a junior synonym of belgica). To avoid this inconvenience, the committee decided, at the request of Nielsen et al. (1977), to continue using the names sensu Malmgren. This meant announcing the neotype belgica. There were objections to codify the misidentification (Holthuis 1978), but the request was supported by a large number of marine biologists (Nielsen & Kirkegaard 1978). The final decision was issued by the ICZN in 1982 (Opinion 1225), putting an end to the discussion. The committee ruled that Malmgren's species of Pectinaria belgica Pallas from Bohuslän was to be regarded as a valid type, with P. belgica retaining the original author – Pallas. However, it was overlooked at that time that a candidate name for the new species – misidentified by Malmgren as P. belgica – was actually available. French Professor Giard (1899, 1913) attempted to solve the problem earlier by introducing Pectinaria septentrionalis as an alternative to Malmgren's P. belgica and preserving the name P. belgica as originally intended by Pallas. Since then the name has been overlooked (or neglected), even by Lucas & Holthuis (1975), and consequently also by the committee. Giard (1899: p. 489) and Giard (1913: p. 122) writes (translated by the author from French): “I have preserved the name Pectinaria belgica assigned by Bouchard-Chantereaux for an Amphictenien so common on our sandy beaches. It is indeed the species that Pallas had before his eyes when describing Nereis cylindrica var. belgica. It is the same species that Malmgren has since called Lagis koreni, reserving the name of Pectinaria belgica Pallas for a species of the Arctic seas whose presence on the coasts of France and Belgium seems to us very doubtful. To avoid any confusion, it would be advisable to restore the name of Pectinaria belgica Pallas to the species from the coasts of Holland, Belgium and France and to give the name Pectinaria septentrionalis to the species from Norwège, which does not penetrate the Baltic”. This available name P. septentrionalisGiard, 1899 does not change the validity of both L. koreni and P. belgica. The committee ruled to maintain the prevailing use of L. koreniMalmgren, 1866 (type locality Finnmark, Norway) and P. belgica (Pallas, 1766) (type locality Bohuslän, Sweden) as valid species.
History of Pectinariidae from the Dutch west coast
There are numerous reports on Pectinariidae washed up on Dutch shores, especially after SE storms and in spring. They mostly report empty tubes, which cannot be linked to any Pectinariid species, but sometimes they provide some information on the worm itself.
17th century: Dutch biologist Jan Swammerdam (1637–1680) worked until his death on his major work, published posthumously in 1737–1738 by Leiden University Professor Herman Boerhaave as Bijbel der natuure. In Volume 2 (1738), Jan Swammerdam describes a trumpet worm (as ‘pijpken’) from material he found on the beaches of the North Sea. This means that Pectinariidae (in this case most likely L. koreni) were already washed up on Dutch beaches in the second half of the 17th century. The material he found, although circumstantial, probably originated from a beach in Zuid-Holland, as he probably made his observations while studying at Leiden University (1661–1667).
1762: In Natuurkundige Uitspanningen, Baster writes about Amphitrite (translated from Dutch by the author): “When walking along the beaches on the isle of Schouwen in February and March, one may notice a number of small ‘blisters’ filled with pink or reddish-brown eggs. From these eggs ‘zand-aaltjes’ hatch, Amphitrite, which hide in the sand immediately after hatching” (Baster 1762, p. 44). It is impossible to tell from his figure (Table V, Figure 1) what species this might be.
1766: Pallas describes Nereis cylindraria var. belgica, which he collected on the beach at Scheveningen (see above).
1826:Bennet & van Olivier (1826) report three species of tube-dwelling worms from the Dutch beaches, Amphitrite auricoma (p. 81), Sabella chrysodon (p. 159) and Sabella belgica (p. 160). The current status of the A. auricoma specimen they reported is not certain, but it is definitely a Pectinariid [translated from Dutch by the author: “slanting head, with 4 palps and with 2 transversely arranged, comb-shaped and gold-colored ‘leafs’ that glisten very strongly. Body tapering and a row of brush-shaped nipples on both sides (…) second and third segment with four comb-shaped gills. Length 2–2.5 inches. Tube of fine sand.”]. Since their A. auricoma is reported from Dutch beaches, this may be the current concept of L. koreni. Their S. chrysodon belongs to the current concept of Lanice conchilega (Pallas). Nereis conchilega is on their list of synonyms and they describe the typical habitus of L. conchilega (e.g. tube of sand and shell debris, flexible). Their Sabella belgica is definitely the current concept of L. koreni, as they describe it as having a sandy tube and 12 feet. They also write (translated from Dutch by the author): “They live along the Dutch western beaches along the North Sea, especially at Scheveningen, were they can be found among shrimps”. The last sentence, at least, is noteworthy, as this is exactly what Pallas wrote – “(…) among shrimps”. It is worth noting that Bennet & van Olivier (1826: p. 160) gave Sabella belgica the vernacular name “Belgische zandkoker”, while the species does not even occur in Belgium. Again a misinterpretation of the species epithet belgica.
1851:Maitland (1851: p. 204) reports Amphitrite auricoma from the Netherlands with a long list of synonyms, including Sabella granulata L. and Nereis cylindraria Pallas, and he also refers to the record by Bennet & van Olivier (1826). The species is reported from Schouwen (Baster) and Scheveningen (Bennet & Olivier). Baster's record from Schouwen is probably not a Pectinariid (see above) and the latter record may still refer to the original Pallas record. Maitland also writes (translated from Dutch by the author): “sometimes in spring, the beaches of our coasts are covered with tubes of these animals”. From his very short Latin description it is impossible to tell which Pectinariid he actually described.
1866: In a note on Dutch caddisflies (Fam. Phryganeidae), van Hasselt (1866) compares the beautiful cases with sandy tubes of saltwater annelids he often found on Dutch beaches. He liked the paleae (“gulden lokken”) of our “Amphitrite Belgica (Amphictena, ook Pectinaria, auricoma)” (Hasselt, 1866, p. 211). This quotation shows that Hasselt (1866) considered Amphitrite belgica and Pectinaria auricoma to be the same species.
1898: In Horst's checklist of Dutch Polychaetes, two species are reported on pages 26–27, Pectinaria belgica Pallas and Pectinaria auricoma Müll. He writes about P. belgica (translated from Dutch by the author): “The ‘belgische zandkoker’ is very common along our coast and sometimes can be found in large numbers on our beaches”. However, in his chapter on Pectinaria auricoma he writes that “Hornell (Report upon the fauna of Liverpool bay) claims that Malmgren's description of the species having 17 bundles of hairs and 13 comb-brushes is false, and that there are only 15 pairs of hair bundles and 12 pairs of the latter are present, like in P. belgica. This is, however, not the case. P. auricoma and belgica have 17 feet, but in the latter species, the two last segments are without bristles, unlike auricoma”. This is interesting as we may conclude from this that Horst's and Hornell's belgica are in fact koreni. Horst's P. belgica can be assigned to L. koreni for he describes it as having 15 pairs of notochaetae and 12 neurochaetae. His P. auricoma has an extra pair of neurochaetae, so this one probably belongs to A. auricoma (see also Horst 1922). P. belgica is reported to be common along the Dutch coast and can sometimes be found in large numbers on the beach. P. auricoma is rare and was only found in dredge samples from the Zuiderzee at Wieringen.
1913: In a popular book on beach species by Heinsius & Jaspers (1913, p. 7 & Figure 1F), Pectinaria belgica is reported as very common, but from the drawings made by these authors it is impossible to tell which species of Pectinariidae they actually described. They even made an erroneous observation by calling the paleae “comb-shaped gills”.
1922: In the chapter on polychaetes in the Zuiderzee (Horst 1922, p. 274), Pectinaria auricoma is reported from Wieringen again. In this paragraph, it is very clear that the specimens described are indeed A. auricoma (translated from Dutch by the author): “This Pectinaria can be distinguished at first sight from Pectinaria belgica, common on our coast, by the slightly bent tube that it inhabits, which in the latter species is entirely straight. Furthermore, in Pectinaria auricoma, the opercular rim behind the paleae is equipped with fringe-like appendages, while it is smooth in Pectinaria belgica (Fig. 17a). There are 17 hair bundles on each side of the body, while the number of rows of combs is 13. The combs (Fig. 17b) are pedunculate, furnished with five to six larger teeth along the free edge and a group of finer teeth and an incision below that. At the base of the tail lobe (scaphe), which ends in a thin tip, there is a row of six anal hooks”. He clearly describes the current concept of A. auricoma and confirms the current knowledge about its distribution in the (northern) Dutch North Sea. It was already clear from Horst's earlier publication (Horst 1898) that his Dutch material of belgica is the same as the current concept of L. koreni, with 15 notochaetae and 12 neurochaetae.
1939:Hana (1949, pp. 81–84) describes all that is washed up on the beach after a storm (from the northwest) and spring tide. Based on his story (numerous straight tubes with worms having golden fans of stiff corneous bristles) and a photograph, there is no clear species description to determine which of the Pectinariidae species he may be referring to. An exact copy of his photograph from Hana (1949) is available in the “Nederlands Fotomuseum”. According to the available information, this is the original photograph by Kees Hana taken in April 1939 on the beach at Scheveningen.
1951:Korringa (1951) reports one juvenile specimen of Pectinaria koreni from his culture of experimental oysters in the Eastern Scheldt on 25 June 1941 with an additional comment that “it apparently does not like that habitat”.
1942–1968: In the Dutch magazine “Het Zeepaard”, there are paragraphs entitled “C.S. feuilleton” where enthusiastic biologists could report on interesting marine species they found in the Netherlands. Pectinariidae (or in Dutch “Goudkammetjes”) are reported in large numbers from Dutch beaches. Mass strandings of “Goudkammetjes” were reported on 15 February 1942 (Scheveningen, Katwijk: Bremer 1942), 11 April 1947 (Scheveningen: Leenhouts 1947a), September 1947 (Katwijk-terheijde: Leenhouts 1947b), 9 October 1955 (Toorne, De Beer, Hoek van Holland, Scheveningen, Wassenaarse slag, Katwijk, Bloemendaal, IJmuiden: Swennen 1955), 14 January 1958 (Hoek van Holland : Wolff 1966), 6 February 1960 (Katwijk-Wassenaar: Spaink 1960), 17 November 1963 (Tholen: Compaan 1965), February–March 1965 (Scheveningen, Terheide: Compaan 1965) and 20 January 1968 (between Noordwijk and Noordwijkerhout: Heerebout 1968). These mass strandings are either reported as Pectinaria sp., Pectinaria belgica, Pectinaria koreni or Pectinaria auricoma. This seems strange as Bloklander (1947) had already written that specimens from Dutch beaches were in fact Pectinaria koreni, not Pectinaria belgica. For some reason, however, most reporters neglected this information until Wolff (1970) clarified the issue (again). For example, Wolff (1970) writes (translated from Dutch by the author): “If one now tries to name the ‘Goudkammetjes’ from the Dutch beach using the most common identification keys for polychaetes (Fauvel, 1927; Friedrich, 1940), one always arrives at one and the same species: Pectinaria koreni Malmgren 1865. Also all material from Zeeland and from the North Sea off the coast at the Delta area belongs, as far as I have seen, to this species”. As Fauvel (1927) and Friedrich (1938, Wolff incorrectly indicated 1940) are correct according to the current knowledge about European Pectinariidae, one may conclude that all coastal species from the SW region are indeed L. koreni. This is confirmed by more recent findings (this paper).
1957: In his popular book on what can be found on Dutch beaches, Prud’homme van Reine (1957, p. 74) writes about two species of “Goudkammetjes” Pectinaria auricoma (Müller) and Pectinaria koreni (Malmgr.). He gives a very brief description of both species and depicts them on Plate XXII, Figure 7. It is not possible to tell from the descriptions which species is involved (about 15 tentacles on both sides of the mouth in P. auricoma and about 20 in P. koreni; the latter also being slightly larger). It is clear from aforementioned Figure 7 that his P. auricoma has a crenulated dorsal edge, while P. koreni has a smooth one. Apparently Prud’homme van Reine (1957) caught or expected both species on the Dutch beaches, but did not provide any data.
1973:Entrop (1973) writes about Pectinaria koreni washed up on the Dutch beaches. His short description (15 segments, 12 with hook shaped chaetae) leaves no doubt as to their identity: L. koreni.
1999:Prins (1999) reports on Pectinariidae (with original drawings) living in shallow waters along the Dutch coast, washed up on beaches after storms. According to him, the specimens had 15 fleshly protuberances, each ending in a brush of hairs. The original drawings show a smooth opercular rim, 15 notochaetae and a straight tube. It is impossible to determine as to how many neurochaetae they have, but they are also clearly L. koreni.
2002/2019: In two guides on marine species, Leewis (2002) and de Bruyne & Gmelig Meyling (2019) report L. koreni from the beaches along the Dutch coast. Since they both mention 15 segments, they clearly refer to L. koreni.
Dutch fauna: Species list and identification key
Based on the material from the Dutch North Sea, especially the descriptions by Malmgren (1866; see also Table 1), which is widely accepted as the correct diagnosis of the European species, the following list of Dutch species of Pectinariidae can be presented, together with a list of synonyms. The list is strictly based on morphological descriptions from the historical papers. C. granulata L. is not a Dutch species. It is an amphiboreal-Arctic species (Holthe 1977) and should not to be expected in the Dutch North Sea. However, due to misinterpretations of the original and subsequent synonymization with other species of Pectinariidae, this species is also included.
At least Malmgren (1866), Levinsen (1884), Meyer (1912), Wollebaek (1912), Hessle (1917), McIntosh (1922), Fauvel (1927), Friedrich (1938), Holthe (1986), Hartmann-Schröder (1996), Kirkegaard (1996), Böggemann (1997) and Jirkov (2001) gave the correct species diagnosis for the NW European species. However, the generic characters need to be revised. Hutchings & Peart (2002, p. 109 + Table 6) are incorrect in claiming that 16 notochaetae is a generic character of the genus Lagis. This would be impossible as the type species L. koreniMalmgren, 1866 (by monotypy) has only 15 notochaetae. It is therefore worth noting that Icelandic specimens of L. koreni (Parapar et al. 2020) have 16 notochaetal bundles, which may suggest another Lagis species. Furthermore, Zhang & Hutchings (2019) indicated in their generic diagnosis of Lagis that these species should have 13 pairs of neurochaetae, but again Lagis koreni (the type species) have only 12 of them. On the other hand, they also write that the neuropodia are present on segments 8–19, i.e. on 12 segments. Furthermore, Fauchald (1977) and Hutchings & Peart (2002) are incorrect about Cistenides. At least C. hyperborea and C. granulata are species that have 12 neurochaetae, while the aforementioned papers report that they should have 13 neurochaetae. The number of neurochaetae in C. granulata is not reported in the original paper by Linné (1767), but Malmgren (1866) clearly writes that both species of Cistenides have 12 neurochaetae. Thus, it is unclear why Fauchald (1977) considered 13 neurochaetae as a generic character of Cistenides. Also the generic characters of Pectinaria (including belgica) and Petta in Hutchings & Peart (2002) are incorrect, as P. belgica does not have 14 but 13 neurochaetae, and P. pusilla does not have 18 but 17 notochaetae (see Malmgren, 1866). Zhang et al. (2019) corrected the number of notochaetae for Petta to 17. The number of vertical rows of the major teeth on the neuropodial uncini is reportedly of diagnostic value for distinguishing between Cistenides (1 row) and Pectinaria (at least 2 rows; Hutchings & Peart 2002, Hutchings et al. 2020). According to Annenkova (1929), however, C. hyperborea has 2–3 vertical rows, which places this species within Pectinaria. Thus, the number of vertical rows is not used in the key.
The six (possible) North Sea species can be identified using the information from Table 1 and the following simplified key:
Originally as Pectinarea on page 327 (Quatrefages, 1865), but according to Nielsen et al. (1977) this was corrected to Pectinariidae by Hartman (1941). However, it was already Hartman (1938: p. 16) who used the family name Pectinariidae and even Perrier (1897) used both Pectinaridae (p. 1556) and Amphictenidae (p. 1640).
Neotype: Swedish Museum of Natural History (Stockholm), SMNH 3138, designated by Nielsen et al. (1977) (type no. verified by curator Lena Gustavsson, 26 June 2020). Holthe (1986: p. 21) wrote wrong type no. 3133.
Please note that Malmgren made his redescription based on material he obtained from the Swedish Kattegat area (Bohuslän, Väderöarne isle and Kosterfjord).
? 1780 Amphitrite auricoma Müller; Fabricius 1780: pp. 289–291
Oyster Grounds, 54°47′0″N, 4°25′0″E, 9 April 2018, 52 m, Reineck box corer, n = 1, leg. Eurofins AquaSense, det. T. van Haaren. Naturalis Biodiversity Collection RMNH.VER.19971
Oyster Grounds at either 54°8′58″N, 5°7′35″E (appr. 44.3 m) or 53°57′3″N, 4°52′27″E (41.1 m) or 53°51′4″N, 4°52′32″E (39.9 m), last week of October 2019, trawl survey, n = 1, leg. NIOZ (Texel), det. T. van Haaren. Collection NIOZ.
Description of the Dutch species
The specimens match the characters of the genus Pectinaria and the species Pectinaria belgica. The length of the specimen from the 2018 Reineck box corer is 6.8 cm and the one from 2019 trawl survey is 5.2 cm. In the largest specimen, the cephalic veil is free of the operculum and has 18 cirri, while in the smaller specimen the cephalic veil is damaged medially and has 14 visible cirri. The opercular rim entirely smooth. On both sides with 10 (large specimen) or 11 (smaller specimen) paleae with a fine thin tip. Two pairs of branchiae in 3–4, 17 pairs of notopodia in segments 5–21, neuropodia 13 pairs in 8–20. Scaphe distinctly separated from the posterior body, with small rounded lobes on the lateral edge basally and a short and rounded anal (ventral) flap, which is damaged in the smaller specimen. Seven or eight scaphal hooks, most with a broken tip, but a few whole with an almost straight blunt tip, only very slightly bent. Uncini with about two vertical rows of larger teeth.
Although Pallas’ Nereis cylindraria var. belgica is in fact morphologically the same as Malmgren's Lagis koreni, the neotype of P. belgica (Pallas) from Bohuslän, Sweden is designated as the correct type. Consequently, and ironically, P. belgica (Pallas) is a new species for the Netherlands.
Only recently a single, fully grown specimen of P. belgica (Pallas) sensu Malmgren (non Pallas) was found in a sample collected at the Dutch Oyster Grounds (9 April 2018; Figs 4–5). Another single, somewhat smaller, specimen was collected in the last week of October 2019 at the Oyster Grounds during a trawl survey by NIOZ, Texel. The exact location and date is not known, but it was collected at three possible locations (see the material section; L. Kleine Schaars, NIOZ, pers. comm.).
Figure 4
Two species of Pectinariidae, L. koreni and P. belgica (preserved), both from the Dutch North Sea (Oyster Grounds 9 April 2018; photo courtesy of T. van Haaren)
Figure 5
Live material of P. belgica just after being collected at the Oyster Grounds site (9 April 2018)
Figure 6
Distribution map of Pectinaria belgica in the Dutch North Sea. The record from 2018 is marked with the black dot (•) and the single specimen from 2019 is located in one of the open dots (o; exact location unknown)
Figure 7
Distribution map of Amphictene auricoma in the Dutch North Sea based on recent records (1991–2019). The original record of A. auricoma from Wieringen (Horst 1898, 1922) is shown on the map with another symbol (+)
There is no evidence that P. belgica (Pallas) sensu Malmgren was ever found in the Netherlands prior to this study. It seems likely that the northern part of the Dutch North Sea (north of the Frisian Front) may be the southernmost limit of this species. It is therefore unlikely that P. belgica will ever be found washed up on the Dutch coast, also due to the southern current.
Distribution in Europe
Due to the confusion with other species, the distribution of P. belgica is not well determined. Most likely its southernmost limit is around latitude 55 in subtidal sediments, usually at a depth of 40–50 m. It occurs in the North Sea, the western Baltic (Kattegat, Skagerrak) and Norway up to the North Polar Basin (Holthe 1986; Kirkegaard 1969, Jirkov, pers. comm.). According to Kirkegaard (1969), it is common in all deeper parts of the Danish waters, in the Øresund Region to the Copenhagen coast, but he recorded it only once during his investigation (1950–1955; Dogger Bank, 1952; 55°40′N, 6°31′E; 42 m). Parapar et al. (2020) did not report the species from Iceland. The records from France (Fauvel 1927), the Mediterranean Sea, the Black Sea and the Pacific coast of the USA and Canada (Kirkegaard 1969; Holthe 1986; Gil 2011) require examination of the material. According to Holthe (1977), P. belgica is an amphiboreal species.
It does not occur and is not to be expected in the Netherlands.
Distribution in Europe
C. granulata and C. hyperborea are restricted to subtidal sediments in boreal and Arctic regions (north of 65 latitude). According to Igor Jirkov (pers. comm.), C. granulata occurs only in the Barents Sea along the Kola Peninsula, while C. hyperborea is more widely distributed in the North Polar Basin, but only in the Arctic regions. In the boreal regions, from the southwestern Barents Sea and further to the south, C. hyperborea is replaced by A. auricoma. In the White Sea, along the coast of Norway and further to the south, C. granulata is replaced by L. koreni. The original record of C. granulata from the deeper parts of the North Atlantic Ocean (Linné 1767) and Greenland (Malmgren 1866) does seem to fit within the current known distribution of this species. Recently, Parapar et al. (2020) reported both Cistenides species in cold water of the Greenland Sea along the northern border of Iceland. They were unable to locate any P. belgica in Iceland. These studies may indicate that Linné, when describing his Sabella granulata, may have had a specimen with characters of the current concept of Cistenides. According to Holthe (1986), C. granulata and C. hyperborea may be more widely distributed in the Arctic regions up to a depth of about 250 m. However, Zhang et al. (2019) concluded, based on molecular studies, that specimens of Cistenides granulata from the Pacific (British Columbia) and Atlantic (Manitoba) coasts of Canada belong to different species. Nevertheless, it is unlikely that Cistenides will ever be found in the Dutch North Sea.
Type locality: not given. Holthe (1986) says Denmark, but Müller (1776) describes species from Denmark and Norway and the material is probably the same as Linné's S. granulata from Greenland (see above). Type locality is therefore most likely Greenland (see history of Amphictene auricoma).
A. auricoma is more or less restricted to the Northern part of the Dutch North Sea, north of the Frisian Front, with only one recent coastal record near the Wadden Isle Terschelling. The single record from Wieringen originates from Horst (1898, 1922) and even now this would be a fairly rare finding. The Wadden Sea is not its favorite habitat. In the Dutch North Sea, it has its southernmost limit at latitude 53.4.
Distribution in Europe
Amphictene auricoma is more common in the NE Atlantic in subtidal sediments, usually at a depth of more than 20 m to about 500 m (Holthe 1986). It has been recorded from the North Sea, the western Baltic Sea (Kattegat, Skagerrak) and the Norwegian coast up to the North Polar Basin (Holthe 1986) and the Icelandic part of the North Atlantic Ocean (Parapar et al. 2020). In the south, the species has been reported from the European Atlantic coast up to Morocco and the Mediterranean (Claparède 1868; Giard 1913; Fauvel 1927; Fauvel 1936; Holthe 1986; Gil 2011; Le Mao et al. 2019). According to Holthe (1977), A. auricoma is an amphiboreal species.
Lectotype: Swedish Museum of Natural History (Stockholm) SMNH 3137, designated by Nielsen et al. 1977 (type no. verified by curator Lena Gustavsson, 26 June 2020). Holthe (1986: p. 18) wrote wrong type no. 3132.
Type locality: Finnmark, Kalfjord, 50fv (fathoms), 1861; Goës & A.J. Malmgren (Nielsen et al. 1977). Karlfjord is located at about 69°47′29″N, 18°41′52″E.
Note that Malmgren (1866) originally described L. koreni based on the material from three sites along the west coast of Norway (Kalfjord in 1861; Karlsö in 1864 and Stavanger, unknown year) and one site in France (La Rochelle), and the material from Kalfjord was supposedly collected (or published?) by Goës & Ipse 1861, according to Malmgren (1866). Zhang & Hutchings (2020) incorrectly stated that the type locality is Stavanger. All of Malmgren's material is still present in SMNH, including that from France (SMNH176327).
Lagis koreni is widely distributed in the Dutch North Sea. It is very common in the coastal zone and has scattered records north of the Frisian Front. It is rare in the Wadden Sea, the delta area and offshore locations, and has only one record in the North Sea Canal.
Figure 8
Distribution map of Lagis koreni in the Dutch North Sea based on recent records (1991–2019). The original record of Nereis cylindraria var. belgica from Scheveningen (Pallas 1766) is shown on the map with another symbol (+)
Distribution in Europe
L. koreni is widely distributed in the North Sea and the Baltic Sea and most likely restricted to shallower depths (< 50 m). It reaches the North-East Atlantic along the Norwegian coast up to the Polar Basin (Holthe 1986) and Iceland (Parapar et al. 2020). However, based on the description of these Icelandic specimens with 16 notochaetae (incorrectly written as 16 uniramous segments), these L. koreni specimens require further examination. According to Holthe (1977), L. koreni is an Atlantic and boreal species. To the south, L. koreni is reported from the European Atlantic coast up to Namibia and possibly South Africa, the Mediterranean and the Black Sea (Fauvel 1936; Fauvel 1927; Day 1967; Gil 2011). However, Jolly et al. (2005) suggest that L. koreni from Brittany represents another haplotype than specimens from the English Channel, indicating that cryptic diversity exists.
Discussion and conclusions
Since the first description of Pectinaridae species in the 18th century by Pallas (1766: Nereis cylindraria), Linné (1767: Sabella granulata) and Müller (1776: Amphitrite auricoma) there have been numerous misinterpretations of characteristics and assumptions in the original and subsequent descriptions. This has led to numerous mistakes in identification and, consequently, incorrectly reported distribution of the species. All specimens originally viewed by their describers (types) are lost, except perhaps Nereis cylindraria (var. capensis and var. belgica), which may turn up somewhere in some hidden part of the collection (Paris, Leiden or Apeldoorn: see Pieters, 1980). About a century later, Malmgren (1866) created L. koreni and redescribed P. belgica, unaware of the fact that he introduced another error creating an even greater problem. This issue was brought before the committee and it was finally decided that the species descriptions in Malmgren (1866), although false for L. koreni vs P. belgica, were considered correct and the neotype from Bohuslän (Sweden) was assigned. So far so good. The committee created stability and finally every researcher knew, or thought they knew, how to recognize P. belgica and L. koreni. The neotype was not a Dutch specimen, but perhaps that was the best thing to do. The discovery of two specimens on the Dutch continental shelf forced the author to clarify the issue. After all, the species epithet “belgica” was introduced by Pallas (1766) for Dutch Pectinariids specimens. A review of the literature revealed that specimens of P. belgica have never been found in the Netherlands before, and reports of this species in more southern regions (Belgium, France, and English Channel) need to be re-examined. Old reports of P. belgica not only refer to P. belgica, but also to L. koreni and partly to C. granulata and A. auricoma. In NW Europe there are now five genera with six species, three of which are also known from the Netherlands. Pectinariidae, however, need to be revised, not only in terms of species diagnosis but also in terms of generic characters. The generic diagnosis of the five genera has changed over time, and in some recent papers the characters of the type species do not match the generic diagnosis, especially for Lagis and Cistenides. It seems as if the generic diagnosis is based on recent material and new species, neglecting the original descriptions. Furthermore, DNA barcoding is recommended for the European fauna and may prove that more species occur in the North Sea.
Basic morphology of Pectinariidae (L. koreni); outside and inside the tube (photo courtesy of Hans Hillewaert). The white line represents 1 cm.
Figure 2
Details of the abdominal segment in Lagis koreni showing notopodial hairs and neuropodial uncini (photo courtesy of T. van Haaren)
Figure 3
Nereis cylindraria var. belgica, Fig. 3 (tube) and Fig. 5 (animal) in Pallas 1766 (source: Iconographia Zoologica)
Figure 4
Two species of Pectinariidae, L. koreni and P. belgica (preserved), both from the Dutch North Sea (Oyster Grounds 9 April 2018; photo courtesy of T. van Haaren)
Figure 5
Live material of P. belgica just after being collected at the Oyster Grounds site (9 April 2018)
Figure 6
Distribution map of Pectinaria belgica in the Dutch North Sea. The record from 2018 is marked with the black dot (•) and the single specimen from 2019 is located in one of the open dots (o; exact location unknown)
Figure 7
Distribution map of Amphictene auricoma in the Dutch North Sea based on recent records (1991–2019). The original record of A. auricoma from Wieringen (Horst 1898, 1922) is shown on the map with another symbol (+)
Figure 8
Distribution map of Lagis koreni in the Dutch North Sea based on recent records (1991–2019). The original record of Nereis cylindraria var. belgica from Scheveningen (Pallas 1766) is shown on the map with another symbol (+)
Annenkova, N. (1929). Beiträge zur Kenntnis der Polychaeten-Fauna der USSR. I. Fam. Pectinariidae Quatrefages (Amphictenidae Malmgren) und Ampharetidae Malmgren. Annuaire du Musee Zoologique de l'Académie des Sciences de l'URSS 30(3): 477–502.AnnenkovaN.1929Beiträge zur Kenntnis der Polychaeten-Fauna der USSR. I. Fam. Pectinariidae Quatrefages (Amphictenidae Malmgren) und Ampharetidae MalmgrenAnnuaire du Musee Zoologique de l'Académie des Sciences de l'URSS303477502Search in Google Scholar
Baster, J. (1762). Natuurkundige uitspanningen behelze eenige waarneemingen over sommige zee-planten en zee-insecten benevens derzelver zaadhuisjes en eijernesten. J. Bosch, Haarlem. DOI: 10.5962/bhl.title.120202.BasterJ.1762Natuurkundige uitspanningen behelze eenige waarneemingen over sommige zee-planten en zee-insecten benevens derzelver zaadhuisjes en eijernestenJ. BoschHaarlem10.5962/bhl.title.120202Open DOISearch in Google Scholar
Bennet, J.A. & van Olivier, G. (1826). Naamlijst der wormen In Nederland [list of worms from The Netherlands]. Natuurkundige verhandelingen van de Hollandsche Maatschappij der wetenschappen te Haarlem 15 (2): 1–256.BennetJ.A.van OlivierG.1826Naamlijst der wormen In Nederland [list of worms from The Netherlands]Natuurkundige verhandelingen van de Hollandsche Maatschappij der wetenschappen te Haarlem1521256Search in Google Scholar
Bloklander G. (1947). Het Zeegat uit !!. Het Zeepaard 7(6–7): 4–8.BloklanderG.1947Het Zeegat uit !!Het Zeepaard76–748Search in Google Scholar
Boddaert, P. (1770). Dierkundig mengelwerk in het welke de nieuwe of nog duistere zoorten van Dieren. Door nauwkeurige afbeeldingen, beschrijvingen en verhandelingen opgehelderd worden. In het latijn beschreeven door den hooggel. heer P.S. Pallas, M.D. Abraham van Paddenburg & J. van Schoonhoven, Utrecht. DOI: 10.5962/bhl.title.46936.BoddaertP.1770Dierkundig mengelwerk in het welke de nieuwe of nog duistere zoorten van Dieren. Door nauwkeurige afbeeldingen, beschrijvingen en verhandelingen opgehelderd worden. In het latijn beschreeven door den hooggel. heer P.S. Pallas, M.D.Abraham van Paddenburg & J. van SchoonhovenUtrecht10.5962/bhl.title.46936Open DOISearch in Google Scholar
Böggemann, M. (1997). Polychaeten aus der Deutschen Bucht. Courier Forschungsinstitut Senckenberg 202: 1–315.BöggemannM.1997Polychaeten aus der Deutschen BuchtCourier Forschungsinstitut Senckenberg2021315Search in Google Scholar
Bremer, J. (1942). Van de C.S. (Centraal systeem) man. Het Zeepaard 3BremerJ.1942Van de C.S. (Centraal systeem) manHet Zeepaard3Search in Google Scholar
Bruguiere, M. (1791). Tableaux encyclopedique et methodique des trois regnes de la Nature. Contenant L'helminthologie, ou les vers infusoires, les vers intestins, les vers Mollusques, & c. Part 7. Panckoucke, Paris. 180pp + 92 plates. DOI: 10.5962/bhl.title.64072.BruguiereM.1791Tableaux encyclopedique et methodique des trois regnes de la Nature. Contenant L'helminthologie, ou les vers infusoires, les vers intestins, les vers Mollusques, & c. Part 7PanckouckeParis180pp + 92 plates.10.5962/bhl.title.64072Open DOISearch in Google Scholar
Bruyne, R. de & Gmelig Meyling, A. (2019). Basisgids Strandvondsten. KNNV Uitgeverij Zeist.BruyneR. deGmelig MeylingA.2019Basisgids StrandvondstenKNNV Uitgeverij ZeistSearch in Google Scholar
Claparède, R.-É. (1868). Les annelides chétopodes du Golfe de Naples. pp. 499 plus 31 plates, Ramboz et Schuchardt, Genève. DOI: 10.5962/bhl.title.105355.ClaparèdeR.-É.1868Les annelides chétopodes du Golfe de Naplespp. 499 plus 31 plates,Ramboz et SchuchardtGenève10.5962/bhl.title.105355Open DOISearch in Google Scholar
Colgan, N. (1917). A note on Pectinaria koreni from Dublin Bay. The Irish Naturalist 26(8): 136–138ColganN.1917A note on Pectinaria koreni from Dublin BayThe Irish Naturalist268136138Search in Google Scholar
Compaan, H. (1965). CS-Feuilleton. Het Zeepaard 25(2): 25–28CompaanH.1965CS-FeuilletonHet Zeepaard2522528Search in Google Scholar
Cunningham, J.T. & Ramage, G.A. (1888). The Polychaeta Sedentaria of the Firth of Forth. Transactions of the Royal Society of Edinburgh 33(3): 635–684. DOI: 10.1017/s0080456800017488.CunninghamJ.T.RamageG.A.1888The Polychaeta Sedentaria of the Firth of ForthTransactions of the Royal Society of Edinburgh33363568410.1017/s0080456800017488Open DOISearch in Google Scholar
Cuvier G. (1816 [1817]). Le règne animal distribué d'après son organisation, pour servir de base à l'histoire naturelle des animaux et d'introduction à l'anatomie comparée. Tome 1, 540 pp.; Tome 2, 528 pp.; Tome 3, 653 pp.; Tome 4, 255 pp., 15 pl. Deterville, Paris. DOI: 10.5962/bhl.title.41460.CuvierG.1816 [1817]Le règne animal distribué d'après son organisation, pour servir de base à l'histoire naturelle des animaux et d'introduction à l'anatomie comparéeTome 1, 540 pp.; Tome 2, 528 pp.; Tome 3, 653 pp.; Tome 4, 255 pp., 15 pl.DetervilleParis10.5962/bhl.title.41460Open DOISearch in Google Scholar
Day, J. H. (1967). A monograph on the Polychaeta of Southern Africa. Part 2. Sedentaria. British Museum (Natural History), London. pp. 459–842. DOI: 10.5962/bhl.title.8596.DayJ. H.1967A monograph on the Polychaeta of Southern Africa. Part 2. SedentariaBritish Museum (Natural History)London45984210.5962/bhl.title.8596Open DOISearch in Google Scholar
Dicquemare, l'abbé [aka Jacques François Dicquemare/Dicquemarre]. (1779). Ver a fourreau conique. Observations et Mémoires sur la Physique, sur l'Histoire Naturelle et sur les Arts et Métiers 14: 54–56, plate 2 figures 1–12 (following page 88).Dicquemare, l'abbé [aka Jacques François Dicquemare/Dicquemarre]1779Ver a fourreau coniqueObservations et Mémoires sur la Physique, sur l'Histoire Naturelle et sur les Arts et Métiers145456plate 2 figures 1–12 (following page 88).Search in Google Scholar
Donovan E. (1802–1803). The natural history of British shells: including figures and descriptions of all the species hitherto discovered in Great Britain, systematically arranged in the Linnean manner, with scientific and general observations on each. Vol. 4: pl. 109–126 [1802]; pl. 127–144, index 8 pp [1803].DonovanE.1802–1803The natural history of British shells: including figures and descriptions of all the species hitherto discovered in Great Britain, systematically arranged in the Linnean manner, with scientific and general observations on eachVol. 4: pl. 109–126 [1802]; pl. 127–144, index 8 pp [1803].Search in Google Scholar
Donovan, E. (1845). Histoire Naturelle des coquilles D’Angleterre, comprenant la figure et la description de toutes les especes decouvertes jusqu’ici le system de Linné. A. Frank, Paris.DonovanE.1845Histoire Naturelle des coquilles D’Angleterre, comprenant la figure et la description de toutes les especes decouvertes jusqu’ici le system de LinnéA. FrankParisSearch in Google Scholar
Entrop, B. (1973). Goudkammetjes - Pectinaria koreni Malmgr. Vita Marina. Wormen: 17–20.EntropB.1973Goudkammetjes - Pectinaria koreni MalmgrVita Marina. Wormen1720Search in Google Scholar
Fabricius, O. (1780). Fauna Groenlandica, systematice sistens animalia groenlandiae occidentalis hactenus indagata, quoad nomen specificium, triviale, vernaculumque, synonyma auctorum plurimum, descriptionem, locum, victum, generationem, mores, usum capturamque singuli, pro ut detegendi occasio fuit, maximaque parte secundum proprias observationes. Hafniae [= Copenhagen] & Lipsiae [= Leipzig]: J.G. Rothe. xvi + 452 pp., 1 pl. DOI: 10.5962/bhl.title.13489.FabriciusO.1780Fauna Groenlandica, systematice sistens animalia groenlandiae occidentalis hactenus indagata, quoad nomen specificium, triviale, vernaculumque, synonyma auctorum plurimum, descriptionem, locum, victum, generationem, mores, usum capturamque singuli, pro ut detegendi occasio fuit, maximaque parte secundum proprias observationesHafniae [= Copenhagen] & Lipsiae [= Leipzig]J.G. Rothexvi + 452 pp., 1 pl.10.5962/bhl.title.13489Open DOISearch in Google Scholar
Fauchald, K. (1977). The polychaete worms, definitions and keys to the orders, families and genera. Natural History Museum of Los Angeles County: Los Angeles, CA (USA), Science Series. 28: 1–188.FauchaldK.1977The polychaete worms, definitions and keys to the orders, families and generaNatural History Museum of Los Angeles County: Los Angeles, CA (USA), Science Series281188Search in Google Scholar
Fauvel, P. (1927). Polychètes sédentaires. Addenda aux errantes, Arachiannélides, Myzostomaires. Faune de France Volume 16. Paul Lechevalier. Paris. 1–494.FauvelP.1927Polychètes sédentaires. Addenda aux errantes, Arachiannélides, MyzostomairesFaune de France Volume 16Paul LechevalierParis1494Search in Google Scholar
Fauvel, P. (1936). Contribution a la faune des Annélides Polychètes du Maroc. Mémoires de la Société des Sciences Naturelles du Maroc 43: 1–143.FauvelP.1936Contribution a la faune des Annélides Polychètes du MarocMémoires de la Société des Sciences Naturelles du Maroc431143Search in Google Scholar
Friedrich, H. (1938). Polychaeta. Tierwelt der Nord- und Ostsee 32(VI.b): 1–201.FriedrichH.1938PolychaetaTierwelt der Nord- und Ostsee32VI.b1201Search in Google Scholar
Giard, M.A. (1899). Coup d'oeil sur la Faune du Boulonnais. In: Boulogne-Sur-Mer et la region Boulonnaise 1899. Tomes I / ouvrage offert par la ville de Boulogne-sur-Mer aux membres du XXVIIIe congrès de l'Association française pour l'avancement des sciences réunis à Boulogne du 14 au 21 septembre 1899.”: Annelides on pp. 488–491.GiardM.A.1899Coup d'oeil sur la Faune du BoulonnaisIn:Boulogne-Sur-Mer et la region Boulonnaise 1899. Tomes I / ouvrage offert par la ville de Boulogne-sur-Mer aux membres du XXVIIIe congrès de l'Association française pour l'avancement des sciences réunis à Boulogne du 14 au 21 septembre 1899.”: Annelideson488491Search in Google Scholar
Giard, A. (1913). Coup d'oeil sur la Faune du Boulonnais. In: Ouvres diverses. Réunies et rééditees par la soins d’un groupe d’eleves et d’amis. II. Faune et Flore de Wimereaux. Notes diverses de zoologie. Paris. p1. 21–123.GiardA.1913Coup d'oeil sur la Faune du BoulonnaisIn:Ouvres diverses. Réunies et rééditees par la soins d’un groupe d’eleves et d’amis. II. Faune et Flore de WimereauxNotes diverses de zoologieParis121123Search in Google Scholar
Gil, J. (2011). The European Fauna of Annelida Polychaeta, Vol II. PhD Thesis. Faculdade de Ciências, Universidade de Lisboa. Family Pectinariidae, pp. 633–638.GilJ.2011The European Fauna of Annelida PolychaetaIIPhD Thesis.Faculdade de Ciências, Universidade de Lisboa. Family Pectinariidae633638Search in Google Scholar
Grube, A.E. (1850). Die Familien der Anneliden. Archiv für Naturgeschichte 16(1): 249–364.GrubeA.E.1850Die Familien der AnnelidenArchiv für Naturgeschichte161249364Search in Google Scholar
Grube, A.E. (1851). Die familien der Anneliden, mit Angabe ihrer Gattungen und Arten. Ein systematischer Versuch. Berlin, Nicolai'schen Buchhandlung. 1–164. DOI: 10.5962/bhl.title.46818.GrubeA.E.1851Die familien der Anneliden, mit Angabe ihrer Gattungen und Arten. Ein systematischer VersuchBerlinNicolai'schen Buchhandlung116410.5962/bhl.title.46818Open DOISearch in Google Scholar
Grube, A.E. (1871). Bemerkungen über die Amphicteneen und Amphareteen Mgn. Jahres-Bericht der Schlesiche Gesellschaft für vaterländische Cultur, Breslau 48: 68–84.GrubeA.E.1871Bemerkungen über die Amphicteneen und Amphareteen MgnJahres-Bericht der Schlesiche Gesellschaft für vaterländische Cultur, Breslau486884Search in Google Scholar
Hana, K. (1949). Goudkammetjes. De Levende Natuur 52(5): 81–84HanaK.1949GoudkammetjesDe Levende Natuur5258184Search in Google Scholar
Hartman, O (1938). Annotated list of the types of Polychaetous Annelids in the Museum of comparative Zoology. Bulletin of the Museum of Comparitive Zoology at Harvard College 85(1):3–31 + Plates 1–3.HartmanO1938Annotated list of the types of Polychaetous Annelids in the Museum of comparative ZoologyBulletin of the Museum of Comparitive Zoology at Harvard College851331+ Plates 1–3.Search in Google Scholar
Hartman, O. (1941). Polychaetous annelids. Part IV. Pectinariidae. With a review of all species from the Western Hemisphere. Allan Hancock Pacific Expeditions 7(5): 325–345.HartmanO.1941Polychaetous annelids. Part IV. Pectinariidae. With a review of all species from the Western HemisphereAllan Hancock Pacific Expeditions75325345Search in Google Scholar
Hartmann-Schröder, G. (1996). Annelida, Borstenwürmer, Polychaeta [Annelida, bristleworms, Polychaeta]. 2nd revised ed. The fauna of Germany and adjacent seas with their characteristics and ecology, 58. Gustav Fischer: Jena, Germany. ISBN 3-437-35038-2.Hartmann-SchröderG.1996Annelida, Borstenwürmer, Polychaeta [Annelida, bristleworms, Polychaeta]2nd revised ed.The fauna of Germany and adjacent seas with their characteristics and ecology, 58.Gustav FischerJena, GermanyISBN 3-437-35038-2.Search in Google Scholar
Hasselt, A.W.M. (1866). Kleine entomologische mededeelingen. No.6. Iets over de Phryganiden. Tijdschrift voor Entomologie 9: 211–215.HasseltA.W.M.1866Kleine entomologische mededeelingen. No.6. Iets over de PhryganidenTijdschrift voor Entomologie9211215Search in Google Scholar
Heerebout, G. (1968). C.S. Feuilleton. Het Zeepaard 28(3): 33–34.HeereboutG.1968C.S. FeuilletonHet Zeepaard2833334Search in Google Scholar
Heinsius, W.H. & Jaspers, J. (1913). Het strandboekje. 1e deel, 2e stuk. Bibliotheek van de Levende Natuur. AmsterdamHeinsiusW.H.JaspersJ.1913Het strandboekje. 1e deel, 2e stukBibliotheek van de Levende NatuurAmsterdamSearch in Google Scholar
Hessle, C. (1917). Zur Kenntnis der terebellomorphen Polychaeten. Zoologiska bidrag från Uppsala 5: 39–258, plates I–V.HessleC.1917Zur Kenntnis der terebellomorphen PolychaetenZoologiska bidrag från Uppsala539258plates I–V.Search in Google Scholar
Holthe, T. (1977). The zoogeography of the Terebellomorpha (Polychaeta) of the northern European waters, Sarsia 63(3): 191–198. DOI: 10.1080/00364827.1978.10411339.HoltheT.1977The zoogeography of the Terebellomorpha (Polychaeta) of the northern European watersSarsia63319119810.1080/00364827.1978.10411339Open DOISearch in Google Scholar
Holthe, T. (1986). Polychaeta Terebellomorpha. Marine Invertebrates of Scandinavia 7: 1–192. DOI: 10.1017/S0025315400026783.HoltheT.1986Polychaeta TerebellomorphaMarine Invertebrates of Scandinavia7119210.1017/S0025315400026783Open DOISearch in Google Scholar
Holthuis, L.B. (1978). Comments on the proposed validation of the generic name Pectinaria Lamarck, 1818, and the specific names P. belgica (Pallas, 1766) and P. koreni (Malmgren, 1866). Z.N.(S.) 2202. Bulletin of Zoological Nomenclature 35(1): 18–23.HolthuisL.B.1978Comments on the proposed validation of the generic name Pectinaria Lamarck, 1818, and the specific names P. belgica (Pallas, 1766) and P. koreni (Malmgren, 1866). Z.N.(S.) 2202Bulletin of Zoological Nomenclature3511823Search in Google Scholar
Hornell, J. (1892). Report on the polychaetous annelids of the L.M.B.C. district. Fauna of Liverpool Bay and the neigbouring seas 3: 126–171 + plates XII–XV.HornellJ.1892Report on the polychaetous annelids of the L.M.B.C. districtFauna of Liverpool Bay and the neigbouring seas3126171+ plates XII–XV.Search in Google Scholar
Horst, R. (1898). Naamlijst der tot de nederlandsche fauna behoorende Annelida Polychaeta. Tijdschrift der Nederlandsche Dierkundige Vereeniging 5(2): 15–28.HorstR.1898Naamlijst der tot de nederlandsche fauna behoorende Annelida PolychaetaTijdschrift der Nederlandsche Dierkundige Vereeniging521528Search in Google Scholar
Horst, R. (1922). Polychaete Anneliden. In H.C. Redeke (Ed.) Flora en Fauna der Zuiderzee. Monografie van een Brakwatergebied (pp. 262–275).HorstR.1922Polychaete AnnelidenInRedekeH.C.(Ed.)Flora en Fauna der Zuiderzee. Monografie van een Brakwatergebied262275Search in Google Scholar
Houttuyn, M. (1771). Natuurlyke historie of uitvoerige beschryving der dieren, planten en mineraalen, volgens het samenstel van den Heer Linnæus. Met naauwkeurige afbeeldingen. 1(16). De hoorens. Amsterdam (erven van F. Houttuyn). pp. [1–4], 1–629, Pl. CXX-CXXV [= 120–125]. DOI: 10.5962/bhl.title.66921.HouttuynM.1771Natuurlyke historie of uitvoerige beschryving der dieren, planten en mineraalen, volgens het samenstel van den Heer Linnæus. Met naauwkeurige afbeeldingen116De hoorens. Amsterdam (erven van F. Houttuyn). pp. [1–4], 1–629, Pl. CXX-CXXV [= 120–125].10.5962/bhl.title.66921Open DOISearch in Google Scholar
Hutchings, P.A. & Peart, R. (2002). A Review of the genera of Pectinariidae (Polychaeta) together with a description of the Australian Fauna. Records of the Australian Museum 54(1): 99–127. DOI: 10.3853/j.0067-1975.54.2002.1356.HutchingsP.A.PeartR.2002A Review of the genera of Pectinariidae (Polychaeta) together with a description of the Australian FaunaRecords of the Australian Museum5419912710.3853/j.0067-1975.54.2002.1356Open DOISearch in Google Scholar
Hutchings, P., Carrerette, O., Nogueira, J.M. de Matos (2020). Pectinariidae. In W. Westheide, G. Purschke & M. Böggemann (Eds.), Handbook of Zoology online. A Natural history of the phyla of the Animal kingdom. Annelida: Pleistoannelida, Sedentaria III and Errantia I. De Gruyter.HutchingsP.CarreretteO.NogueiraJ.M. de Matos2020PectinariidaeInWestheideW.PurschkeG.BöggemannM.(Eds.),Handbook of Zoology online. A Natural history of the phyla of the Animal kingdom. Annelida: Pleistoannelida, Sedentaria III and Errantia IDe GruyterSearch in Google Scholar
International Commission on Zoological Nomenclature (ICZN). (1982). OPINION 1225: Pectinaria Lamarck, 1818, Nereis cylindraria belgica Pallas, 1766 and Lagis koreni Malmgren, 1866 (Polychaeta): conserved. Bulletin of Zoological Nomenclature 39(3): 186–191.International Commission on Zoological Nomenclature (ICZN)1982OPINION 1225: Pectinaria Lamarck, 1818, Nereis cylindraria belgica Pallas, 1766 and Lagis koreni Malmgren, 1866 (Polychaeta): conservedBulletin of Zoological Nomenclature393186191Search in Google Scholar
Jirkov, I.A. (2001). Polychaeta of the Arctic Ocean. [Polikhety severnogo Ledovitogo Okeana]. Yanus-K Press, Moscow. (In Russian).JirkovI.A.2001Polychaeta of the Arctic Ocean[Polikhety severnogo Ledovitogo Okeana]Yanus-K PressMoscow(In Russian).Search in Google Scholar
Jirkov, I. A. & Leontovich, M. K. (2013). Identification keys for Terebellomorpha (Polychaeta) of the eastern Atlantic and the North Polar basin. Invertebrate Zoology [Zoologiya Bespozvonochnykh] 10(2): 217–243. DOI: 10.15298/invertzool.10.2.02.JirkovI. A.LeontovichM. K.2013Identification keys for Terebellomorpha (Polychaeta) of the eastern Atlantic and the North Polar basinInvertebrate Zoology [Zoologiya Bespozvonochnykh]10221724310.15298/invertzool.10.2.02Open DOISearch in Google Scholar
Johnston, G. (1865). A catalogue of the British non-parasitical worms in the collection of the British Museum. British Museum. London. DOI: 10.5962/bhl.title.20834.JohnstonG.1865A catalogue of the British non-parasitical worms in the collection of the British MuseumBritish MuseumLondon10.5962/bhl.title.20834Open DOISearch in Google Scholar
Jolly, M.T., Jollivet, D., Gentil, F., Thiébaut, E. & Viard, F. (2005). Sharp genetic break between Atlantic and English Channel populations of the polychaete Pectinaria koreni, along the North coast of France. Heredity 94: 23–32. DOI: 10.1038/sj.hdy.6800543.JollyM.T.JollivetD.GentilF.ThiébautE.ViardF.2005Sharp genetic break between Atlantic and English Channel populations of the polychaete Pectinaria koreni, along the North coast of FranceHeredity94233210.1038/sj.hdy.680054315305173Open DOISearch in Google Scholar
Kirkegaard, J.B. (1969). A quantitative investigation of the central North Sea Polychaeta. Spolia zool. Mus. haun 29.KirkegaardJ.B.1969A quantitative investigation of the central North Sea PolychaetaSpolia zool. Mus. haun29Search in Google Scholar
Kirkegaard, J.B. (1996). Havbørsteorme II. Sedentaria. Danmarks Fauna 86. Family 54 Pectinariidae (pp. 278–286). Kopenhagen.KirkegaardJ.B.1996Havbørsteorme II. SedentariaDanmarks Fauna 86. Family 54 Pectinariidae278286KopenhagenSearch in Google Scholar
Klein, J.T. (1731). Descriptiones tubulorum marinorum : in quorum censum relati lapides caudæ cancri Gesneri, & his similes : Belemnitæ, eorum que alueoli, secundum dispositionem musei Kleiniani : addita est Dissertatio epistolaris de pilis marinis : cum figuris. DOI: 10.5962/bhl.title.66797.KleinJ.T.1731Descriptiones tubulorum marinorum : in quorum censum relati lapides caudæ cancri Gesneri, & his similes : Belemnitæ, eorum que alueoli, secundum dispositionem musei Kleiniani : addita est Dissertatio epistolaris de pilis marinis : cum figuris10.5962/bhl.title.66797Open DOISearch in Google Scholar
Korringa, P. (1951). The shell of Ostrea edulis as a habitat. - Arch. néerl. Zool. 10(1): 32–152. DOI: 10.1163/036551654x00122.KorringaP.1951The shell of Ostrea edulis as a habitat.Arch. néerl. Zool.1013215210.1163/036551654x00122Open DOISearch in Google Scholar
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 desautres corps naturels, enfin, l'Exposition des Principes fondamentaux de la Zoologie. Paris, Deterville. vol 5: 612 pp. DOI: 10.5962/bhl.title.12712.LamarckJ.B.1818Histoire 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 desautres corps naturels, enfin, l'Exposition des Principes fondamentaux de la ZoologieParisDeterville5612 pp.10.5962/bhl.title.12712Open DOISearch in Google Scholar
Lameere, A. (1895). Manuel de la Faune de Belgique. Tome I, Animaux non insectes. H. Lamertin, Bruxelles. 639 pp. DOI: 10.5962/bhl.title.127632.LameereA.1895Manuel de la Faune de Belgique. Tome I, Animaux non insectesH. LamertinBruxelles639 pp.10.5962/bhl.title.127632Open DOISearch in Google Scholar
Leach, W.E. (1816). Annulosa. In Encyclopedia Britannica, supplement to the fourth, fifth and sixth editions 1: 401–453, pls. 20–26. (June 1816).LeachW.E.1816AnnulosaInEncyclopedia Britannica, supplement to the fourth, fifth and sixth editions 1401453pls. 20–26. (June 1816).Search in Google Scholar
Leenhouts, P. (1947a). Het C.S. Feuilleton (III). Het Zeepaard 7(4–5): 7–12LeenhoutsP.1947aHet C.S. Feuilleton (III)Het Zeepaard74–5712Search in Google Scholar
Leenhouts, P. (1947b). Het C.S. – feuilleton VI. Het Zeepaard 7(10–11): 6–9.LeenhoutsP.1947bHet C.S. – feuilleton VIHet Zeepaard710–1169Search in Google Scholar
Leewis, R. (2002). Veldgids Flora en Fauna van de Zee. KNNV Uitgeverij, Utrecht.LeewisR.2002Veldgids Flora en Fauna van de ZeeKNNV UitgeverijUtrechtSearch in Google Scholar
Le Mao, P., Godet, L., Fournier, J., Desroy, N., Gentil, F. et al. (2019). Atlas de la faune marine invertébrée du golfe Normano-Breton. Volume 2 – Annélides. Editions de la Station biologique de Roscoff.Le MaoP.GodetL.FournierJ.DesroyN.GentilF.2019Atlas de la faune marine invertébrée du golfe Normano-BretonVolume 2 – Annélides.Editions de la Station biologique de RoscoffSearch in Google Scholar
Leuckart, R. (1871). Bericht über die wissenschaftlichen Leistungen in der Naturgeschichte der niederen Thiere während der Jahre 1870 und 1871. Archiv für Naturgeschichte 37(2): 367–484.LeuckartR.1871Bericht über die wissenschaftlichen Leistungen in der Naturgeschichte der niederen Thiere während der Jahre 1870 und 1871Archiv für Naturgeschichte372367484Search in Google Scholar
Levinsen, G.M.R. (1884). Systematisk-geografisk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi. Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjöbenhavn 45(1883): 92–350, plates II–III. DOI: 10.5962/bhl.title.16117.LevinsenG.M.R.1884Systematisk-geografisk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og BalanoglossiVidenskabelige Meddelelser fra den naturhistoriske Forening i Kjöbenhavn45188392350plates II–III10.5962/bhl.title.16117Open DOISearch in Google Scholar
Linné, C. von (1767). Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Ed. 12. 1., Regnum Animale. 1 & 2. Holmiae, Laurentii Salvii. Holmiae [Stockholm], Laurentii Salvii (pp. 533–1327). DOI: 10.5962/bhl.title.37256.LinnéC. von1767Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locisEd. 12. 1., Regnum Animale. 1 & 2. Holmiae, Laurentii Salvii.Holmiae [Stockholm]Laurentii Salvii533132710.5962/bhl.title.37256Open DOISearch in Google Scholar
Lucas, J.A.W. & Holthuis, L.B. (1975). On the identity and nomenclature of ‘Pectinaria belgica Pallas, 1766)’ (Polychaeta, Amphictenidae). Zoologische Mededeelingen 49(9): 85–90.LucasJ.A.W.HolthuisL.B.1975On the identity and nomenclature of ‘Pectinaria belgica Pallas, 1766)’ (Polychaeta, Amphictenidae)Zoologische Mededeelingen4998590Search in Google Scholar
Maitland, R.S. (1851). Fauna Belgii septentrionalis: systematische beschrijving der dieren, welke in Noord Nederland of aan deszelfs kusten voorkomen met bijvoeging derzelver synonymen, benevens der plaatsen waar dezelve gevonden zijn, gerangschikt naar Prof. J. van der Hoeven's Handboek der Dierkunde (2de uitgave). C.G. van der Hoek: Leyden, Nederland. XII, XXXVIII, 234p. DOI: 10.5962/bhl.title.11256.MaitlandR.S.1851Fauna Belgii septentrionalis: systematische beschrijving der dieren, welke in Noord Nederland of aan deszelfs kusten voorkomen met bijvoeging derzelver synonymen, benevens der plaatsen waar dezelve gevonden zijn, gerangschikt naar Prof. J. van der Hoeven's Handboek der Dierkunde (2de uitgave)C.G. van der HoekLeyden, NederlandXII, XXXVIII, 234p.10.5962/bhl.title.11256Open DOISearch in Google Scholar
Malmgren, A.J. (1866). Nordiska Hafs-Annulater. [part three of three]. Öfversigt af Königlich Vetenskapsakademiens förhandlingar 22(5): 355–410, plates XVIII–XXIX.MalmgrenA.J.1866Nordiska Hafs-Annulater. [part three of three]Öfversigt af Königlich Vetenskapsakademiens förhandlingar225355410plates XVIII–XXIX.Search in Google Scholar
Malmgren, A.J. (1867). Annulater polychaeta Spetsbergiae, Grönlandiae, Islandiae et Scandinaviae hactenus cognita. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 24(4): 127–235, plates II–XV. DOI: 10.5962/bhl.title.13358.MalmgrenA.J.1867Annulater polychaeta Spetsbergiae, Grönlandiae, Islandiae et Scandinaviae hactenus cognitaÖfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar244127235plates II–XV.10.5962/bhl.title.13358Open DOISearch in Google Scholar
Martini, F.H.W. (1769). Neues systematisches Conchylien-Cabinet. Vol 1: 1–408 + Plate I–XXXI.MartiniF.H.W.1769Neues systematisches Conchylien-Cabinet11408+ Plate I–XXXI.Search in Google Scholar
McIntosh, W.C. (1914). Notes from the Gatty Marine On the ventral furrows of the lesser rorqual (Balaenoptera acutorostrata, Lacépède). 2. On some of the species of Prionospio, Malmgren. 3. On the british Amphictenidae. 4. On the british Ampharetidae. Annals and Magazine of Natural History 13: 77–110.McIntoshW.C.1914Notes from the Gatty Marine On the ventral furrows of the lesser rorqual (Balaenoptera acutorostrata, Lacépède). 2. On some of the species of Prionospio, Malmgren. 3. On the british Amphictenidae. 4. On the british AmpharetidaeAnnals and Magazine of Natural History1377110Search in Google Scholar
McIntosh, W.C. (1922). A monograph of the British marine annelids. Polychaeta: Hermellidae to Sabellidae. London, Ray Society Vol. 4(1) pp.1–250. DOI: 10.5962/bhl.title.54725.McIntoshW.C.1922A monograph of the British marine annelids. Polychaeta: Hermellidae to SabellidaeLondonRay Society41125010.5962/bhl.title.54725Open DOISearch in Google Scholar
Meyer, A.H. (1912). Die Amphicteniden, Ampharetiden und Terebelliden der Nord- und Ostsee. Inaugural-Dissertation zur Erlangung der Doktorwürde der hohen philosophischen Fakultät der Königlichen Christian-Albrechts-Universität zu Kiel. Heider Anzeiger, Heide. 1–68 pp.MeyerA.H.1912Die Amphicteniden, Ampharetiden und Terebelliden der Nord- und Ostsee. Inaugural-Dissertation zur Erlangung der Doktorwürde der hohen philosophischen Fakultät der Königlichen Christian-Albrechts-Universität zu KielHeider AnzeigerHeide168Search in Google Scholar
Michaelsen, W. (1897). Die Polychaetenfauna der deutschen Meere einschliesslich der benachbarten und verbindenden Gebiete. Wissenschaftliche Meeresuntersuchungen herausgegeben von der Kommission zur wissenschaftlichen Untersuchung der deutschen Meere in Kiel und der Biologischen Anstalt auf Helgoland, Neue Folge 2(1–1): 1–216, plate I.MichaelsenW.1897Die Polychaetenfauna der deutschen Meere einschliesslich der benachbarten und verbindenden GebieteWissenschaftliche Meeresuntersuchungen herausgegeben von der Kommission zur wissenschaftlichen Untersuchung der deutschen Meere in Kiel und der Biologischen Anstalt auf Helgoland, Neue Folge21–11216plate ISearch in Google Scholar
Möbius, K. (1873). Die faunistischen Untersuchungen. A. Die wirbellosen Thiere der Ostsee. In: Die Exp. zur physikalisch-chem. und biol. Untersuchung der Ostsee im Sommer 1870 S.M. Avisodampfer Pommerania nebst physikalischen Beob. an den Stat. der preus. Ostseeküste. Erster Jahresber. Comm. Wiss. Unters. Dtsch. Meere Kiel 1: 97–144.MöbiusK.1873Die faunistischen Untersuchungen. A. Die wirbellosen Thiere der Ostsee. In: Die Exp. zur physikalisch-chem. und biol. Untersuchung der Ostsee im Sommer 1870 S.M. Avisodampfer Pommerania nebst physikalischen Beob. an den Stat. der preus. Ostseeküste. Erster JahresberComm. Wiss. Unters. Dtsch. Meere Kiel197144Search in Google Scholar
Montagu, G. (1803). Testacea Britannica or natural history of British shells, marine, land, and fresh-water, including the most minute: Systematically arranged and embellished with figures. J. White, London, Vol. 1, xxxvii + 291 pp;; Vol. 2, pp. 293–606, pl. 1–16. DOI: 10.5962/bhl.title.33927.MontaguG.1803Testacea Britannica or natural history of British shells, marine, land, and fresh-water, including the most minute: Systematically arranged and embellished with figuresJ. WhiteLondon1xxxvii + 291 pp;; Vol. 2, pp. 293–606, pl. 1–16.10.5962/bhl.title.33927Open DOISearch in Google Scholar
Müller, O.F. (1776). Zoologiæ Danicæ Prodromus, seu Animalium Daniæ et Norvegiæ indigenarum characteres, nomina, et synonyma imprimis popularium. Havniæ [Copenhagen]: Hallageri. xxxii + 274 pp. DOI: 10.5962/bhl.title.13268.MüllerO.F.1776Zoologiæ Danicæ Prodromus, seu Animalium Daniæ et Norvegiæ indigenarum characteres, nomina, et synonyma imprimis populariumHavniæ [Copenhagen]Hallagerixxxii + 274 pp.10.5962/bhl.title.13268Open DOISearch in Google Scholar
Müller, O.F. (1779). Zoologica danica sev Animalum daniae et norvegiae. Rariorum ac minus notorum descriptiones et historia. Volumen Primum. DOI: 10.5962/bhl.title.12923.MüllerO.F.1779Zoologica danica sev Animalum daniae et norvegiae. Rariorum ac minus notorum descriptiones et historiaVolumen Primum10.5962/bhl.title.12923Open DOISearch in Google Scholar
Müller O.F. (1788). Zoologia Danica seu animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historia. Volumen primum: pp. 1–5, 1–46, pls. 121–160. Havniae [Copenhague], N. Möller. DOI: 10.5962/bhl.title.49477.MüllerO.F.1788Zoologia Danica seu animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historiaVolumen primum: pp. 1–5, 1–46, pls. 121–160.Havniae [Copenhague]N. Möller10.5962/bhl.title.49477Open DOISearch in Google Scholar
Nielsen, C., Kirkegaard, J.B. & Lemche, H. (1977). Pectinaria Lamarck, 1818 (Polychaeta), and the specific names P. belgica (Pallas, 1766) and P. koreni (Malmgren, 1866) to be validated under the plenary powers. Z.N. (S.) 2202. Bulletin of Zoological Nomenclature 34(2): 112–122.NielsenC.KirkegaardJ.B.LemcheH.1977Pectinaria Lamarck, 1818 (Polychaeta), and the specific names P. belgica (Pallas, 1766) and P. koreni (Malmgren, 1866) to be validated under the plenary powers. Z.N. (S.) 2202Bulletin of Zoological Nomenclature342112122Search in Google Scholar
Nielsen, C. & Kirkegaard, J.B. (1978). Amended proposal for validating Pectinaria Lamarck, 1818 (Polychaeta), Pectinaria belgica (Pallas, 1766) and P. koreni (Malmgren, 1866) under the plenary powers Z.N.(S.) 2202. Bulletin of Zoological Nomenclature 35(1): 25–29. DOI: 10.5962/bhl.part.14601.NielsenC.KirkegaardJ.B.1978Amended proposal for validating Pectinaria Lamarck, 1818 (Polychaeta), Pectinaria belgica (Pallas, 1766) and P. koreni (Malmgren, 1866) under the plenary powers Z.N.(S.) 2202Bulletin of Zoological Nomenclature351252910.5962/bhl.part.14601Open DOISearch in Google Scholar
Nilsson, D. (1928). Neue und alte Amphicteniden. Göteborgs Königlich vetenskaps- och vitterhetssamhälles handlingar 4(33): 1–96.NilssonD.1928Neue und alte AmphictenidenGöteborgs Königlich vetenskaps- och vitterhetssamhälles handlingar433196Search in Google Scholar
Pallas, P.S. (1766). Miscellanea zoologica. Quibus novae imprimis atque obscurae animalium species describuntur et observationibus iconibusque illustrantur. Petrum van Cleef. Hagí Comitum., xii + 224 pp., 14 pls. DOI: 10.5962/bhl.title.69851.PallasP.S.1766Miscellanea zoologica. Quibus novae imprimis atque obscurae animalium species describuntur et observationibus iconibusque illustranturPetrum van Cleef. Hagí Comitum.xii + 224 pp., 14 pls.10.5962/bhl.title.69851Open DOISearch in Google Scholar
Parapar, J., Palomanes, V., Helgason, G.V. & Moreira, J. (2020). Taxonomy and distribution of Pectinariidae (Annelida) from Iceland with a comparative analysis of uncinal morphology. European Journal of Taxonomy 666: 1–32. DOI: 10.5852/ejt.2020.666.ParaparJ.PalomanesV.HelgasonG.V.MoreiraJ.2020Taxonomy and distribution of Pectinariidae (Annelida) from Iceland with a comparative analysis of uncinal morphologyEuropean Journal of Taxonomy66613210.5852/ejt.2020.666Open DOISearch in Google Scholar
Pennant, T. (1777). British Zoology, vol. IV. Crustacea. Mollusca. Testacea. London. i–viii, 1–154, Plates 1–93. DOI: 10.5962/bhl.title.62481.PennantT.1777British Zoology, vol. IV. Crustacea. Mollusca. TestaceaLondoniviii1–154, Plates 1–93.10.5962/bhl.title.62481Open DOISearch in Google Scholar
Pennant, T. (1812). British Zoology. A new edition in four volumes. vol IV. Class V. Crustacea. Class VI. Vermes. Wilkie & Robinson. London. xvi + 379 pp. 95 pls. DOI: 10.5962/bhl.title.51161.PennantT.1812British Zoology. A new edition in four volumes. vol IV. Class V. Crustacea. Class VI. VermesWilkie & RobinsonLondonxvi + 379 pp. 95 pls.10.5962/bhl.title.51161Open DOISearch in Google Scholar
Perrier, E. (1897). Traité de Zoologie IV: Vers (suite) – Mollusques – Tuniciers. Paris. Chapter on Anneles, Chetopodes, Polychetes (pp. 1345–1643). DOI: 10.5962/bhl.title.11522.PerrierE.1897Traité de Zoologie IV: Vers (suite) – Mollusques – TuniciersParisChapter on Anneles, Chetopodes, Polychetes1345164310.5962/bhl.title.11522Open DOISearch in Google Scholar
Pieters, F.F.J.M. (1980). Notes on the menagerie and zoological cabinet of Stadholder William V of Holland, directed by Aernout Vomaer. J. Soc. Biblphy nat. Hist 9(4): 539–563. DOI: 10.3366/jsbnh.1980.9.4.539.PietersF.F.J.M.1980Notes on the menagerie and zoological cabinet of Stadholder William V of Holland, directed by Aernout VomaerJ. Soc. Biblphy nat. Hist9453956310.3366/jsbnh.1980.9.4.539Open DOISearch in Google Scholar
Prins, B. (1999). 17. Oorlog langs de Nederlandse kust. In Koninklijke Nederlandse Natuurhistorische Vereniging, Kwartaalbericht Zoetermeer nr 23 (jan. 1999): 32–39PrinsB.199917. Oorlog langs de Nederlandse kustInKoninklijke Nederlandse Natuurhistorische Vereniging, Kwartaalbericht Zoetermeer23jan.19993239Search in Google Scholar
Prud’Homme van Reine, W.J. (1957). Wat vindt ik aan het strand? Excursieboekje voor de Nederlandse Noordzeekust, met 450 afbeeldingen van planten en dieren. Vijfde Druk. Thieme & Cie, Zutphen.Prud’Homme van ReineW.J.1957Wat vindt ik aan het strand? Excursieboekje voor de Nederlandse Noordzeekust, met 450 afbeeldingen van planten en dierenVijfde DrukThieme & Cie, ZutphenSearch in Google Scholar
Quatrefages, A. (1865). Histoire naturelle des Annelés marins et d'eau douce. Annélides et Géphyriens. Volume 2. Première partie. 1–336. Explication des planches pp. 1–24. planches 1–20. Librarie Encyclopédique de Roret. Paris. DOI: 10.5962/bhl.title.122818.QuatrefagesA.1865Histoire naturelle des Annelés marins et d'eau douce. Annélides et GéphyriensVolume 2. Première partie. 1–336. Explication des planches pp. 1–24. planches 1–20.Librarie Encyclopédique de RoretParis10.5962/bhl.title.122818Open DOISearch in Google Scholar
Rathke, H. (1843). Beiträge zur Fauna Norwegens. Nova Acta Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, Breslau & Bonn 20: 1–264c. DOI: 10.5962/bhl.title.11613.RathkeH.1843Beiträge zur Fauna NorwegensNova Acta Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, Breslau & Bonn201264c10.5962/bhl.title.11613Open DOISearch in Google Scholar
Read, G. & Fauchald, K. (Ed.) (2020, October 16). World Polychaeta database. Pectinariidae Quatrefages, 1866. From: http://www.marinespecies.org/aphia.php?p=taxdetails&id=980ReadG.FauchaldK.(Ed.)2020October16World Polychaeta database. Pectinariidae Quatrefages, 1866From: http://www.marinespecies.org/aphia.php?p=taxdetails&id=980Search in Google Scholar
Renier, S.A. (1804). Prospetto della Classe dei Vermi, nominati el ordinati secondo il Sistemo de Bosc. [Page proofs of introductory book text in folio sheets, never published]. xv–xxvi.RenierS.A.1804Prospetto della Classe dei Vermi, nominati el ordinati secondo il Sistemo de Bosc. [Page proofs of introductory book text in folio sheets, never published]xvxxviSearch in Google Scholar
Rouse, G.W. & Pleijel, F. (2002). Polychaetes. Oxford University Press.RouseG.W.PleijelF.2002PolychaetesOxford University PressSearch in Google Scholar
Saint-Joseph, A.A. de. (1898). Annélides polychètes des côtes de France (Manche et Océan). Annales des Sciences Naturelles, Zoologie et Paléontologie, Paris Série 8(5): 209–464, plates XIII–XXIII.Saint-JosephA.A. de.1898Annélides polychètes des côtes de France (Manche et Océan)Annales des Sciences Naturelles, Zoologie et Paléontologie, Paris Série85209464plates XIII–XXIIISearch in Google Scholar
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 ou Recueil des Observations et des Recherches qui ont été faites en Égypte pendant l'Expédition de l'Armée Française, publié par les Ordres de sa Majesté l'Empereur Napoléon le Grand, Histoire Naturelle, Paris 1(3):1–128.SavignyJ.-C.1822Système 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 EspècesDescription de l'Égypte ou Recueil des Observations et des Recherches qui ont été faites en Égypte pendant l'Expédition de l'Armée Française, publié par les Ordres de sa Majesté l'Empereur Napoléon le Grand, Histoire Naturelle, Paris131128Search in Google Scholar
Southern, R. (1910). The marine worms (Annelida) of Dublin Bay and the adjoining district. Proceedings of the Royal Irish Academy 28B(6): 215–246.SouthernR.1910The marine worms (Annelida) of Dublin Bay and the adjoining districtProceedings of the Royal Irish Academy28B6215246Search in Google Scholar
Southern, R. (1914). Clare Island Survey. Archiannelida and Polychaeta. Proceedings of the Royal Irish Academy 31(47): 1–160.SouthernR.1914Clare Island Survey. Archiannelida and PolychaetaProceedings of the Royal Irish Academy31471160Search in Google Scholar
Sowerby, J. (1806). The British miscellany; or, Coloured figures of new, rare, or little known animal subjects : many not before ascertained to be inhabitants of the British Isles: and chiefly in the possession of the author, James Sowerby. R. Taylor & Co., London, Vol. 1–2 136 pp., 76 plates. DOI: 10.5962/bhl.title.120056.SowerbyJ.1806The British miscellany; or, Coloured figures of new, rare, or little known animal subjects : many not before ascertained to be inhabitants of the British Isles: and chiefly in the possession of the author, James SowerbyR. Taylor & Co.London1–2136 pp., 76 plates.10.5962/bhl.title.120056Open DOISearch in Google Scholar
Spaink, G. (1960) C.S. Feuilleton. Het Zeepaard 20(3): 38–42.SpainkG.1960C.S. FeuilletonHet Zeepaard2033842Search in Google Scholar
Swammerdam, J. (1737–1738). Bijbel der natuure. Posthumously published by H. Boerhaave and translated from Latin by H.D.Gaubius. Volume 1 (1737): 1–362 + Plate I–XVI, Vol. 2 (1738): pp. 364–910 + Plate LIII, Legenda (1738): p1–124 + Plates XVII-LII. DOI: 10.5962/bhl.title.119987.SwammerdamJ.1737–1738Bijbel der natuurePosthumously published by H. Boerhaave and translated from Latin by H.D.Gaubius.117371362+ Plate I–XVI, Vol. 2 (1738): pp. 364–910 + Plate LIII, Legenda (1738): p1–124 + Plates XVII-LII.10.5962/bhl.title.119987Open DOISearch in Google Scholar
Swennen, C. (1955). Van de C.S. Man. Het Zeepaard 15(6): 86–89.SwennenC.1955Van de C.S. ManHet Zeepaard1568689Search in Google Scholar
Turton, W. (1807). British Fauna, containing a compendium of the zoology of the British Islands; arranged according to the Linnean system. Vol. 1, Mammalia, birds, Amphibia, fishes and worms (pp. 1–230). DOI: 10.5962/bhl.title.27635.TurtonW.1807British Fauna, containing a compendium of the zoology of the British Islands; arranged according to the Linnean system1Mammalia, birds, Amphibia, fishes and worms123010.5962/bhl.title.27635Open DOISearch in Google Scholar
Ushakov, P.V. (1955). Polychaeta of the far eastern seas of the U.S.S.R. Fauna SSSR, Akademiya Nauk SSSR, Zoologicheskii Institut 56: 1–443. (In Russian).UshakovP.V.1955Polychaeta of the far eastern seas of the U.S.S.R. Fauna SSSRAkademiya Nauk SSSR, Zoologicheskii Institut561443(In Russian).Search in Google Scholar
Van Hasselt, A.W.M. (1866). Kleine Entomologische mededeelingen No 6: Iets over de Phryganiden. Tijdschrift voor Entomologie 9(2): 211–215.Van HasseltA.W.M.1866Kleine Entomologische mededeelingen No 6: Iets over de PhryganidenTijdschrift voor Entomologie92211215Search in Google Scholar
Verhas, G. (1925). Le long de no plages. Flore et Faune du littoral Belge. Maurice Lamertin, Bruxelles. pp. 139 + plates 1–20.VerhasG.1925Le long de no plages. Flore et Faune du littoral BelgeMaurice LamertinBruxelles139+ plates 1–20.Search in Google Scholar
Von Marenzeller, E. (1874). Ueber Lagis (Pectinaria) koreni Mgr., aus den Mittelmeere und die Hakenborsten der Amphicteneen. Verhandlungen der Zoologisch-Botanische Gesellschaft, Wien 24: 217–224.Von MarenzellerE.1874Ueber Lagis (Pectinaria) koreni Mgr., aus den Mittelmeere und die Hakenborsten der AmphicteneenVerhandlungen der Zoologisch-Botanische Gesellschaft, Wien24217224Search in Google Scholar
Wolff, W.J. (1966). Het voorkomen van de fluwelen zwemkrab, Macropipus puber in Nederland. Het Zeepaard 26(3): 42–49.WolffW.J.1966Het voorkomen van de fluwelen zwemkrab, Macropipus puber in NederlandHet Zeepaard2634249Search in Google Scholar
Wolff, W.J. (1970). De goudkammetjes van het Nederlandse strand. Het zeepaard 30(6): 100–101WolffW.J.1970De goudkammetjes van het Nederlandse strandHet zeepaard306100101Search in Google Scholar
Wollebaek, A. (1912). Nordeuropaeiske annulata Polychaeta I. Ammocharidae, Amphictenidae, Ampharetidae, Terebellidae og Serpulidae. Skrifter utgit av Videnskapsselskapet i Kristiania. I, Matematisknaturvidenskabelig klasse. year 1911 part 2 no. 18: 1–144. DOI: 10.5962/bhl.title.11634.WollebaekA.1912Nordeuropaeiske annulata Polychaeta I. Ammocharidae, Amphictenidae, Ampharetidae, Terebellidae og SerpulidaeSkrifter utgit av Videnskapsselskapet i Kristiania. I, Matematisknaturvidenskabelig klasseyear 1911 part 2 no. 18: 1–144.10.5962/bhl.title.11634Open DOISearch in Google Scholar
Zhang, J. & Hutchings, P. (2019). A revision of Australian Pectinariidae (Polychaeta), with new species and new records. Zootaxa 4611(1): 1–70. DOI: 10.11646/zootaxa.4611.1.1.ZhangJ.HutchingsP.2019A revision of Australian Pectinariidae (Polychaeta), with new species and new recordsZootaxa4611117010.11646/zootaxa.4611.1.131717089Open DOISearch in Google Scholar
Zhang, J., Hutchings, P. & Kupriyanova, E. (2019). A revision of the genus Petta Malmgren, 1866 (Annelida: Pectinariidae), with two new species from deep waters of southeastern Australia, and comments on phylogeny of the family. Zootaxa 4614 (2): 303–330. DOI: 10.11646/zootaxa.4614.2.3.ZhangJ.HutchingsP.KupriyanovaE.2019A revision of the genus Petta Malmgren, 1866 (Annelida: Pectinariidae), with two new species from deep waters of southeastern Australia, and comments on phylogeny of the familyZootaxa4614230333010.11646/zootaxa.4614.2.331716379Open DOISearch in Google Scholar
