ZooKeys 241: 21–31, doi: 10.3897/zookeys.241.3820
Oxydromus Grube, 1855 reinstated over Ophiodromus Sars, 1862 (Polychaeta, Hesionidae)
Tulio F. Villalobos-Guerrero 1, Leslie H. Harris 2
1 Geomare A. C., Mazatlán, Sinaloa, México
2 Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA, 90007

Corresponding author: Tulio F. Villalobos-Guerrero (tulio1786@msn.com)

Academic editor: Chris Glasby

received 12 August 2012 | accepted 24 October 2012 | Published 12 November 2012

(C) 2012 Tulio F. Villalobos-Guerrero. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.


The hesionid polychaete genera Oxydromus Grube, 1855 and Ophiodromus Sars, 1862 have been regarded as synonyms with the former considered as invalid since it was thought to be a junior homonym of Oxydromus Schlegel, 1854. However, Schlegel’s name is an incorrect subsequent spelling for Ocydromus Wagler, 1830 (Aves, Gruiformes, Rallidae) and is not an available name. Consequently, Oxydromus Grube, 1855 must be reinstated for this hesionid polychaete genus. A check-list of valid species of Oxydromus including 30 new combinations is provided.


Nomenclature, taxonomy, hesionid, Phyllodocida, Annelida


Grube (1855) proposed Oxydromus within the polychaete family Phyllodocidae for Oxydromus fasciatus Grube, 1855, a new species capable of rapid movement from two Mediterranean Sea localities: Trieste (Italy) and Villa Franca (probably Villefranche-sur-Mer, France). Later, Sars (1862) established the genus Ophiodromus for a Norwegian species, Ophiodromus vittatus Sars, 1862. He also transferred Oxydromus to the family Hesionidae and distinguished it from Ophiodromus by the presence of articulated palps and biramous parapodia. Both features were present in Ophiodromus fasciatus but misinterpreted by Grube when he defined them as simple palps and uniramous parapodia (von Marenzeller 1874, Pleijel 2011 pers. comm.).

Pleijel (1998) examined syntypes of Oxydromus fasciatus (ZMB 3825), the type species of the genus, and specimens of Nereis flexuosa delle Chiaje, 1825 (currently Ophiodromus flexuosus fide Pleijel 1998) from near the type locality (type material of this species does not exist). He agreed with McIntosh (1908:116) that the former is a junior synonym of the latter although he didn’t go into detail. Nevertheless, regarding the defining generic characters, Ophiodromus and Oxydromus are synonymous. Ophiodromus flexuosus is possibly a senior synonym of the type species Ophiodromus vittatus (fide von Marenzeller 1874, McIntosh 1908, Pleijel 1998, Fauchald 2011). An examination of specimens from the type localities, Gulf of Naples and Norway respectively, is required to resolve their status.

Pleijel (1998) pointed out that Oxydromus has seniority over Ophiodromus but, as first stated by Hartman (1965), concluded that the former genus name was preoccupied in the class Aves, Oxydromus Schlegel, 1854, and for which reason Ophiodromus must be used. However, Viéitez et al. (2004) argued that Oxydromus is an available genus name and must be considered as valid. Then, following their suggestion, we proposed to reinstate Oxydromus over Ophiodromus to standardize the worldwide use of both generic names.


Viéitez et al. (2004:521) realized that Oxydromus Schlegel is an incorrect subsequent spelling of Ocydromus Wagler, 1830 (Aves, Gruiformes, Rallidae) (itself a junior homonym of the beetle genus Ocydromus Clairville, 1806 [Insecta, Carabidae], and replaced by Gallirallus Lafresnaye, 1841). Although Schlegel’s name was used in later publications (e. g. Reischek 1886, Röse 1890), as an incorrect subsequent spelling it remains unavailable according to Article 33.3 of the International Code of Zoological Nomenclature. This states “Any subsequent spelling of a name different from the correct original spelling, other than a mandatory change or an emendation, is an ‘incorrect subsequent spelling’; it is not an available name and, like an incorrect original spelling [Article 32.4], it does not enter in homonymy and cannot be used as a substitute name…”. Viéitez et al. erroneously stated that Sars proposed Ophiodromus to replace Oxydromus Grube due to his mistaken belief that Grube’s genus name is a junior homonym of Oxydromus Schlegel. Sars actually didn’t mention the homonymy; instead he discussed the morphological discrepancy between both genera.

Preservation of the genus-group name Ophiodromus as the senior synonym would require a reversal of precedence according to Article 23.9 (ICZN 1999). This states that in order to maintain the prevailing usage the following conditions must be met: 1) the senior synonym (in this case Oxydromus Grube, 1855) has not been used as a valid name after 1899 (Article, and 2) the junior synonym (namely Ophiodromus Sars, 1862) has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years (Article The first condition is not applicable which negates the need for a reversal of precedence. Oxydromus has also been listed as a valid name after 1900 in many publications (e.g. Gravier 1900, McIntosh 1908, Chamberlin 1919, Fauvel 1923, Augener 1927, Uschakov 1955, Hartmann-Schröder 1959, 1965, Hartman 1961, Day 1963, Averincev 1972).

In accordance with Article 23.9.3 (ICZN 1999), we consider that the Oxydromus/Ophiodromus situation does not threaten the stability of nomenclature or the universality of a widespread use, thus no referral to the Commission for a ruling should be required. We propose to reinstate Oxydromus over Ophiodromus based on the Principle of Priority (Article 23, ICZN 1999).

Systematics Family HESIONIDAE Grube, 1850 Subfamily OPHIODROMINAE Pleijel, 1998 Tribe OPHIODROMINI Pleijel, 1998
Oxydromus Grube, 1855, reinstated
Oxydromus Grube, 1855: 98.
Ophiodromus Sars, 1862: 87; Pleijel, 1998: 137–143, figs. 31–33 (synonymy).
Type species.

Oxydromus fasciatus Grube, 1855, by monotypy.

Oxydromus adorsosetosus (Hartmann-Schröder, 1985), comb. n. (basionym of Ophiodromus adorsosetosus Hartmann-Schröder, 1985)

Type locality: Port Lincoln, South Australia.

Oxydromus adspersus (Grube, 1874), comb. n. (basionym of Ophiodromus adspersus Grube, 1874)

Type locality: Dalmatia, Croatia.

Oxydromus agilis (Ehlers, 1864), comb. n. (basionym of Ophiodromus agilis Ehlers, 1864)

Type locality: Adriatic Sea.

Oxydromus angolaensis (Hartmann-Schröder, 1974), comb. n. (basionym of Podarke angolaensis Hartmann-Schröder, 1974)

Type locality: Lobito, Angola.

Oxydromus angustifrons (Grube, 1878), comb. n. (basionym of Irma angustifrons Grube, 1878)

Type locality: Philippines.

Oxydromus berrisfordi (Day, 1967), comb. n. (basionym of Ophiodromus berrisfordi Day, 1967)

Type locality: Walvis Bay, Namibia.

Oxydromus brevipodius (Uchida, 2004), comb. n. (basionym of Ophiodromus brevipodius Uchida, 2004)

Type locality: Wakayama, Japan.

Oxydromus bunbuku (Uchida, 2004), comb. n. (basionym of Ophiodromus bunbuku Uchida, 2004)

Type locality: Shikoku, Japan.

Oxydromus constrictus (Uchida, 2004), comb. n. (basionym of Ophiodromus constrictus Uchida, 2004)

Type locality: Wakayama, Japan.

Oxydromus didymocerus (Schmarda, 1861), comb. n. (basionym of Cirrosyllis didymocerus Schmarda, 1861)

Type locality: New South Wales, Australia.

Oxydromus fasciatus Grube, 1855 (possible junior synonym of Oxydromus flexuosus (delle Chiaje, 1825))

Type locality: Adriatic Sea (Trieste) and Western Mediterranean Sea (Villefranche-sur-Mer, France).

Oxydromus fauveli (Uchida, 2004), comb. n. (basionym of Ophiodromus fauveli Uchida, 2004)

Type locality: Wakayama, Japan.

Oxydromus flexuosus (delle Chiaje, 1825) (basionym of Nereis flexuosa delle Chiaje, 1825)

Type locality: Gulf of Naples.

Oxydromus furcatus (Hartmann-Schroder, 1962), comb. n. (basionym of Podarke furcatus Hartmann-Schroder, 1962)

Type locality: Peru.

Oxydromus guanicus (Hoagland, 1919), comb. n. (basionym of Podarke guanica Hoagland, 1919)

Type locality: Guanica, Puerto Rico.

Oxydromus latifrons (Grube, 1878), comb. n. (basionym of Irma latifrons Grube, 1878)

Type locality: Philippines.

Oxydromus limicolus (Willey, 1905), comb. n. (basionym of Irma limicola Willey, 1905)

Type locality: Sri Lanka.

Oxydromus longifundus (Uchida, 2004), comb. n. (basionym of Ophiodromus longifundus Uchida, 2004)

Type locality: Okinawa, Japan.

Oxydromus longicirratus (Knox and Cameron, 1971), comb. n. (basionym of Nereimyra longicirratus Knox and Cameron, 1971; not a senior homonym to Oxydromus longocirratus (Tenerelli, 1973) as Pleijel (1998) suggested, but this name is considered as nomen dubium)

Type locality: Melbourne, Australia.

Oxydromus microantennatus (Hutchings and Murray, 1984), comb. n. (basyonymy of Podarke microantennata Hutchings and Murray, 1984)

Type locality: New South Wales, Australia.

Oxydromus minutus (Hartmann-Schröder, 1959), comb. n. (basyonymy of Podarke minuta Hartmann-Schröder, 1959)

Type locality: San Juan, El Salvador.

Oxydromus mutilatus (Treadwell, 1901), comb. n. (basyonymy of Castalia mutilata Treadwell, 1901)

Type locality: Puerto Rico.

Oxydromus notospinosus (Rosito, 1983), comb. n. (basionym of Ophiodromus notospinosus Rosito, 1983)

Type locality: Philippines.

Oxydromus obscurus (Verrill, 1873), comb. n. (basyonymy of Podarke obscura Verrill, 1873)

Type locality: Massachusetts, United States.

Oxydromus okudai (Uchida, 2004), comb. n. (basionym of Ophiodromus okudai Uchida, 2004)

Type locality: Nagasaki, Japan.

Oxydromus pallidus Claparède, 1864

Type locality: Golfe du Lion, France.

Oxydromus parapallidus (Uchida, 2004), comb. n. (basionym of Ophiodromus parapallidus Uchida, 2004)

Type locality: Wakayama, Japan.

Oxydromus pelagicus (Rioja, 1923), comb. n. (basionym of Ophiodromus pelagicus Rioja, 1923)

Type locality: Pontavedra, Spain.

Oxydromus pugettensis (Johnson, 1901), comb. n. (basyonymy of Podarke pugettensis Johnson, 1901) (Figure 1)

Type locality: Washington, United States.

Oxydromus spinapandens (Storch and Niggemann, 1967), comb. n. (basyonymy of Podarke pugettensis spinapandens Storch and Niggemann, 1967)

Type locality: Red Sea.

Oxydromus spinosus (Ehlers, 1908), comb. n. (basyonymy of Orthodromus spinosus Ehlers, 1908)

Type locality: Angola.

Oxydromus viridescens (Ehlers, 1864), comb. n. (basyonymy of Podarke viridescens Ehlers, 1864)

Type locality: Adriatic Sea.

Oxydromus vittatus (Sars, 1862), comb. n. (basionym of Ophiodromus vittatus Sars, 1862)

Type locality: Norway.

Figure 1.

A representative living specimen of Oxydromus collected in California. Species: Oxydromus pugettensis (Johnson, 1901) (photo: Leslie Harris).

Remarks and discussion

Pleijel (1998) revised the phylogeny and classification of the family Hesionidae based on available type and non-type material. He provided descriptions and diagnoses for all supraspecific taxa and world-wide species lists, including 24 nominal taxa in Ophiodromus. He newly synonymized Orseis Ehlers, 1864 and Schmardiella Czerniavsky, 1882, and continued the prior synonymy of Oxydromus Grube, 1855, Podarke Ehlers, 1864, Mania de Quatrefages, 1866 and Irma Grube, 1878 with Ophiodromus Sars, 1862, whose type species is Ophiodromus vittatus Sars, 1862 (possibly Ophiodromus flexuosus fide Pleijel 1998). However, Oxydromus is herein reinstated due to the priority of Oxydromus over Ophiodromus Sars, with Ophiodromus fasciatus as the type species.

Oxydromus (from the Greek oxys= fast, quick; dromus= runner) (Figure 1) is currently represented by 32 species and is distinguished from other genera by the presence of six pairs of enlarged anterior cirri, prostomium bearing three antennae with the median one sited anteriorly, and biarticulated palps (see Salazar-Vallejo and Orensanz 2006). Uchida (2004) suggested that the species of this genus are among the most difficult groups to identify in the family and considered that the form of parapodia is a more useful character for identification of the species than prostomium and anterior part structures. A detailed revision of the genus Oxydromus is required to redefine species, especially those considered to be widely distributed.


Special thanks are given to Sergio I. Salazar-Vallejo, María Ana Tovar-Hernández and an anonymous reviewer whose important suggestions on nomenclatural issues improved this contribution. We wish to also thank Fredrik Pleijel and José Viéitez and colleagues whose papers contributed so much to this study. In particular, Viéitez et al’s (2004) comment on the availability of Oxydromus inspired TFVG to carry out their suggestion that the conflict between the two names be formally resolved. Workspace for TFVG was kindly facilitated by Nuria Méndez in ICMyL, UNAM, and was greatly appreciated.

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