Hydroides Gunnerus, 1768 (Annelida, Serpulidae) is feminine: a nomenclatural checklist of updated names.

As a service to taxonomists and ecologists using names in the well-known and species-rich ship-fouling serpulid genus Hydroides we present an update of all 107 non-synonymised scientific names, with additional information on Hydroides nomenclature, original names, etymologies, and type localities derived from original literature, and in accord with the World Register of Marine Species (WoRMS) database. An update is needed because the gender of genus Hydroides has from 1 January 2000 reverted to the original feminine, due to a change in the wording of International Code of Zoological Nomenclature which was overlooked at that time, and is contrary to the usage in practice of Hydroides as masculine which had started about 1992, although Code-required from the 1960s. We match 31 further original names of current WoRMS subjective junior synonyms to each non-synonymised name, and also report on the world distribution of the genus as illustrated by type localities of the valid names. We include notes on seven species inquirenda. The correct rendering is given of six names that have been altered for gender agreement for the first time herein. Hydroides gottfriedinom. n. replaces junior homonym Hydroides rostrata Pillai, 1971. Currently there are 41 non-synonymised species-group names in Hydroides which should be gender invariant, and 23 names which would only change if moved to a neuter genus; the remaining 43 names are fully gender variable. Place-names (23), and personal names (16) make up more than a third (36%) of the species names, with most of the remainder (68) being descriptive of species character states, usually of operculum morphology (54). All species, except Hydroides norvegica (63°N), have type localities in shallow-water coastal locations in temperate to tropical waters below latitude 44°, with the highest number of new species (54) from the adjoining Western Pacific and Indian Ocean areas. The other concentration of new species (31) are those first found on the Pacific and Atlantic coasts of North America and in the Caribbean.

Introduction

An unusual situation has arisen concerning the correct formulation and spelling of historic species-group names in Gunnerus, 1768 () with respect to the established requirement of the International Code of Zoological Nomenclature (hereafter the Code) that the suffix spelling of a Latin or Latinized adjectival species-group name must agree in gender with its genus (ICZN 1999: Article 31.2). The stability of names is important for tracking name usages as it is the largest serpulid genus, currently with 107 non-synonymised names, and also a further 31 original names currently placed as subjective synonyms. includes (Haswell, 1883), a model organism for settlement and genetic study (e.g., Hadfield 1998), and some other economically important species such as Okuda, 1934, (Verrill, 1873), Mörch, 1863, and Krøyer [in] Mörch, 1863, which are foulers of ship hulls, harbour structures, and aquaculture equipment (Sun et al. 2015).

species are easily recognisable by the morphology of the plug which closes the mouth of their calcareous tubes. It is a two-tier operculum with a basal funnel and a distal spinous structure called the verticil. The distinctive and varied structure of the verticil spines has enabled many species to be detected. Identification from tubes alone is problematic, thus past diversity is difficult to determine from the fossil record in the absence of the opercula. has no current subgenera, but Philippi 1844, the most prominent junior synonym of , and the little-used Mörch, 1863, were both at times used as subgenera defined by verticil morphology. The taxonomic history is reviewed in Bastida-Zavala and ten Hove (2002).

We recently realised that the gender agreement serpulid taxonomists had been applying to adjectival species-group names for the last 16 years was the opposite of that required by the fourth edition of the Code (ICZN 1999, effective 1 January 2000), in that authors since that date had continued using or creating masculine instead of feminine Latin forms. Additionally, taxonomists had not consistently observed the different Code rule applicable prior to 1999, with one author producing new names seemingly of both genders in the same publication (Straughan 1967a). Therefore we have compiled a checklist of name spellings we believe are currently correct, including also identifying the names not subject to gender agreement.

The derivation of as an genus name is known. Stearn (1983: 266) explains that substantives derived from -oides were commonly used for new genera before and during Linnæus’ era to indicate resemblance to a genus already known, sometimes just as temporary names. The -oides suffix is originally adjectival, transliterated from Greek οειδης. Gunnerus had at first used the cnidarian genus of (named for the creature of Greek myth) as the genus name for his new tube-PageBreakdwelling worm in 1766 correspondence with Linnæus, but shortly thereafter changed it to for his new species as published in 1768, “until Mr. v. Linné makes its genus known” (Gunnerus 1768, Moen 2006). The link to hydrozoans was spurious, but the genus name persisted. Usage of the same spelling applied to true hydrozoans persists in Romance language works, where “hydroïdes” (hydroids) can substitute for the formal higher group name, . The pair of usages are not homonyms in the strict Code sense, but there is some false positive reporting of the annelid genus in bioinformatics search results (e.g., from some of the works of Billard, such as Billard 1907).

was feminine because Gunnerus clearly treated it as feminine when he used the feminine ‘’ as epithet for the worm instead of the masculine ‘’ (likewise the calcareous tube was separately named by Gunnerus). However, in recent years Gunnerus’s original feminine gender assignment for became not obvious to most. This is exemplified by Moen (2006), who in the summary of her historical paper on Gunnerus reports without qualification that “in 1768 J. E. Gunnerus first described the species ”. Although Moen was well aware Gunnerus did not use that spelling she perhaps believed the ‘correct’ masculine ending (although incorrect since 2000) was always to be used, regardless of what was originally written.

The Code in its first edition indicated that genus names ending in -oides were to be treated as masculine (ICZN 1961: 33, Article 30(a)(ii) Examples), whereas in botany they were treated as feminine (Stearn 1983: 265). By the time of the Code third edition (ICZN 1985: 30, Article 30(b) Examples) the article text was the same, with the examples text explaining that these masculine genus names were substantivated adjectives, thus for the adjectival descriptive of ‘hydra-like’ was functioning as a noun. Mandatory gender agreement, although much debated, was retained in the Code fourth edition, but changes were made in order “to simplify the identification of gender in genus-group names” (ICZN 1999: XXVI).

Unexpectedly one of the qualifying clauses now included in the Code fourth edition (ICZN 1999) had a major effect on Gunnerus by reverting it to feminine status after almost 40 years as the opposite gender. The wording of Article 30.1.4.4 in full is “A compound genus-group name ending in the suffix -ites, -oides, -ides, -odes, or -istes is to be treated as masculine unless its author, when establishing the name, stated that it had another gender or treated it as such by combining it with an adjectival species-group name in another gender form.” Why the Code editorial group thought the refinement was necessary is unknown, but presumably it was regarded as better matching contemporary practice with the original 18–19th century usages.

began as feminine in 1768, and feminine adjectival endings matching this were usual for over 220 years but not universal (e.g., Pixell, 1913). Hartman (1965: 79) had maintained original feminine endings in her supplementary world catalogue, although not long later she had used the masculine for Hartman (Hartman 1969). Masculine endings, which the Code had required from the early 1960s onwards, otherwise only became common around 1992 (Moen 2006: PageBreak121), although Imajima, 1982 and Imajima & ten Hove, 1989 were newly described somewhat earlier. Ben-Eliahu and ten Hove (1992: 37) correctly pointed out that the Code third edition (ICZN 1985) had as masculine (actually in place since the first edition). Serpulid taxonomists then adopted the use of masculine endings and continued with this right up to August 2015, unaware of the change back to feminine required from 1 January 2000 when the new Code came into effect. The fourth edition Code was incorrectly cited as continuing masculine endings in ten Hove and Ben-Eliahu (2005: 128). In summary, for nearly 40 years species names in were required to have masculine endings according to the ICZN Code, although largely ignored for about 30 years, and now for the last 16 years they were required to have the feminine ending as begun by Gunnerus, also ignored. The conflict was first reported in WoRMS by one of us (GBR) in July 2015, after a misinterpretation of the Code requirement for names was published in Tovar-Hernández et al. (2016, as first online July 2015, p.8). Gender-corrected names were subsequently used in Sun et al. (2015), Kupriyanova et al. (2015), and Sun et al. (2016).

Code Article 31 (ICZN 1999) explains some of the requirements and exceptions regarding species-group Latin name formation. In general, most species-group names ending in the suffixes -us, -a, -um are declinable and likely to be adjectives. There are some exceptions applicable here such as -spina, which is a noun in apposition and should not change with gender, and most other name endings will not change. The only possible endings of changeable adjectives are -a, -us (these two make up over 60% of all names), -is, -um, -e, -er, -ior, whereas nouns can have all endings (Welter-Schultes 2013). Personal names as species-group names are usually (exceptions) formed as genitive-case nouns (ICZN 1999, Article 31.1). Nouns with Latin adjectival suffixes can become adjectives, notably non-Latin place-names with the suffixes -ensis (masculine/feminine) or -ense (neuter), indicating of that place, or suffix ‘-anus -a’, indicating belonging to. However, nouns compounded with dictionary Latin adjectives are treated as noun phrases in apposition (ICZN 1999, glossary).

Here we present an update of all non-synonymised names, and additional information on nomenclature, as derived in conjunction with the database (Read and Fauchald 2016), where further details are available. Certain species inquirenda (seven names which are otherwise valid but require taxonomic clarification due to inadequate original descriptions) are included and examined in the checklist for analytic purposes, while noting (as explained by ICZN 1999, Article 23.9.6) that the inclusion of these names should not later be considered as new usages.

World Register of Marine Species

We have taken this opportunity to investigate type localities of all the species, and to geolocate them to modern standards if possible. Prior to satellite-based navigation only vessel-based collections were likely to provide type-locality geolocations, and the descriptions used to pinpoint coastal sites could be vague or problematic. For instance Treadwell (1939: 164) gave an update that the “precise position” of his Mayagüez Bay station 6062 of 1902 could be relocated based on using a red buoy at the harbour PageBreakentrance as a reference point, but we are doubtful of the 100 year longevity of this buoy. We have been able to suggest placements for at least three species for which only a vague location was previously available. Type localities are mapped to show the world distribution.

Methods

The checklist is based on a review of original literature for all species-group names, and a review of about 250 -related name records at WoRMS (Read and ten Hove 2016). Under Code Article 34.2 (ICZN 1999), prior usages in literature are not required—here we simply formulate and present correct spellings. Gender agreement is mandatory, which means that non-agreeing scientific names strictly do not exist as valid spellings, and can be updated without explanation (to the bewilderment of many in the past, so we strongly advise annotation of new gender-spelling variants to avoid uncertainty). It is also worth noting that, while gender-agreement variants obviously are minor spelling changes, the Code is worded so that these different spellings are not treated as separate usages under prevailing usage rules.

All original literature for names was examined. Names as given are our derivations of correct endings for gender agreement, and are followed as necessary with the original binominal combination and comments on current status. The etymology (author’s and/or interpreted dictionary entries) is given, followed by our evaluation of the type of name (adjective, invariant noun in apposition, etc.) from available information. The derivation of names is unambiguous when authors give full etymologies, but this is rare for old names, and often sketchy for modern ones. Derivations are frequently only evident by matching likely character states mentioned, and occasionally there seems no obvious basis for the name chosen. The sources we have used to study derivations include online dictionaries and meta Greek/Latin language resources (Harper’s Etymonline; Logeion; Lexilogos), the Lewis and Short (1891) Latin dictionary (print, also online), as well as analytical dictionaries on the classical languages in science (Brown 1956; Stearn 1983). We have included the current subjective synonyms at WoRMS (if any) of each name (and their type localities), but have not included the other superseded recombinations, nor any misspellings of the valid name (these are fully listed at WoRMS and links to the current and original name records at WoRMS are in the Suppl. material 1 which also summarises the name analysis).

The type locality names have been investigated and their geolocations are included, usually derived by retrospective georeferencing. They are mapped to place the original discoveries in a geographic context and to locate where topotype material could be sought. Current place-names were geolocated using several web-based gazetteers (e.g., GeoNames, GEOLocate, Marineregions (WoRMS), Wikipedia). Disused historic names were sought via general web searches and Wikipedia. Holotype georeference PageBreakinformation in online collections databases and in subsequent publications was evaluated if available (these data can be based on retrospective approximations, rather than information supplied by authors on labels). Occasionally modern authors have published geolocations that are obviously imprecise or displaced, and we have pointed these out. The point-geolocations of the older taxa are our informed coastal assignments (indicated as map estimates) if derived from place-names which are towns, islands, or occasionally only known as strips of coast or other imprecise geographic extents. Sometimes positioning was assisted by further information from or about authors. A few times we were unable to narrow the collection site to any point and we indicate when we have given a general geolocation instead. We are unable to calculate the uncertainties (in extent) of our derived coordinates, and caution that each is a precise point location of the possible site, the nearest logical coastal geolocation at this time, as adjusted with satellite image overlay of terrain using the Wikipedia GeoLocator mapping tool. A list of geolocations is in the Suppl. material 1.

Results

Checklist of species original names

Family

Type species. Gunnerus, 1768 (original binomen)

Includes Philippi, 1844, type species Philippi, 1844 (by subsequent designation), Mörch, 1863 (as ), type species uncertain (full synonymy in WoRMS)

(original binomen)

Etymology: The author states the name for is derived from Latin adamas ‘diamond’ in reference to the diamond-shaped appearance of the verticil spines “although their distal ends are curved inwards”. The suffix -formis ‘shaped’ is used to form an adjective.

Evaluation: Masculine/feminine invariant adjective (formis, neuter forme) (Stearn 1983: 94).

Type locality: Lucas Island (south west corner), near Dampier Archipelago, Kimberley region, Western Australia.

Geolocation: (author, but is east of Lucas).

WoRMS: 555194

Synonyms: No subjective synonyms.

Status: Name now disused and representing a species inquirenda possibly senior to (Iroso, 1921).

Etymology: Not stated, but is named from the Latin adjective ‘related to’.

Evaluation: Masculine/feminine invariant adjective ( -e) (Stearn 1983: 94).

Type locality: Golfe de Marseille, France, Mediterranean Sea. No further precision, but likely to be coastal close to Marseille, possibly at or near Île Ratonneau, which the author mentions frequently (Marion and Bobretzky 1875). However, a stone pier off Arenc, Marseille is also mentioned (as habitat).

Geolocation: (map estimate, Île Ratonneau).

WoRMS: 383237

Synonyms: As species inquirenda has no synonyms although has been suggested (Zibrowius 1971: 713–714).

Etymology: The author states that is named for “limbations that are to be found on the sides of the spines of the distal opercular circlet”, thus combining the Latin adjectives alata ‘furnished with wings’ and lateralis ‘lateral’.

Evaluation: Masculine/feminine invariant adjective (lateralis -e) (Stearn 1983: 94).

Type locality: Port Royal, Jamaica, Caribbean Sea.

Geolocation: (map estimate).

WoRMS: 369228

Synonyms: No subjective synonyms.

Etymology: Not stated, but may be named for the operculum album mentioned by Grube (1870: 521) by combining the Latin adjective albus ‘white’ with -ceps derived from Latin noun caput ‘head’.

Evaluation: Noun in apposition (cf. noun ‘quadriceps’), or if treated as adjectival -ceps endings are invariant.

Type locality: ‘Tor’ (El Tor), Gulf of Suez, Red Sea.

Geolocation: (map estimate).

WoRMS: 130997

Synonyms: Pillai, 2009 (Fenelon Island, Kimberley, Australia)

Fischli, 1903 (Ternate, Indonesia)

Etymology: The authors state the name for is in honour of the Australian Museum Research Institute (AMRI).

Evaluation: Invariant non-Latinized noun in apposition ‘’ from an acronym, pronounced as a single word, not letter by letter as if an initialism (ICZN 1999, Article 11.3).

Type locality: Bass Point south, south of Wollongong, NSW, Australia.

Geolocation: (authors).

WoRMS: 852781

Synonyms: No subjective synonyms.

Etymology: Not stated, but may be named from Latin nouns ancora ‘anchor’, spina ‘thorn’, referring to the fact that both radii and verticil spines have anchor shaped tips.

Evaluation: Invariant noun in apposition.

Type locality: Wellawate, Colombo, Sri Lanka.

Geolocation: (map estimate).

WoRMS: 328434

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after one of its collectors, Arnold Schultze.

Evaluation: Invariant genitive noun from modern personal name of Arnold.

Type locality: Reported as two worms from two collection sites, Lome, Togo and Isla Annobón, Equatorial Guinea, both in Gulf of Guinea, West Africa. However, only the Annobón occurrence remains in (see WoRMS for further explanation).

Geolocation: (Annobón, map estimate).

WoRMS: 338000

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after Hermann Augener.

Evaluation: Invariant genitive noun from modern personal name of Augener.

Type locality: “Malembe” in Zaire (now Democratic Republic of Congo), but a coastal instance of the name could not be found, either in DR Congo or its neighbours. Zaire (DR Congo) has a very narrow access to the coast, and a coastal georeference was derived from “Vista”, the only other named collection site.

Geolocation: (map estimate for Vista).

WoRMS: 328435

Synonyms: No subjective synonyms.

Etymology: Not stated, but evidently is named after the Azores archipelago where collected.

Evaluation: Latinized adjectival form with correct feminine ending. usages exist (e.g., Bellan 2001).

Type locality: On shipwreck “Doria” east of Ponta Delgada port, Ilha de Sao Miguel, Açores (Azores).

Geolocation: (map estimate).

WoRMS: 328436

Synonyms: No subjective synonyms.

Etymology: Not stated, but evidently is named after the Banda Islands where collected.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from non-Latin geographic name Banda.

Type locality: Banda Islands (exact location unknown), Banda Sea, Indonesia.

Geolocation: (gazetteer, for Banda Islands).

WoRMS: 369229

Synonyms: No subjective synonyms.

Etymology: The author named after biologists Albert H. (Hank) and Dora May (Dee) Banner.

Evaluation: Invariant plural genitive adjective from Banner family name.

Type locality: Near Banners Point (Kalaeloa) sewage outfall, near Pearl Harbour, Oahu, Hawaii, Pacific Ocean.

Geolocation: (map estimate).

WoRMS: 328437

Synonyms: No subjective synonyms.

Status: The synonymy of and Straughan, 1967a with Treadwell, 1929 was re-confirmed by Sun et al. (2015: 63), but is being re-examined, and we provisionally include the original name analysis separately.

Etymology: Not stated, but the compound name for means ‘spiny-pedestal’ as derived from Latin (originally Greek) feminine noun basis ‘pedestal’ and adjective -a -um ‘spiny’. Basal internal spinules on opercular spines are mentioned (not figured).

Evaluation: Gender-variable adjective (in practice). Elsewhere in the article Straughan used feminine adjectival new species names, and in relation to the basal spinules Straughan probably intended another adjectival compound name. Her error in gender ending can be corrected to ‘-spinosa’. However, she used the feminine Latin noun ‘basis’ (pedestal), not the adjectival ‘basalis’ (basal) which would have become ‘basalispinosa’. If a noun phrase with a feminine noun then ‘’ was incorrect Latin (it should also have been ‘’), and the original spelling must be maintained (ICZN 1999, Article 31.2.1). While this can be noted, Straughan is not the only author to adopt ‘basis’ as if adjectival, and it seems best not to apply the strictest interpretation here. Usage as ‘’ already exists (e.g., Sun et al. 2015: 63).

Type locality: Mouth of Ross River, Townsville, Queensland, Australia.

Geolocation: (map estimate).

WoRMS: 881640

Synonyms: See comments.

Etymology: Not stated, but the name for is adjectival from Latin furcatus ‘forked’, likely referring to the bifid verticil spines.

Evaluation: Gender-variable adjective. The original incorrect masculine ending as , repeated in Day (1951: 64), was silently corrected to ‘’ in Day (1967: 808).

Type locality: Minicoy/Maliku (as Minikoi), south Lakshadweep archipelago, north of the Maldive Islands.

Geolocation: (map estimate).

WoRMS: 873900

Synonyms: No subjective synonyms.

Etymology: Not stated, but is likely named based on ‘bisected’, a New Latin past participle used as an adjective, derived from Latin bis ‘two’, secare ‘to cut’, and referring to the bifid tips of verticil spines.

Evaluation: Gender-variable adjective, corrected herein to ‘’.

Type locality: off Sesoko Marine Station, Sesoko Island, Okinawa Islands, Japan.

Geolocation: (map estimate).

WoRMS: 880526

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is likely referring to the pair of lateral spinules on the verticil spines described by Bush, based on Latin bis ‘two’ with adjective ‘spined’. Bush compared with .

Evaluation: Gender-variable adjective with correct original feminine ending. Usage as ‘bispinosus’ exists (e.g., Bastida-Zavala and ten Hove 2002: 125).

Type locality: Bermuda. The Yale Peabody Museum type (syntype? YPM IZ 001367.AN) from the Verrill Bermuda Expedition in 1898, evidently has no further location data, but Castle Harbour is a collection site mentioned by Bush (1910).

Geolocation: Imprecisely known (possible place of origin, Castle Harbour, , Bermuda).

WoRMS: 421083

Synonyms: No subjective synonyms.

Etymology: Not stated, and the description of does not indicate why the name derives from Greek βραχυ (brachy) ‘short’, ακανθα (akantha) ‘spine’, feminine noun, thus short-spine. In New Latin acantha has frequently been used as part of feminine compound names in both genera and species-group names. An identical spelling might be expected to be a noun form in both, but species-group names ending as -acantha -acanthus have regularly been treated as Latinized Greek adjectives, and that may have been the intention of the author.

Evaluation: Gender-variable adjective with correct feminine ending. Usages as ‘brachyacanthus’ exist (e.g., Bastida-Zavala and ten Hove 2003: 73).

Type locality: Marina Mazatlán, Mazatlán, Sinaloa, Gulf of California, Mexico.

Geolocation: (original author, with neotype of Sun et al. (2016: 49) from the same geolocation).

WoRMS: 328441

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named for the bulbous (Latin -a -um) dorsal verticil spine.

Evaluation: Gender-variable adjective, corrected herein to ‘’.

Type locality: Khor Ghubb ’Ali, Musandam Peninsula, Oman, Strait of Hormuz, in a sheltered bay at 18 m.

Geolocation: (map estimate).

WoRMS: 882354

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is a compound noun which may be referring to the operculum, from Greek καλος (kalos) ‘beautiful’, πωμα (poma) ‘lid’.

Evaluation: Invariant noun in apposition (indeclinable because ending in a transliterated Greek word).

Type locality: Isla Tortuga, off Isla Annobón, Equatorial Guinea, Gulf of Guinea.

Geolocation: (map estimate).

WoRMS: 369230

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after the Cape Provinces of South Africa.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from non-Latin geographic name.

Type locality: Offshore from Lambert’s Bay, north of Cape Town, western coast of South Africa.

Geolocation: (author).

WoRMS: 338003

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after the country of collection.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from non-Latin geographic name.

Type locality: Arica (coastal port city), Chile.

Geolocation: (map estimate).

WoRMS: 328444

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is likely referring to the verticil spines, which are cross-bearing, from feminine Latin noun crux, crucis ‘cross’, with Latin suffix ger, gera ‘to bear’.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as incorrect suffix ‘’ and as masculine ‘cruciger’ exist (e.g., de León González 1990: 336, Bastida-Zavala and ten Hove 2003: 78). Names ending in -ger may be nouns or masculine adjectives (ICZN 1999, Article 31.2.2). The usage of Mörch was adjectival as he used feminine -gera.

Type locality: Puntarenas, Gulf of Nicoya, Costa Rica Pacific coast (Mörch 1863 as “oceano pacifico, juxta Puntarenas”).

Geolocation: (map estimate).

WoRMS: 333637

Synonyms: [sic] Treadwell, 1929 (“Lower California” (Baja California) Mexico)

Etymology: Not stated, but as “trouvé par Mr. Dafni” is evidently named after the collector, Yaacob Dafni (Amoureux et al. 1978: 60, 148).

Evaluation: Invariant genitive form of the modern personal name Dafni.

Type locality: Eilat, Gulf of Aqaba (of Eilat), Israel, Red Sea, on reef coral. The site is mapped by the authors, but not georeferenced.

Geolocation: (map estimate).

WoRMS: 369231

Synonyms: No subjective synonyms.

Etymology: The authors state is named after Jesús A. de León-González.

Evaluation: Invariant genitive form constructed from the personal name de Leon.

Type locality: Punta San Juanico, Western coast of Baja California Sur, Mexico. Authors’ georeference (, inland, ~26 km off target) is herein corrected to ).

Geolocation: (San Juanico, map estimate).

WoRMS: 328445

Synonyms: No subjective synonyms.

Etymology: Verrill (1873: 28) states for that the name alludes to the resemblance to flowers as the colours of its branchiae “recalls the varied hues and forms of different kinds of pinks, (.)”. The botanical generic name (flower of Zeus) is New Latin (Linnæus and earlier) from Greek Δηοσ (Dios), genitive of Zeus, and ανθος (anthos) ‘flower’. As is feminine and is masculine it seems Verrill intended the name as a noun ( Ellis, 1768 is an earlier similar pairing).

Evaluation: Invariant noun in apposition.

Type locality: Great Egg Harbor to New Haven and Cape Cod, Atlantic coast USA.

Geolocation: Unknown (New Haven, , as a central possible place of origin on Atlantic coast USA).

WoRMS: 131000

Synonyms: Possibly sensu Pratt, 1916 and others [non Bosc, 1802]

Verrill, 1873 (for Verrill’s colour variant specimens)

Augener, 1922 [partim, fide Bastida-Zavala and ten Hove 2002: 143) (Haiti, Caribbean Sea)

Status: Name now disused and representing a species inquirenda. A single subsequent valid usage of the name was later identified as an occurrence of (a junior name), but it is uncertain that Grube’s original serpulid (type missing) was the same (fide Zibrowius 1978: 144; Sun et al. 2015: 37).

Etymology: Not stated, but the name for derives from Greek Latinized as diplos ‘two-fold’ and feminine Greek noun χοάνη (choani) ‘funnel’, thus double funnel, evidently in reference to the two-tier operculum that Grube describes (a generic character). There are no other names based on chone in , but it is part of several generic names in .

Evaluation: Invariant noun in apposition.

Type locality: Askold Island, outer Peter the Great Gulf, North Japan Sea. We infer this to be the type locality. Grube does not present location information beyond that the material was from “nordjapanischen Meeres”, but it is also mentioned that the collector was the Siberian-based Polish naturalist Dybowski, whose travels in the region are documented. In 1874 Benedykt Dybowski collected fauna at Askold Island, near Vladivostok, Primorsky Krai (fide Zoological Museum, University of Lliv [no date]).

Geolocation: (map estimate, Askold Island).

WoRMS: 333639

Synonyms: As species inquirenda has no synonyms, although has been suggested.

Etymology: Not stated, but in the description for Schmarda (1861: 29) describes in Latin “Operculum duplex infundibuliforme” (double funnel lid) and in German “Das Thier hat zwei Deckel” (has two lids), evidently referring to the two opercula figured in his plate, and based on Greek δις (dis) ‘twice’, and πωμα (poma) ‘cap’ (see ten Hove and Ben-Eliahu (2005) for an analysis of records of bi-operculate specimens in ).

Evaluation: Invariant compound noun in apposition (indeclinable because ending in a transliterated Greek word).

Type locality: Cape of Good Hope, South Africa (“Vorgebirge der Guten Hoffnung”).

Geolocation: (Cape of Good Hope (gazetteer), although Schmarda more likely was indicating a general coastal area).

WoRMS: 369232

Synonyms: Pixell, 1913 (Gulf of Suez)

Ehlers, 1913 (Simonstown, False Bay, South Africa)

Etymology: Not stated, but for Mörch describes “utrinque inflexione obsoleta, unde lateraliter adunco-rostrato” (rudimentary bend both sides, hence laterally-curved beak), with the Latinization evidently referring to the twin sharp lateral points of the blunt tip of the verticil spines, ultimately from Greek δις (dis) ‘twice’, PageBreakand ραμφος (rampos) ‘beak’, the latter modified through New Latin masculine noun forms rhamphus and the lesser-used ramphus (both with a number of usages as part of compound genus names) to rampha, a usage seemingly unique to Mörch.

Evaluation: Treated here as an invariant noun in apposition, because an incorrect Latinization. While Mörch consistently modified his names as feminine, and the name seems intended as feminine adjectival rather than a noun, it looks like a misspelled Latinization which should be left unaltered. Usages in as ‘diramphus’ exist (e.g., Bastida-Zavala and ten Hove 2002: 161).

Type locality: Saint (St.) Thomas Island, United States Virgin Islands, Lesser Antilles (“in portu urbis St. Thomae Antillarum”), most likely the Saint Thomas port town of Charlotte Amalie.

Geolocation: Imprecisely known (near to if harbour at Charlotte Amalie, St Thomas Island).

WoRMS: 131001

Synonyms: Bush, 1910 (Bermuda, western Atlantic)

Claparède, 1870a (Gulf of Naples, Italy, Tyrrhenian Sea)

Mörch, 1863 (Bahia coast, Brazil)

Mörch, 1863 (Philippines unspecified)

Mörch, 1863 (New Zealand unspecified)

[sic] Rioja, 1942 (Mazatlán, Gulf of California, Mexico Pacific coast)

Etymology: The authors state for that “ is from the Latin dolabra, a sort of pickaxe that resembles the shape of the verticil spines”.

Evaluation: Invariant noun in apposition. The Latin dolabra is a feminine noun, and cannot become a masculinised adjective as ‘’ to match a masculine . This is not a word as listed in classical Latin dictionaries, and should be considered as an invariant combination of letters. A suitable adjectival equivalent would have been dolabratus -ata.

Type locality: Mazatlan Marina, Mazatlan, Gulf of California, Mexico Pacific coast.

Geolocation: (authors).

WoRMS: 851651

Synonyms: No subjective synonyms.

Status: The much-used name is nomen protectum with respect to nomen oblitum H. Krøyer [in] Mörch, 1863 (Bastida-Zavala and ten Hove 2002).

Etymology: Not stated, but the name is likely derived from the Latin adjective -antis (genitive) ‘elegant’.

Evaluation: Invariant adjective (masculine/feminine/neuter ‘-ans’).

Type locality: Port Jackson, NSW, Australia (not further specified).

Geolocation: (gazetteer).

WoRMS: 131002

Synonyms: Krøyer [in] Mörch, 1863 [nomen oblitum] (Saint Croix island, Virgin Islands, Caribbean Sea)

Monro, 1938 (Shoreham Harbour Canal, Sussex, England)

Hartman, 1969 (Velero station 1452-42, Ship hull & pier, Long Beach, California)

Straughan, 1967a (Shoal Point, Mackay, Queensland, Australia)

Etymology: Not stated, but the name is likely derived from Latin adjective -antis ‘elegant’, combined with the Latin suffix -ulus, a male diminutive adjectival form.

Evaluation: Gender-variable adjective, corrected in to feminine ‘’ by Zibrowius (1971: 695).

Type locality: Bermuda. The Yale Peabody Museum holotype YPM IZ 001323.AN from the Verrill Bermuda Expedition in 1898, evidently has no further location data, but Castle Harbour is a collection site mentioned by Bush (1910).

Geolocation: Imprecisely known (possible place of origin, Castle Harbour, , Bermuda).

WoRMS: 873929

Synonyms: No subjective synonyms.

Status: Name now disused and representing a species inquirenda that has been compared to Zibrowius, 1968. It is not a candidate nomen oblitum (used as valid by Zibrowius 1972c: 116–117). If suppression is desirable prevailing usage of would be maintained (ICZN 1999, Recommendation 23A).

Etymology: Not stated, but was evidently named after the Latin name for Caiola Island, Naples, where Delle Chiaje states it was collected. Caiola is modern day Gaiola, in Roman times known as Euplaea. The name as combined with feminine adjectival suffix -ana indicates from Euplaea.

Evaluation: Gender-variable geographical Latin adjective (-anus, -ana) from place-name, with correct original feminine ending.

Type locality: Caiola (Gaiola/Euplaea) Island, Naples, Italy, Tyrrhenian Sea, Mediterranean.

Geolocation: (map estimate).

WoRMS: 381073

Synonyms: As species inquirenda has no synonyms, although has been suggested.

Etymology: Not stated, but Marenzeller (1885: 217) described the character of the opercular verticil spines for as being elevated on a central column (“einer centralen Säule”). The Latin ‘(up) lifted’ species-group name is perhaps in reference to this.

Evaluation: Gender-variable adjective, corrected to feminine ‘’ as recombined (e.g., Imajima 1976b: 232). Usages in as ‘’ exist (e.g., Dew 1959: 27).

Type locality: East coast of Enoshima Island (“Ostküste der Insel Eno-sima”), Sagami Bay, Honshu, Japan. There is an Enoshima-rettō Island, also off Honshu, but the Sagami Bay Enoshima Island (only ~0.5 km long) is the most likely visited.

Geolocation: (map estimate).

WoRMS: 873938

Synonyms: No subjective synonyms.

)

Status: Name now disused and representing a species inquirenda. Similarities of the original description to have been noted, but the name is yet to be synonymised.

Etymology: Not stated, but was evidently named for its vesicular dorsal verticil spine.

Evaluation: Gender-variable adjective with incorrect original ending, corrected to feminine (e.g., Monro 1937: 316).

Type locality: Gatavaké (Baie de Gatavaké), Mangareva Island, Mangareva/Gambier Islands, French Polynesia, South Pacific.

Geolocation: (map estimate).

WoRMS: 875068

Synonyms: As species inquirenda has no synonyms.

Etymology: Not stated, but for it is likely the Latin spina ‘thorn’ refers to the external (curved outwards) spines of the verticil.

Evaluation: Invariant noun in apposition.

Type locality: Heron Island, Queensland, Australia, collected close to the marine station by Dew (map in Straughan 1967b).

Geolocation: (map estimate).

WoRMS: 328446

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after its area of collection, as Ezo (also as Yezo) is a former name for the island of Hokkaido, Japan. The species-group name ‘’ has often been used for Japanese taxa, along with ‘yezoensis’.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from non-Latin geographic name.

Type locality: Not fixed by author. Original records are from “Akkeshi, Muroran, and Oshoro”, which are widely separated places around the coast of Hokkaido Island, with the first two having marine stations.

Geolocation: Imprecisely known (possibly as for Akkeshi Marine Station).

WoRMS: 131003

Synonyms: See comments for .

Etymology: Not stated, but is evidently named after its purported region of collection. ‘Florida’ is Spanish for flowery land and is here combined with the Latin adjectival suffix -anus -a -um, indicating from Florida.

Evaluation: Gender-variable adjective based on a non-Latin geographic name, corrected to feminine herein. Usages in as ‘’ exist (e.g., Bastida-Zavala and ten Hove 2002: 118) but not previously as ‘’.

Type locality: Unknown, not certain to be off namesake Florida. When Ehlers (1887: 286) described the collected from voyages of Coast Survey Steamer “Blake” he wrote in his native German but recorded the two locations for the specimens literally in English as “inside fishing ground Cape Rear” and also “off W. down Cape Dear Rio” (both at 7 fathoms). However, these place-names seem to be misreadings as they could not be found in the Caribbean or Florida, nor do the “Blake” voyage reports include the names. The similarity of names suggests the location is possibly off Cape Fear, North Carolina, with its associated Cape Fear (Rio) River, disregarding that Ehlers’ monograph title appears to exclude Atlantic coast voyages the “Blake” also made. As the types are believed lost the original label cannot be checked. No specimens are currently listed in the Yale Peabody Museum online catalogue although Bush (1910: 498) earlier saw a mass of several hundred tubes, indicating an aggregation.

Geolocation: Unknown (if off Cape Fear, North Carolina then that place-name is at (gazetteer) ).

WoRMS: 369234

Synonyms: Treadwell, 1931 (Grand Isle, Louisiana, Gulf of Mexico)

Iroso, 1921 [junior objective synonym (same specimen)]

Etymology: Not stated but Grube described for forked spines in the opercular funnel as well as the verticil, thus Latin furca ‘fork’, combined with adjectival PageBreaksuffix -fer -a -um ‘bear’. Lewis and Short (1891: 795) include as a feminine noun meaning phallus, but it is unlikely this was Grube’s intention. A more common adjectival form would be furcillata ‘forked’.

Evaluation: Gender-variable adjective with correct original feminine ending. Names ending in -fer may be nouns or masculine adjectives (ICZN 1999, Article 31.2.2 example). The usage of Grube was adjectival as he used feminine -fera. A listing-only usage in as ‘furcifer’ exists (ten Hove and Kupriyanova 2009: 53).

Type locality: “Ubay, Pandanon”, Philippines. Ubay () is a small port on Bohol Island, and Pandanon Island () is a small reef ~45 km to the west of Ubay.

Geolocation: (map estimate, Ubay).

WoRMS: 369235

Synonyms: [sic] Imajima, 1982 (off Arumonogui, Palau Islands, Micronesia)

Etymology: Not stated, but the species-group name for from Latin fuscus ‘dark’ is evidently in reference to the “glossy black” verticil spines.

Evaluation: Gender-variable adjective with correct original feminine ending. A listing-only usage in as ‘fuscus’ exists (ten Hove and Kupriyanova 2009: 53).

Type locality: Offshore east off northern tip of Tanegashima (island), Southern Japan, 80 m.

Geolocation: (map estimate from author’s map).

WoRMS: 369236

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after the gastropod genus (now ) combined with -cola ‘dweller’, as it was found attached to a ‘Fuso’ sp.

Evaluation: Invariant compound noun in apposition with -cola as a substantival suffix. The Code has a stipulation (ICZN 1999, Article 30.1.4.2) that genera with -cola endings be treated as masculine compound nouns (or mostly so treated, similar to the -oides situation). It has no advice for species-group names with -cola suffixes, but they are recommended to be treated similarly (David and Gosselin 2002: 34), not declined to agree with the first noun or the genus.

Type locality: Japan (not further specified). Mörch only knew the specimen was from the collection of Wessel in Hamburg.

Geolocation: Unknown (gazetteer Japan central point as ).

WoRMS: 369237

Synonyms: Pillai, 1972 [nom. n. for “ (sensu Okuda et Uschakov)”] (location not fixed by author, but the Okuda (1937: 63) usage was for Ishihama, Japan, a name for at least four possible Honshu coastal locations)

Status: Candidate nomen protectum against senior name Mörch, 1863 (fide Bastida-Zavala and ten Hove, 2002: 132). Prevailing usage maintained pending proof of sufficient usage of , but is a nomen oblitum, not used as a valid name after 1899 (listings are excluded as usages under ICZN 1999, Article 23.9.6).

Etymology: Not stated, but is evidently named after its place of collection, Gairaca.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin place-name.

Type locality: Gairaca, near Santa Marta, Caribbean Sea coast of Colombia.

Geolocation: (map estimate).

WoRMS: 369238

Synonyms: Mörch, 1863 [nomen oblitum] (La Guayra, Panama, Caribbean Sea)

Status: Type taxon by monotypy of Rioja 1941b, a synonym of .

Etymology: Not stated, but as Rioja (1941a: 174) writes of “una robusta protuberancia . . .en forma de glande” for , it is likely to be a functional name for the bulbous dorsal verticil spine from Latin glans ‘acorn’ combined with adjectival suffix -fer -a -um ‘bear’. It is unclear why Rioja (incorrectly) used the neuter form ‘’ at first, but he later (Rioja 1941b: 733) modified the spelling for his feminine combination.

Evaluation: Gender-variable adjective with corrected feminine ending. Usages in as ‘glandifer’ and ‘’ exist (e.g., Bastida-Zavala and ten Hove 2003: 89).

Type locality: Caleta (Playa Caleta), Acapulco, Mexico.

Geolocation: (map estimate).

WoRMS: 338016

Synonyms: No subjective synonyms.

Etymology: The authors dedicated to Christopher J. Glasby.

Evaluation: Invariant genitive form of the personal name Glasby.

Type locality: Fort Hill Wharf, Darwin, Northern Territory, Australia.

Geolocation: (authors, ).

WoRMS: 852813

Synonyms: No subjective synonyms.

Etymology: Not stated, but the Latin ‘slender’ name for is likely referring to the simple verticil spines.

Evaluation: Invariant adjective (masculine/feminine ‘’).

Type locality: Pacific Grove, California, Pacific coast USA.

Geolocation: (map estimate).

WoRMS: 333640

Synonyms: Chamberlin, 1919 (Laguna Beach, California coast)

Status: The synonymy of and Straughan, 1967a with Treadwell, 1929 was followed by Sun et al. (2015: 63), but is being re-examined, and we provisionally include the record separately.

Etymology: Not stated, but for the Latin ‘gradual’ is evidently describing the gradual size change of the ring of opercular spines.

Evaluation: Gender-variable adjective with correct original feminine ending.

Type locality: Pretty Beach, 40 km north of Cairns, Queensland, Australia

Geolocation: (map estimate).

WoRMS: 384604

Synonyms: See comments.

Status: Zibrowius (1971: 713–714) synonymised an older name, Marion, 1875, under . This is not possible on priority, nor does qualify as a nomen oblitum as it was used as valid (Zibrowius 1968: 115) post 1899. We include both names (see entry for as species inquirenda).

Etymology: Not stated, but is likely named after the larger helmet-like dorsal verticil spine as Iroso (1921: 54) describes “che ricade sugli altri ad elmo” (which falls on others [spines] helmet-like). Helm and helmet are not from Latin, though the author’s construction appears to be intended as adjectival, with adjectival suffix -atus added to mean helm-like.

Evaluation: Gender-variable adjective corrected to feminine in in Zibrowius (1971: 713).

Type locality: Unspecified Gulf of Naples (Golfo di Napoli), Italy.

Geolocation: Imprecisely known (a Golfo di Napoli mid-point (gazetteer) is ).

WoRMS: 131004

Synonyms: No subjective synonyms, but has been linked to (see above).

Etymology: Not stated but the name for is likely describing the dimorphism in verticil spines, from Greek hετεροσ (heteros) ‘different’ and κερασ (keras) ‘horn’. The Latinized is an adjectival form to be declined.

Evaluation: Gender-variable adjective corrected to feminine in Zibrowius (1971: 715). Grube (1868: 639) originally incorrectly created a masculine ‘’ in agreeing with the masculine subgenus rather than the feminine genus . Usages exist in as ‘’ (e.g., Ben Eliahu and ten Hove 2011: 26), and as the misspelling ‘heteroceros’ (e.g., Day 1967: 807).

Type locality: Unspecified Red Sea. Grube’s report title refers to Red Sea worms collected by Georg Ritter von Frauenfeld. Grube states in his opening sentence that the worms were handed to him without any other information, and it seems he did not investigate this further. In the narrative of his visit von Frauenfeld (1855) mentions Suez, the Sinai Peninsula, and seeing countless annelids on the Red Sea shore, but he does not match observation to locality.

Geolocation: Imprecisely known, but perhaps northern Red Sea (a gazetteer Red Sea mid-point is ).

WoRMS: 851900

Synonyms: No subjective synonyms. However, the misidentification non Philippi, sensu Gravier, 1906, has been assigned to (e.g., Pixell 1913: 75).

Etymology: Not stated, but evidently is named because there are two types of furcate verticil spines of the operculum (Pillai 1971: 114).

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘heterofurcatus’ exist (e.g., ten Hove and Kupriyanova 2009: 53).

Type locality: near Talaimannar Pier, Sri Lanka, 4 m depth.

Geolocation: (map estimate from author map).

WoRMS: 328449

Synonyms: No subjective synonyms.

Status: A name disused by taxonomists and representing a species inquirenda. The original description and figure are rudimentary and the species Bosc saw will remain indeterminable unless original specimens are found (unlikely). However, the name cannot be a nomen oblitum as it was revived as [sic] in three widely used manuals on invertebrates of the United States eastern coast (Pratt 1916, Grave 1937, Costello et al. 1957). These instances should be considered misidentifications, and might be referable either to the junior name (fide Zibrowius 1971: 697, Bastida-Zavala and ten Hove 2002: 108), or to other similar species. Nevertheless, there are multiple PageBreakmodern citations of the research on sperm (e.g., Colwin and Colwin 1961) in which the name appeared.

Etymology: Bosc described the tube of as “montrant la moitié d’un prisme hexagone …”, and the name is a New Latin adjectival form for six-sided, modified from Greek. Bosc’s figure shows two ridges so the tube cross-section would be trapezoidal, not literally hexagonal as named, but half (la moitié) of that.

Evaluation: Gender-variable adjective with correct feminine ending herein. Usages in as ‘’ and ‘hexagonis’ exist (e.g., Pratt 1916: 302, Grave and Oliphant 1930: 234) but not previously as ‘’.

Type locality: Charleston Harbour, Charleston, South Carolina, Atlantic coast USA.

Geolocation: (gazetteer).

WoRMS: 384606

Synonyms: As species inquirenda has no synonyms.

Etymology: Not stated, but for it is likely that the Greek hομοσ (homos) ‘uniform’ and κερασ (keras) ‘horn’, refers to the opercular verticil spines. Pixell appears to have named ‘’ as the opposite to ‘heteroceros’ (her error for the existing ‘’) which she mentions.

Evaluation: Incorrect Latinization treated here as an unchanging noun in apposition. Usages exist as ‘homocera’ (e.g., Ben-Eliahu and ten Hove 1992: 35) and ‘homocerus’ (e.g., Bellan 2001: 226).

Type locality: Multiple Indian Ocean localities as the syntypes (aggregated as only one NHM specimen lot 1924.6.13.147 received from the Cyril Crossland Collection) came both from the Maldive area (specified as Miladhunmadulu Atoll and Minikoi), and from off Zanzibar.

Geolocation: Unknown (map estimate for Noonu, the southern Miladhunmadulu Atoll).

WoRMS: 238212

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after the sea in which the worm was collected as “Huanghai” means Yellow Sea in Chinese.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin geographic name, Huanghai.

Type locality: Northern Yellow Sea, off the Chinese coast near Dalian.

Geolocation: (as authors, [? error for 7']).

WoRMS: 328450

Synonyms: No subjective synonyms.

Etymology: Not stated, but for possibly the name, from Latin ‘humble’ (or ‘low’), is referring to the small size of the single specimen collected.

Evaluation: Masculine/feminine invariant adjective ( -e) (Stearn 1983: 94).

Type locality: Guaymas, Gulf of California coast, Sonora state, Mexico. Bush provides no other details other than the name Guaymas (Mexico).

Geolocation: (map estimate).

WoRMS: 369239

Synonyms: No subjective synonyms.

Etymology: Not stated, but for it is likely that the Latin ‘unarmed’, is referring to the verticil spines without spinules. Monro stated the operculum “lacks spines both on the lower and the upper calix”.

Evaluation: Masculine/feminine invariant adjective ( -e).

Type locality: James Bay, Isla Santiago (was James Island), Galapagos, Ecuador.

Geolocation: (map estimate).

WoRMS: 338017

Synonyms: No subjective synonyms.

Status: The current synonymy of with is being re-evaluated, and meantime it is included separately here.

Etymology: Not stated, but for it is likely that the Latin inornatus ‘unadorned’ is referring to the verticil spines without side spinules.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘inornatus’ exist (e.g., Amoureux et al.: 57).

Type locality: Maha Alamba (not found, perhaps disused), “about a mile” from the Negombo Lagoon entrance (an aquatic research institute is nearby), north of Colombo, west coast of Sri Lanka.

Geolocation: (map estimate).

WoRMS: 338018

Synonyms: No subjective synonyms, and has been regarded as junior to (e.g., Sun et al. 2015: 62).

Etymology: The author named after the Kimberley region of Western Australia.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin geographic name.

Type locality: Off east side of Fenelon Island (main island of Institut Islands) at 6 m, Kimberley, Western Australia.

Geolocation: (author).

WoRMS: 555195

Synonyms: No subjective synonyms.

Etymology: The authors named after Hugh J.P. Lambeck (entomologist, deceased, one time assistant to ten Hove), who first noted this as a species different from .

Evaluation: Invariant genitive form of the personal name Lambeck.

Type locality: Vaarsenbaai (cove), Boca Sami, Curaçao, Netherlands Antilles.

Geolocation: (gazetteer).

WoRMS: 328452

Synonyms: No subjective synonyms.

Etymology: The authors named after the Australian Museum’s .

Lizard Island Research Station

Evaluation: Invariant non-Latinized noun in apposition ‘’ from an acronym, pronounced as a single word.

Type locality: Front of reef between Bird and South Islands, Lizard Island, Queensland, Australia, (station MI QLD 2354 in Ribas and Hutchings, 2015).

Geolocation: (station list).

WoRMS: 877990

Synonyms: No subjective synonyms.

Etymology: Not stated, but evidently is named after the “conspicuous, long central spine” (long in comparison with ), based on Latin adjectives longus ‘long’ with ‘spined’.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘longispinosus’ exist (e.g., Bailey-Brock 1987: 282).

Type locality: Koniya, Amami-Oshima, Amami Islands, Southern Japan.

Geolocation: (map estimate).

WoRMS: 328453

Synonyms: Wu & Chen, 1981 (Yulin Harbour, Hainan Island, South China Sea)

Etymology: Not stated, but for evidently the Latin longus ‘long’ and adjectival Latinization of Greek στυλος (stylos) ‘pillar’ refers to the long, elongated basis of the opercular funnel, thus ‘pillar-like’.

Evaluation: Masculine/feminine invariant adjective (-stylaris -e).

Type locality: Shellfish farms, Zhangpu (Zhangzhou), Fujian Province, China.

Geolocation: (map estimate).

WoRMS: 328454

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is evidently a compound noun from Latin malleolus ‘small hammer’, referring to the dorsal hammer-shaped verticil spine, and spina ‘thorn’.

Evaluation: Invariant noun in apposition. Usages as ‘’ exist (e.g., Murray et al. 2010: 393).

Type locality: Pialba, Hervey Bay, Queensland, Australia.

Geolocation: (map estimate).

WoRMS: 369240

Synonyms: “” [sic] Pillai, 2009 [unavailable name (no type-designation), assignment by Murray et al. 2010] (Australia)

Etymology: Not stated, but for the ‘micro’ derives from Greek μικρος (micros) ‘small’, and perhaps is combined with Greek neuter noun genitive οτοσ (otos) ‘ear’. The Latinizations ‘’ and ‘microtus’ are in use as both genus and species-group names for small-eared biota. Whether the same derivation applies for is unclear, as the verticil spines are knob-tipped and not notably small or ear-like.

Evaluation: Invariant whether a noun in apposition or (masculine/feminine) intended as adjectival.

Type locality: North America (unspecified) as “ad Americam borealem” on (was as ), collected by A. B. Mayer, presumably on the Atlantic coast as is the bay scallop of that region.

Geolocation: Unknown (unspecified Atlantic coast of North America, with the mid point of the coastal extent).

WoRMS: 333641

Synonyms: No subjective synonyms.

Etymology: Not stated, but for the Latin adjective -acis meaning ‘jutting out’ is likely referring to the enormous dorsal verticil spine.

Evaluation: Invariant adjective (masculine/feminine ‘’).

Type locality: Philippines (unspecified).

Geolocation: Unknown ( (gazetteer) is central to the Philippines Islands).

WoRMS: 131007

Synonyms: Gravier, 1906 (Bonhoure Recif, Djibouti, Gulf of Aden)

Etymology: The author named after Ricardo Monges López of Veracruz.

Evaluation: Invariant genitive noun from modern personal name of Monges López.

Type locality: On floating pumice, Playa Norte, Isla Santiaguillo, Veracruz, Gulf of Mexico.

Geolocation: (map estimate).

WoRMS: 328456

Synonyms: No subjective synonyms.

Etymology: Not stated, but the species is evidently named after C. C. A. (Charles Carmichael Arthur) Monro, who had studied the specimens earlier.

Evaluation: Invariant genitive noun from modern personal name of Monro.

Type locality: Pointe Noire, Congo, West Africa.

Geolocation: (map estimate).

WoRMS: 328457

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is evidently referring to the pointed (Latin mucronatus) side spines of the verticil spines “que tienen forma de mucron” (Rioja 1958: 256).

Evaluation: Correct original adjectival feminine ending. Usages as ‘mucronatus’ exist (e.g., Bastida-Zavala and ten Hove 2002: 141).

Type locality: Isla de Sacrificios, Veracruz, Gulf of Mexico.

Geolocation: (map estimate).

WoRMS: 328458

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for evidently refers adjectivally to multiple lateral spinules on the verticil spines.

Evaluation: Correct original adjectival feminine ending. Usages as masculine ‘multispinosus’ exist (e.g., ten Hove and Kupriyanova 2009: 54).

Type locality: Shore at Eno-sima (Enoshima), Sagami Bay, Honshu, Japan. There is an Enoshima-rettō Island, also off Honshu, but the Sagami Bay Enoshima is the most likely visited.

Geolocation: (map estimate).

WoRMS: 335316

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named broadly geographically as “Nanhai” is the South China Sea in Chinese.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin geographic area name.

Type locality: Xi River estuary, Pearl River Delta, Macao, Guangdong, China coast, South China Sea, 58m, fixed on rock, stations 6016, 6044 (fide Sun and Yang 2014: 218 (map), 241; no locality in the original text).

Geolocation: (map estimate, Xi River mouth).

WoRMS: 328459

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is evidently referring to the dark colour of the operculum, especially of the opercular constriction (“un anneau noir à la base de l‘opercule”) and the verticil spines, and derived from the Latin adjective niger, , nigrum ‘black’.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘niger’ exist (e.g., Bellan 2001: 226)).

Type locality: Tabarka “au large de l’ile [Tabarka] et de la Pointe Meloula [4 km west]”, Tunisia, Mediterranean Sea.

Geolocation: (map estimate for north end of Tabarka).

WoRMS: 328460

Synonyms: No subjective synonyms.

Etymology: The authors named after Nika Mikhin, daughter of Kupriyanova.

Evaluation: Invariant feminine genitive form of given name Nika.

Type locality: Edithburgh Jetty, Edithburgh, St Vincent Gulf, South Australia.

Geolocation: (adjusted to jetty from authors’ inland (should be 45') E).

WoRMS: 871949

Synonyms: No subjective synonyms.

Etymology: Not stated, but is likely named for the internal “rounded projection” at the base of each verticil spine, from the adjective nodosus -a -um ‘knotty’.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘nodosus’ exist (e.g., ten Hove and Kupriyanova 2009: 54).

Type locality: Tannum Sands, Gladstone, Queensland, Australia.

Geolocation: (map estimate fide Australian Museum holotype W.4013 catalogue record).

WoRMS: 328461

Synonyms: No subjective synonyms.

Status: The type species of the genus (by monotypy).

Etymology: Not stated, but the name for is evidently derived from the country of collection, Norway (Latin Norvegia), from which the feminine-suffix adjective ‘’ is derived.

Evaluation: Gender-variable adjective based on a geographic name. was given a species-group name with a feminine ending. Many usages as ‘’ exist (e.g., Moen 2006: 115).

Type locality: Trøndelag region, Norway. Trondheimsfjord off Statsbygd is one of three locations mentioned by Gunnerus (see Moen, 2006: 118).

Geolocation: Imprecisely known (map estimate for off Statsbygd).

WoRMS: 131009

Synonyms: There is an extensive list by McIntosh (1923: 347) of early serpulid names and usages in , , , and that are suggested to be synonyms. Nine of the placements were repeated later in a world catalogue (Hartman 1959), but only two can be confirmed here (see Read and Fauchald 2016 for status of the remainder). Also Mörch, 1863 named a subspecies , based on a Fabricius MS, but it is a nomen dubium unlikely to be a .

Claparède, 1870b (Gulf of Naples, Italy, Tyrrhenian Sea)

Bean, 1844 (Scarborough, North Yorkshire, England)

Etymology: Not stated, but the name for is evidently a Latinized form of the former name of the island of collection, New Britain, Bismarck Archipelago, now part of Papua New Guinea, once a German colony named Neupommern, after the Baltic (Ostsee) coastal lands besides Pommersche Bucht.

Evaluation: Invariant noun in the genitive case created from a non-Latin geographic name Latinized as ‘novaepommeran’.

Type locality: “Hanam-Hafen” (Hannan or Garua Harbour), north coast of New Britain, Papua New Guinea.

Geolocation: (map estimate).

WoRMS: 131010

Synonyms: Pillai, 1965 (Binakayan, Cavite, Manila Bay, Philippines)

Etymology: Rioja (1941a: 167) stated the name for was “dedicar esta especie al Maestro D. Isaac Ochoterena”, but he used the spelling ‘’ for the species-group name.

Evaluation: Incorrect Latinization to be treated as a noun in apposition. The use of has been regarded as an accidental incorrect original spelling by Bastida-Zavala and ten Hove (2003), who cited Article 32.5 (ICZN 1999) as justification for using ‘ochoterena’, although that would be an unchanged noun in apposition, rather than a genitive. Instead, we cannot reject the likelihood that Rioja had intentionally used the altered ‘-eana’ ending (after all he used it consistently five times but correctly spelled the name of dedicatee Ochotorena) aiming to create an adjectival form of Ochotorena. His adaptation could be intended as a rendering using the suffix ‘-anus’ -ana’ (belonging to), frequently used for adjectival Latinization of nouns based on personal and geographic names. As it was the author who was responsible for an incorrect Latinization (ICZN 1999, Article 32.5.1) his original spelling is not corrected (also see Welter-Schultes 2013: 77). This also avoids the name looking like an authorship (ICZN 1999, recommendation 31A).

Type locality: La Aguada and La Quebrada beaches, Acapulco, Mexico.

Geolocation: (La Quebrada, map estimate).

WoRMS: 328462

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for derives from the Latin verb operculo -avi -atum ‘to cover’, and in New Latin is used as an adjectival form. It is unclear why Treadwell chose the name as all have opercula. His specimen was endowed with two, but he didn’t name it ‘bioperculata’.

Evaluation: Gender-variable adjective with incorrect original feminine ending for . Usages in as ‘operculatus’ exist (e.g., Bellan 2001: 226).

Type locality: Berbera, Somaliland, Gulf of Aden.

Geolocation: (map estimate).

WoRMS: 131011

Synonyms: Straughan, 1967a [re-evaluating, see listing herein]

Straughan, 1967a [re-evaluating, see listing herein]

Pillai, 1960 [re-evaluating, see listing herein]

Etymology: The authors state that is named “for its distribution, as far as known yet restricted to the Pacific side of Panama (and adjacent areas).”

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin geographic name.

Type locality: Paitilla Beach (Punta Paitilla), Panama City, Western Panama.

Geolocation: (map estimate).

WoRMS: 328464

Synonyms: No subjective synonyms.

Etymology: Not stated, but Treadwell (1902: 210) stated the specimens of were “very small” (6 mm) thus Latin ‘small’.

Evaluation: Gender-variable adjective recombined in with correct feminine ending (e.g., Zibrowius 1971: 712, 717). Usages in as ‘’ exist (e.g., Hartman 1956: 250).

Type locality: West coast of Puerto Rico, Caribbean Sea, at both Boqueron Bay and nearby Mayagüez Harbour (station 6062, estimated ), as Treadwell had specimens from both locations. No station geolocations appear to have been available for the various Fish Hawk ‘Porto Rico’ stations (Treadwell 1939). Syntypes (USNM 16173) in the Smithsonian National Museum of Natural History are recorded as from Boqueron Bay (Bahia de Boqueron).

Geolocation: (map estimate, Bahia de Boqueron).

WoRMS: 876557

Synonyms: No subjective synonyms.

Status: Name now disused and representing a species inquirenda. It is not eligible as a candidate nomen oblitum (used in taxonomy by Iroso 1921: 49, Naples), but is indeterminable unless original specimens are found. The operculum figured by Philippi is similar to that of (Haswell, 1883) (fide Zibrowius 1971: 718).

Etymology: Philippi’s brief Latin description of describes the operculum spines as ‘utrinque pectinatis’ (pectinate both sides) with three sharp teeth. The Latin adjective indicates comb-like divisions.

Evaluation: Gender-variable adjective recombined in with correct feminine ending (e.g., Mörch 1863: 377).

Type locality: Unspecified Mediterranean, but can be narrowed to the Tyrrhenian Sea coast of Italy as Philippi’s activities were in western Italy, and plausibly to Naples as he was based there prior to 1844.

Geolocation: Unknown (Tyrrhenian Sea, with Naples shore ( map estimate) a possible point location).

WoRMS: 393822

Synonyms: As species inquirenda has no synonyms although has been suggested.

Etymology: Fauvel announces on the first page of his article that is dedicated to “M. Ch. Pérez”, who collected the worms off the Arabian coast.

Evaluation: Invariant genitive form from personal name Pérez.

Type locality: Pearling banks (within 24°55'N–25°10'N, 54°40'E–55°10'E) dredged ~15 miles from the coast of Oman (currently near Dubai, UAE) (Fauvel 1918: 329).

Geolocation: (map estimate, mid point of bounds given by author).

WoRMS: 209947

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after its La Plata collection station of the Swedish frigate Eugenie expedition.

Evaluation: Invariant genitive from Old Frankish ‘platen’, ultimately from Greek πλατuσ (platus) ‘flat’, relating to the Spanish La Plata placename, which plausibly had derived from a once widespread use of ‘plate’ to signify precious metals. Other ‘’ species group names of the period may relate to the German zoological collector Carl Platen (1843–1899) but clearly not this one.

Type locality: Offshore off the La Plata (“prope ostium fluvii La Plata”) embayment, Argentina/Uruguay (the Eugenie berthed at Montevideo, Uruguay).

Geolocation: Imprecisely known (map estimate for mid La Plata, offshore of Montevideo).

WoRMS: 369242

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named from (serpulid genus) combined with -cola ‘dweller’, because it was fastened on the tube of Benedict, 1887.

Evaluation: Invariant compound noun in apposition with -cola as a substantival suffix.

Type locality: Northeast off Cape Hatteras, North Carolina, Atlantic coast USA, 86 m.

Geolocation: (from author as ).

WoRMS: 338020

Synonyms: No subjective synonyms.

Etymology: The author states he named after the superficial similarity of the operculum to that of .

Evaluation: Gender variable adjective, with usage as corrected feminine ‘’ in Sun et al. (2015: 65).

Type locality: Shoreline on “island off north east Heywood Island” Kimberley, Western Australia. The author’s given geolocation () is oceanic and clearly incorrect. This is not a rounding error. The island north east of Heywood is the closely adjacent and much larger Jungulu Island.

Geolocation: (map estimate, Jungulu shore adjacent Heywood Island).

WoRMS: 882697

Synonyms: No subjective synonyms.

Status: Currently valid but it is possibly the same as the disused (see above).

Etymology: Not stated, but evidently was named because it represents one of the taxa previously confounded under (see below), a name regarded as of indeterminable identity from its original description (fide Zibrowius 1971: 709).

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘pseudouncinatus’ species (or nominal subspecies) exist (e.g., ten Hove and Kupriyanova 2009: 54).

Type locality: East off Île Gaby (also Degaby), Marseille, France, Mediterranean Sea (not in Zibrowius 1968, fide Zibrowius 1971: 708).

Geolocation: (map estimate).

WoRMS: 131012

Synonyms: No subjective synonyms.

Etymology: The author named subspecies after its continent of collection, Africa.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘africanus’ exist (e.g., ten Hove and Kupriyanova 2009: 54).

Type locality: Off Rio de Oro, Mauritania, Atlantic coast of Africa.

Geolocation: (author, ).

WoRMS: 335489

Synonyms: No subjective synonyms.

Etymology: The authors dedicated to Jian-Wen Qiu.

Evaluation: Invariant genitive form from personal name Qiu.

Type locality: East Arm Port, Darwin Harbour, Northern Territory, Australia.

Geolocation: (authors, ).

WoRMS: 852783

Synonyms: No subjective synonyms.

Etymology: Not stated, but is named after Ralum plantation, near its place of collection.

Evaluation: Gender-variable adjective, based on a non-Latin place-name, corrected by Day (1967: 806) from the masculine. The suffix ‘-anus -a’ is frequently used for Latinization of names based on localities and personal names.

Type locality: Ralum, Kokopo, Blanche Bay, New Britain (Neu-Pommern), Bismarck Archipelago of Papua New Guinea. The plantation “Ralum” was briefly the base for Friedrich Dahl, who collected the worms in 1896–97 (fide Augener 1927).

Geolocation: (map estimate).

WoRMS: 209951

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is perhaps a reference to the 8th enlarged dorsal verticil spine with its “pointed process perpendicular to it” from Latin rectus ‘perpendicular’.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as masculine ‘rectus’ exist (e.g., Pillai 2009: 132).

Type locality: Pretty Beach, north of Cairns, Queensland, Australia.

Geolocation: (map estimate, a beach 40 km north of Cairns).

WoRMS: 328466

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is likely referring to the verticil spines which are sharply curving backwards on themselves. Thus the name is formed from Latin recurvus ‘backward curved’ combined with spina ‘thorn’.

Evaluation: Invariant noun in apposition. Bastida-Zavala and ten Hove (2003: 99) maintained the original spelling.

Type locality: La Aguada, Acapulco, Mexico.

Geolocation: (map estimate).

WoRMS: 328467

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for may be referring to the shape of the verticil spines, derived from a combination of Greek ρομβος (rombos) ‘rhombus’, which is a parallelogram with only opposite angles equal, and Latin -ulus, which is a diminutive in masculine-form.

Evaluation: Clearly intended as an adjectival name, so it is corrected herein to feminine .

Type locality: Dongshan, Fujian Province, China

Geolocation: (map estimate).

WoRMS: 882579

Synonyms: No subjective synonyms.

Status: Previously unreplaced junior homonym preoccupied by the invalid Iroso, 1921, which was a n. nom. for the specimen of non Philippi, sensu Ehlers, 1887, but a junior objective synonym of (Bush, 1910) as Bush had already re-named it. Replaced by nom. n. here.

Etymology: Not stated, but is likely named after the large rostrum-like verticil spine figured by the author. The adjective rostratus -a -um, means having a beak. The genitive replacement name is in memory of Telesphore Gottfried Pillai (1930–2013), the original-name author.

Evaluation: Gender-variable adjective with correct original feminine ending. Usages as ‘rostratus’ exist (e.g., ten Hove and Kupriyanova 2009: 54).

Type locality: Hikkaduwa, Sri Lanka. Types were collected at both Hikkaduwa and Wellawatte. These localities are separated by some considerable distance, but the holotype at the Natural History Museum, London BM 1968–148, is from Hikkaduwa.

Geolocation: (map estimate).

WoRMS: 328469

Synonyms: No subjective synonyms.

Etymology: The authors named as a dedication to Sergio Salazar-Vallejo.

Evaluation: Invariant genitive form from personal name Salazar-Vallejo.

Type locality: Cabo de la Aguja, Santa Marta region, Colombia, Caribbean Sea.

Geolocation: (map estimate).

WoRMS: 328470

Synonyms: No subjective synonyms.

Etymology: Not stated, but is clearly named after its type locality, Saint Croix Island, and the syntypes at the Zoological Museum, University of Copenhagen are labelled “Kr. St. Croix, legit Oerstedt”. The genitive of the feminine Latin noun crux ‘cross’ is .

Evaluation: Place-name translated into Latin. The genitive-case noun is invariant.

Type locality: Saint Croix (unspecified further), Virgin Islands, Caribbean Sea.

Geolocation: (map estimate for the port area).

WoRMS: 333645

Synonyms: Augener, 1922 [partim, fide Bastida-Zavala and ten Hove 2002: 147] (Haiti, Caribbean Sea)

Etymology: Not stated, but an instance of the Latin adjective ‘similar to’. Later in the same work Treadwell (1929: 12) considered his (now ) as similar to his , which isn’t compared to any taxon, so the more logical application of the names would have been in reverse.

Evaluation: Masculine/feminine invariant adjective ( -e) (Stearn 1983: 94).

Type locality: Unspecified beyond a “Lower California” location on label (Baja California, Mexico). The collector was Townsend, on the ‘Albatross’ voyage of 1911, and the location is perhaps more likely the Gulf of California than off the Pacific coast. Gulf coast sites mentioned by Treadwell where other polychaetes were collected include PageBreakIsla Carmen and Isla San José, but there are many other possibilities (see Townsend 1916: 399, end map).

Geolocation: Unknown ( (gazetteer) as Baja California general region, but perhaps inner coast).

WoRMS: 369244

Synonyms: No subjective synonyms.

Etymology: The authors state they named for its resemblance to (type locality Baja California) thus combining the Latin adjective ‘similar to’ with the suffix -oides, also ‘similar to’.

Evaluation: Invariant adjectival suffix -oides.

Type locality: La Parguera (jetty of marine institute), Isla Magueyes, Puerto Rico.

Geolocation: (map estimate).

WoRMS: 328471

Synonyms: No subjective synonyms.

Etymology: The author states the name “refers to the simple unmodified spines at the base of the enlarged coronal [verticil] spine”, combining Latin adjectives simplus -a -um ‘simple’ and dentatus -a -um ‘toothed’.

Evaluation: Corrected to the feminine form in Sun et al. (2015: 79) as clearly an adjectival name.

Type locality: Unnamed reef north-west of Buffon Island (but cf. author’s supplied geolocation which is non-reef and east of Buffon Island), Kimberley, Western Australia.

Geolocation: (author as stated, but likely displaced incorrectly by ~13 km to the East).

WoRMS: 882648

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named for its occurrence on the coast of China.

Evaluation: Masculine/feminine invariant Latin adjective (‘’) referring to China, a non-Latin geographic name.

Type locality: Off Qingdao (Zibrowius as ‘Tsindao’), China coast, northern Yellow Sea.

Geolocation: (map estimate).

WoRMS: 328472

Synonyms: No subjective synonyms.

Etymology: Not stated but is evidently named from English ‘sponge’ as stem spongi- combined with -cola ‘dweller’, because of its association as “frail calcareous tubes in living sponges”.

Evaluation: Invariant compound noun in apposition with -cola as a substantival suffix.

Type locality: West offshore from Venice, Florida, Gulf of Mexico, USA, 48 m.

Geolocation: (as from author as ).

WoRMS: 338021

Synonyms: No subjective synonyms.

Etymology: The species is dedicated to Heinz Steinitz.

Evaluation: Invariant genitive form from personal name Steinitz.

Type locality: Sinai bank of Little Bitter Lake, Suez Canal, opposite Al-Kabrit on Egyptian bank.

Geolocation: (opposite Al-Kabrit, map estimate).

WoRMS: 131014

Synonyms: No subjective synonyms.

Etymology: Not stated, but is from Greek Στοιχαδας (Stoichadas), an old name for Îles d’Hyères, an archipelago of small islands near Toulon, Mediterranean coast of France (H. Zibrowius pers. comm.).

Evaluation: Invariant Latinization created from Greek place-name, having the form of a noun in apposition.

Type locality: Cap du Merlan, the south west corner of Parc Nacional de Port Cros (island), off the Mediterranean coast of France.

Geolocation: (map estimate).

WoRMS: 131015

Synonyms: No subjective synonyms.

Etymology: Not stated but is evidently named after its place of collection, Tambalagam.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin place-name.

Type locality: Nachchikuda, Tambalagam Lake (a bay), eastern Sri Lanka.

Geolocation: (map estimate).

WoRMS: 328474

Synonyms: [noun in apposition] Pillai, 2009 (Long Reef, Kimberley, Western Australia)

Etymology: The authors dedicated the name to Harry ten Hove.

Evaluation: Invariant genitive form from personal name ten Hove.

Type locality: Cabo San Lazaro, western coast of Baja California Sur, Mexico.

Geolocation: (authors , adjusted to be coastal).

WoRMS: 328475

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for evidently refers to three vesicular verticil spines (three lobes), which are small ones, hence the diminutive Latin suffix -ulus.

Evaluation: Clearly meant as an adjectival name, so it is corrected herein to .

Type locality: Xisha Islands (Paracel Islands, unspecified further), South China Sea, of which Yongxing Island is the largest.

Geolocation: Imprecisely known ( if Yongxing Island, map estimate).

WoRMS: 882593

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for is evidently referring to the three lobes of the enlarged dorsal spine of the verticil, thus tri with Latin adjective vesiculosus -a -um ‘full of blisters’.

Evaluation: Gender-variable adjective with ending feminine as in Sun et al. (2015: 85) and Kupriyanova et al. (2015; 293), both mistakenly reporting the original name as ‘’). Usages as ‘’ (besides original) exist (e.g., ten Hove and Ben Eliahu 2005: 134).

Type locality: Heron Island, Queensland coast, Australia, collected close to the marine station by Dew (map in Straughan 1967b).

Geolocation: (map estimate).

WoRMS: 882647

Synonyms: No subjective synonyms.

Etymology: The authors named dedicated to Jossy S. Tromp, a student of ten Hove.

Evaluation: Invariant genitive form from personal name Tromp.

Type locality: Lower chamber wall and floor, Miraflores Locks, Panama Canal, Panama.

Geolocation: 8.9967, -79.5964 (authors).

WoRMS: 328478

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name for may refer to tubercles (knobs) on each verticil spine as the Latin noun tuberculum refers to a swelling or lump.

Evaluation: Gender-variable adjectival form of tuberculum with correct original feminine ending. Usages as ‘tuberculatus’ exist (e.g., Bailey-Brock 1987: 282).

Type locality: Urata (beach), Tanegashima (island), Southern Japan. Imajima (1976a) also gives records for Sumiyoshi, and off Nishinoomote Harbour, Tanegashima, but a Urata specimen is the holotype (NSMT-Pol. H-120) at the National Museum of Nature & Science, Tokyo.

Geolocation: (map estimate, Urata).

WoRMS: 871950

Synonyms: No subjective synonyms.

Status: Name now disused and representing a species inquirenda which is the type species of . It is not a candidate nomen oblitum (used in taxonomy as valid by Zibrowius 1968: 109, Hartman 1969: 757, Gibbs 1971: 202, Pillai 1972: 15, Day 1973: 132, and others), but the taxon it represents may be indeterminable unless original specimens are found. Zibrowius (1968) named to establish a separation from , and Pillai (1972) did likewise with . The name does not obviously threaten the validity of subsequent names, but it is notable that several usages have been assigned elsewhere, e.g., non Philippi, sensu Gravier, 1906 to .

Etymology: Not stated, but Philippi described the verticil spines as with “cornubus octo, apice incurvo uncinatis” (eight horns, curved tip hooked), thus the name refers to the hooked spines, from the Latin adjective ‘hooked’.

Evaluation: Gender-variable adjective with correct feminine ending. Usages in as ‘’ exist (e.g., Ehlers 1913: 582).

Type locality: Unspecified Mediterranean, but can be narrowed to the Tyrrhenian Sea coast of Italy as Philippi’s activities were in western Italy, and plausibly to Naples as he was based there prior to 1844.

Geolocation: Unknown (Tyrrhenian Sea, with Naples shore ( map estimate) a possible point location).

WoRMS: 156135

Synonyms: As species inquirenda has no synonyms.

Etymology: Not stated, but the name evidently refers to the straight, thick, unornamented spines of the verticil which are described as “uniform”.

Evaluation: Masculine/feminine invariant adjective ( -e) (Stearn 1983: 94).

Type locality: Kesao, Guadalcanal, Solomon Islands, Pacific Ocean.

Geolocation: (map estimate).

WoRMS: 369245

Synonyms: No subjective synonyms.

Etymology: Not stated, but the name likely derives from the collection location, Visakhapatnam, which has the nickname Vizag.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin place-name.

Type locality: Naval Base (collected off settlement panels), Visakhapatnam Harbour, east coast of India, Bay of Bengal.

Geolocation: (map estimate).

WoRMS: 870503

Synonyms: No subjective synonyms.

Etymology: Not stated, but is evidently named after its area of collection, the Xisha Islands.

Evaluation: Masculine/feminine invariant ‘-ensis’ adjective created from a non-Latin geographic name.

Type locality: Xisha Islands (Paracel Islands, unspecified further), South China Sea, of which Yongxing Island is the largest.

Geolocation: Imprecisely known ( if from Yongxing Island, map estimate).

WoRMS: 328480

Synonyms: No subjective synonyms.

Discussion

Name characteristics and potential variation

Place-names (23), and personal names (16) make up more than a third (36%) of the 107 non-synonymised species-group names in , with most of the remainder (68) being descriptive of species character states, with a remarkable number relating to operculum morphology (54). The (perhaps) ‘small-eared’ was the only species where the reason for the author’s choice was not obvious. Otherwise three species were named for their attractive appearance (, , ), three were named for the animals they were found on PageBreak(, , ), four names are comparative (, , , ), two species names probably relate to body size (, ), and one species name relates to the tube form ().

Currently the 107 names include 41 which should be gender invariant (including 17 nouns in apposition, including two acronyms), and 23 with adjectival masculine/feminine endings in -is, which would only change (to -e) if moved to a neuter genus (13 of these are place-names). The remaining 43 names are fully gender variable. There are 68 adjectival names in total (including 19 adjectival place-names), with only two adjectival names completely invariant.

Type locality distribution

Type localities of the serpulids listed are, with one exception, in shallow-water coastal locations in temperate to tropical waters between 43.3°N and 35.3°S (Figs 1–2). is the exception from deeper water (but still inshore), and occurred at the highest latitude at 63.4°N. It is the most cold tolerant based on type locality, with a 20° latitudinal gap to all other species type localities, although its distribution extends south into the Mediterranean (Zibrowius 1971). The western Pacific Ocean (Australia PageBreakto northern Japan) has the biggest group of new species at 39, with another 15 species in the northern Indian Ocean and Red Sea (Fig. 1), a total of 54 for the greater Indo-Pacific. The Americas north of the equator have 13 new species on the East Pacific coast, and 18 on the Western Atlantic coast and the Caribbean/Gulf of Mexico area (Fig. 2), a total of 31 for North American and Caribbean coasts combined. Europe (including Azores) has only 10 new species, mostly in the Mediterranean Sea (Fig. 1). This leaves only 12 other species described from elsewhere. Notably, few new species (9) have been reported from the major continental coasts of the South American coast south of the equator, and the African coast (outside of the Red Sea and Mediterranean, and including Madagascar), but this may be partly a reflection of lesser sampling effort, and also the consequence of other areas being examined first, given that species are readily translocated on the floating objects and vessel hulls they colonise.

Figure 1.

species type localities of the Eastern Hemisphere (Indian Ocean and Western Pacific). Labels are current species-group names (except homonym renamed herein as nom. n.)

Figure 2.

species type localities of the Western Hemisphere (Americas and Eastern Pacific). Labels are current species-group names. Map grids 30° intervals.