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Acromegalomma, nomen novum, is introduced as a replacement name for the polychaete genus Megalomma Johansson, 1926 (Annelida, Sabellidae), preoccupied by Megalomma Westwood, 1842 (Insecta, Coleoptera, Carabidae). The historical background of the homonymy and a full list with 36 new combinations in the new genus are included, while two species are considered as species inquirenda.
The genus Megalomma (Annelida, Sabellidae) was established by Johansson (1926) for the usage of Branchiomma sensu Claparède (1869), based on the species Branchiomma koellikeri Claparède, 1869. However, the name Megalomma is preoccupied by Megalomma Westwood, 1842 (Insecta, Coleoptera), a well-established genus of tiger beetles from the Mascarene Islands. Megalomma Johansson, 1926 has no known available or potentially valid synonyms, for which reason, and in accordance with Article 60.3 of the ICZN (1999), it must be replaced by a new substitute name.
The name Megalomma was first used as a subgenus of Cicindela Linnaeus, 1758 (Insecta, Coleoptera, Carabidae) by Westwood (1842: 203), for the Mauritian species Cicindela (Megalomma) vigilans Westwood, 1842, and raised to the generic level the following year by Lacordaire (1843: 113). The genus is well established and in current use (see Moravec 2007), comprising five species from the Mascarene Islands (West Indian Ocean): M. fulgens W. Horn, 1892; M. janaki Moravec, 2007; M. oculatum (Fabricius, 1799); M. pierreorum Deuve, 2000; and M. viridulum (Quensel in Schönherr, 1806), which includes as a synonym the type species of the genus, M. vigilans (Westwood, 1842).
A second genus Megalomma was created by Smith (1873: 405) in Hymenoptera (Insecta), to include three new species from Brazil: Megalomma politum Smith, 1873, M. elegans Smith, 1873, and M. nigriceps Smith, 1873. Later Schulz (1906: 200) pointed that it was a junior homonym of Megalomma Westwood, 1842 and replaced it by the new name Megistommum Schulz, 1906.
In polychaetes, the name Megalomma was first used by Johansson (1926: 10), as a replacement name for Branchiomma sensu Claparède (1869), based on a misinterpretation of Claparède’s work. While discussing the validity of the name Dasychone introduced by Sars (1862) for sabellids with eyes on their radioles, Claparède (1869) stated that Kölliker (1858) had already used the name Branchiomma for the same group, to include Amphitrite bombyx Dalyell, 1853 (renamed as Branchiomma dalyellii Kölliker, 1858). However, instead of synonymizing the junior Dasychone into Branchiomma, Claparède (1869: 162) tried to retain both by redefining the genus Branchiomma, with the following justification:
Quoi qu’il en soit, le nom de Dasychone a pris place dans la science, et celui de Branchiomma est à peu près oublié. Je pense pourtant pouvoir ressusciter celui-ci, en tenant compte des scrupules de M. Sars, et sans proscrire le nom de Dasychone. Dans son mémoire sur le genre Branchiomma, M. Kölliker décrit en outre de la Dasychone Bombyx une autre espèce qu’il n’a étudiée que d’une manière très-cursive, il est vrai, dans le golfe de Naples, et qui est caractérisée par des yeux à l’extrémité des branchies. Cette espèce que j’ai retrouvée n’est point une Dasychone. Elle pourra rester dorénavant l’espèce-type du genre Branchiomma.
Hence, Claparède (1869) attempted to transfer Amphitrite bombyx Dalyell, 1853 to Dasychone (see Claparède 1869: 168), while making reference to a short comment by Kölliker (1858: 536) where he recorded that he had observed, but not named nor described in detail, an additional sabellid from Naples with eight compound eyes near the tips of the radioles:
Schon im Jahre 1842 kam mir in Neapel ein kleiner Kopfkiemer unter die Augen, der an seinen Kiemen 8 zusammengesetzte Sehorgane trug. Leider war es mir damals, da ich gerade mit der Verfolgung der Entwicklung der Cephalopoden beschäftigt war, nicht möglich, diese interessante Annelide, von der ich ohnehin nur Ein Individuum erhalten hatte, näher zu verfolgen, und unterliess ich es daher, etwas über dieselbe zu veröffentlichen.
Consequently, Claparède described this species as Branchiomma koellikeri [original spelling köllikeri corrected here to koellikeri according to Article 184.108.40.206 of the ICZN (1999)], based on the specimens collected by him at the Gulf of Naples, citing (Claparède 1869: 164):
Je doute à peine que cette espèce soit la même que M. Kölliker a eue sous les yeux. Elle n’est en effet point rare dans le golfe de Naples. M. Kölliker n’indique, il est vrai, que huit filaments branchiaux, tandis que j’en ai compté jusqu’à trente-deux. Mais cela peut ne tenir qu’à une différence d’âge.
This leaves little doubt that Claparède considered his new species Branchiomma koellikeri to be the same species observed previously by Kölliker from Naples and, moreover, that he proposed B. koellikeri as the type of his emendation of Kölliker’s genus with the sentence (Claparède 1869: 162): “Elle pourra rester dorénavant l’espèce-type du genre Branchiomma.”
Apparently Claparède’s intention was simply to redefine the genus Branchiomma to restrict it to the unnamed Kölliker species (= B. koellikeri Claparède, 1869, the intended new type species of the redefined genus), as can be inferred by the fact that he wrote “Genre Branchiomma Koellkr. char. em.” (Claparède 1869: 162).
However, Johansson (1926: 10) considered erroneously that Claparède was formally establishing a new genus, an interpretation that was followed by other authors (e.g. Hartman 1959, Day 1967, Fauchald 1977).
In this way, Johansson (1926: 10) argued that the generic name Branchiomma sensu Claparède (1869), used for Branchiomma koellikeri, could not be accepted, as it was already preoccupied by Kölliker (1858) for the species Amphitrite bombyx Dalyell, 1853. A new replacement name was thus necessary, and Johansson introduced for the third time in Zoology the name Megalomma, emphasizing the large compound eyes of the genus (Johansson 1926: 10):
Als Claparède 1869 für seine Art Köllikeri die Gattung Branchiomma bildete, war der Name schon präokkupiert. Die Gattung Branchiomma Claparède muss also einen neuen Namen erhalten. Ich schlage Megalomma vor, welcher Name wie Branchiomma auf die grossen zusammengesetzen Augen dieser Gattung hindedeutet, doch ohne ihren Charakter als Branchialaugen hervorzuheben [...].
However, and as stated above, the name Megalomma Johansson, 1926 is itself preoccupied by Megalomma Westwood, 1842, and a replacement name is necessary. The name Acromegalomma, nomen novum is here proposed to accomplish this need.
1842. Westwood (p. 203): Cincidella (Megalomma) new subgenus (Insecta, Coleoptera, Carabidae), for Cicindella (Megalomma) vigilans Westwood, 1842.
1843. Lacordaire (p. 113): Megalomma raised to generic level.
1858. Kölliker (p. 537): Branchiomma new genus (Annelida, Sabellidae), for Amphitrite bombyx Dalyell, 1853 (renamed as Branchiomma dalyellii Kölliker, 1858).
1869. Claparède (p. 162–163): Branchiomma redefined (Annelida, Sabellidae), for Branchiomma koellikeri Claparède, 1869. Amphitrite bombyx Dalyell, 1853 assigned to Dasychone Sars, 1862.
1873. Smith (p. 405): Megalomma new genus (Insecta, Hymenoptera, Crabronidae), for Megalomma politum Smith, 1873, M. elegans Smith, 1873, and M. nigriceps Smith, 1873.
1906. Schulz (p. 200): Megistommum new name (Insecta, Hymenoptera, Crabronidae), replacement name for Megalomma Smith, 1873.
1926. Johansson (p. 10): Megalomma new genus (Annelida, Sabellidae), to include Branchiomma sensu Claparède, 1869 (not Branchiomma Kölliker, 1858).
Present study. Acromegalomma new name (Annelida, Sabellidae), replacement name for Megalomma Johansson, 1926.
To establish the list of new combinations in Acromegalomma new name a list of valid Megalomma species was compiled based on WoRMS (Bellan 2008), and updated with Tovar-Hernández and Carrera-Parra (2011), Mikac et al. (2013), Capa and Murray (2015) and Giangrande et al. (2015). Synonymies were obtained from Tovar-Hernández and Carrera-Parra (2011). The type of synonymy and its author are provided inside square brackets, following the synonym.
Each new combination is accompanied by the reference of the original description, synonymies, type locality of the species and remarks, when necessary. Type locality is based on the original description, except where indicated. Geolocations of type localities are derived from the original descriptions, being considered an “original geolocation” when the authors provided the coordinates, or “estimated geolocation”, when estimated using Google Earth (www.google.com/earth) from the general geographic limits described by the authors. All geolocations were converted to decimal degrees.
The whereabouts of type material of the new combinations were summarised by Tovar-Hernández and Carrera-Parra (2011) and following publications describing new species (Mikac et al. 2013, Capa and Murray 2015, Giangrande et al. 2015).
While the gender of Megalomma and Acromegalomma new name is neuter, some names in Megalomma had incorrect endings and needed to be emended. Following Article 31.2 of the ICZN (1999), the names of the new combinations and the species inquirenda were herein revised to assure they agreed in gender with the generic name with which they are combined. Original names with incorrect endings are indicated with the remark “[sic]” following the specific epithet. Endings corrected herein are: carunculatum for carunculata, inflatum for inflata, interruptum for interrupta, jubatum for jubata, and longoventrale for longoventralis.
, nomen novum
The type species of the new genus is Branchiomma koellikeri Claparède, 1869 (junior synonym of Sabella lanigera Grube, 1846), according to recommendation 60A of the ICZN (1999). Type by monotypy, established by Johansson (1926).
Following his principle of basing observations and descriptions only on live organisms Édouard Claparède did not designate type material or deposit specimens in museums or collections (Fauchald 1989). However, Knight-Jones (1997) refers the existence of a type of Branchiomma koellikeri Claparède, 1869 deposited at the Zoological Museum of Berlin (currently the Museum für Naturkunde Berlin), with the reference number ZMB 6387. Tovar-Hernández and Carrera-Parra (2011) refer this specimen as being a syntype. Although the designation of a lectotype for B. koellikeri is desirable, it is out of the scope of the present work.
The name of the new genus is composed by combining the Greek terms for acro, meaning “tip end” or “extremity of a body”, mega, meaning “big” or “large”, and the suffix –omma, a noun meaning “eye”, and referring to the big compound eyes located on the radiolar subdistal region, typical of the genus.
The publication date of the genus Megalomma Johansson should be considered as “1926”. It was generally accepted as being “1927” until Tovar-Hernández and Salazar-Vallejo (2008) pointed out that the name had been introduced in a previous publication by the same author, referring the date as “1925”. In fact, the last page of this publication states “Tryckt den 5 november 1925” (“Printed the 5 November 1925”) and in the following line “Uppsala 1925. Almquist & Wiksells Boktryckeri-A.-B.” This date is also present in existing reprints of the paper. However, the bounded volume comprising the article provides the publication date as “Häfte 2 inneh. A N:o 6-12, [...] utkom den 5 juni 1926” (“Booklet 2 cont. A No. 6-12, [...] published 5 June 1926”). This booklet includes article 7A by Johansson, where the name Megalomma is introduced for polychaetes. Hence, the work was printed in 5 November 1925, but published only in the following year, on 5 June 1926.
The genus Acromegalomma, nomen novum is represented by 36 valid species, all of them new combinations.
(Grube, 1878) comb. n.
Singapore (1.25°, 103.85°; estimated geolocation) or Philippines (12.22°, 121.77°; estimated geolocation).
(Giangrande, Caruso, Mikac & Licciano, 2015) comb. nov.
Brindisi, Italy, South Adriatic Sea (40.65°, 17.95°; original geolocation).
(Ehlers, 1887) comb. n.
West of Dry Tortugas, Straits of Florida (24.6181°, -83.0517°; original geolocation).
(Tovar-Hernández & Salazar-Vallejo, 2008) comb. n.
Punta Manzanillo, Acapulco, Guerrero, Mexican Pacific (16.842°, -99.910°; estimated geolocation).
(Fitzhugh, 2003) comb. n.
Hungtou Yu (Orchid Island), northern coastline, about 1 km east of Langtao Village, Taiwan, Pacific Ocean (22.0794°, 121.5369°; original geolocation).
(Moore, 1923) comb. n.
Between S. 35° W, 3.5 miles (5.6 km) and S. 43° W, 5.2 miles (8.4 km) off Brockway Point, Santa Rosa Island, Channel Islands, California, Pacific coast of the USA (34.02°, -120.22°; estimated geolocation).
(Gravier, 1906) comb. n.
Syntypes collected at the reef of Marabout (11.611°, 43.132°; estimated geolocation), at Djibouti Bay, and the “Grand Récif” (11.736°, 43.235°; estimated geolocation), Moucha Islands, both at the Gulf of Tadjoura, Gulf of Aden, Indian Ocean.
Gravier introduced the name Branchiomma claparedei as new twice, first in 1906 (Gravier 1906: 39) and again in 1908 (Gravier 1908b: 91). This caused some confusion, inducing some authors in error, by considering the correct publication date as being 1908, while overlooking the smaller 1906 publication (e.g. Tovar-Hernández and Carrera-Parra 2011). The correct publication date is therefore “1906” (see also Wehe and Fiege 2002).
(Chamberlin, 1919) comb. n.
Laguna Beach, California, Pacific coast of the USA (33.542°, -117.786°; estimated geolocation).
(Giangrande, Licciano & Gambi, 2007) comb. n.
Lagoon side of Carrie Bow Cay, Belize, Caribbean Sea (16.803°, -88.085°; estimated geolocation).
(Tovar-Hernández & Carrera-Parra, 2011) comb. n.
Off Georgia, Atlantic coast of the USA (30.95°, -79.9667°; original geolocation).
(Knight-Jones, 1997) comb. n.
La Herradura, Estero Jaltepeque, El Salvador, Pacific Ocean (13.303°, -88.902°; estimated geolocation).
(Perkins, 1984) comb. n.
Hutchinson Island, Florida, Atlantic Ocean (27.345°, -80.2133°; original geolocation).
(Capa & Murray, 2009) comb. n.
Southeast of Bate Bay, New South Wales, Australia (-34.0667°, 151.2167°; original geolocation).
(Capa & Murray, 2009) comb. n.
One Tree Island, Queensland, Australia (-23.5°, 152.0833°; original geolocation).
(Capa & Murray, 2015) comb. n.
MacGillivray Reef, Lizard Island, Queensland, Australia (-14.6569°, 145.4947°; original geolocation).
(Knight-Jones, 1997) comb. n.
Point Kean near Kaikoura, east coast of South Island, New Zealand (-42.425°, 173.715°; estimated geolocation).
(Grube, 1846) comb. n.
Branchiomma köllikeri Claparède 1869: 163–164, plate XXII fig. 4 [subjective synonymy by Knight-Jones (1997)]. Branchiomma vesiculosum neapolitana Claparède 1869: 164–166, plate XXII fig. 5 [subjective synonymy by Giangrande and Licciano (2008)].
The species was described based on a single specimen (T-ZMB 136) deposited at the Zoological Museum of Berlin (currently the Museum für Naturkunde Berlin), from an unknown location (Grube 1846: 51).
(Ehlers, 1887) comb. n.
Key West, Florida, Gulf of Mexico (24.54°, -81.80°; estimated geolocation).
(Giangrande, Caruso, Mikac & Licciano, 2015) comb. n.
Rovinj, coastal station near the Island Banjole, Croatia, North Adriatic Sea (45.095250°, 13.619283°; original geolocation).
(Giangrande & Licciano, 2008) comb. n.
Brindisi, Italy, Adriatic Sea (40.65°, 17.96°; estimated geolocation).
(Nishi, 1998) comb. n.
Ao Tang Khen, Phuket, Thailand, Andaman Sea (7.8185°, 98.4144°; estimated geolocation).
(Quatrefages, 1866) comb. n.
Lima, Peru, Pacific Ocean (-12.07°, -77.15°; estimated geolocation).
(Fitzhugh, 2002) comb. n.
Thailand, Andaman Sea (08.5°, 98.1°; original geolocation).
(Gravier, 1906) comb. n.
“Grand Récif” (11.736°, 43.235°; estimated geolocation), Moucha Islands, Gulf of Tadjoura, Gulf of Aden, Indian Ocean.
As in the case of Branchiomma claparedei explained above, Gravier introduced the name Branchiomma mushaensis [sic] as new twice, first in 1906 (Gravier 1906: 40) and again in 1908 (Gravier 1908b: 94). This incurred some authors into error (e.g. Tovar-Hernández and Carrera-Parra 2011). The correct publication date is “1906” (see also Wehe and Fiege 2002).
(Knight-Jones, 1997) comb. n.
El Bilaiyim (= Ghor Blayim) lagoons, Sinai Peninsula, Gulf of Suez (28.55°, 33.24°; estimated geolocation).
(Grube & Örsted in Grube, 1859) comb. n.
Punta Arenas, Gulf of Nicoya, Costa Rica (9.976°, -84.852°; estimated geolocation).
(Tovar-Hernández & Salazar-Vallejo, 2006) comb. n.
Cape Lookout, North Carolina, Atlantic coast of the USA (34.62°, -76.54°; estimated geolocation).
(Capa & Murray, 2009) comb. n.
Off Townsend Point, Corner Inlet, Victoria, Australia (-38.8°, 146.55°; original geolocation).
(Reish, 1963) comb. n.
Bahía de San Quintín, Baja California, Mexico, Pacific Ocean (30.456°, -115.958°; estimated geolocation).
(Mikac, Giangrande & Licciano, 2013) comb. n.
13 nautical miles off the coast of the Istrian Peninsula, Croatia, Gulf of Venice, Northern Adriatic Sea (45.2833°, 13.2667°; original geolocation).
(Willey, 1905) comb. n.
Aripu (= Arippu) Coral Reef, Sri Lanka, Gulf of Manaar, Indian Ocean (08.78°, 79.87°; estimated geolocation).
(Gravier, 1908) comb. nov.
Payta (= Paita), Peru, Pacific Ocean (-5.083°, -81.111°, estimated geolocation).
(Moore, 1905) comb. n.
Kasaan Bay, center of Round Island, S. 10d W., 0.4 miles, Clarence Strait, Prince of Wales Island, Alexander Archipelago, SE Alaska, North Pacific Ocean (55.51°, -132.39°; estimated geolocation).
Pseudopotamilla anoculata Moore 1905: 566–568, plate XXXVII figs 28–33 [subjective synonymy by Hartman (1959)]. Branchiomma disparoculatum Treadwell 1914: 223–224, plate 12 figs 44–46 [subjective synonymy by Hartman (1956)]. Branchiomma burrardum Berkeley 1930: 71, fig. 1 [subjective synonymy by Knight-Jones (1997)].
(Ehlers, 1904) comb. n.
French Pass, between D’Urville Island and north end of South Island, New Zealand (-40.922°, 173.837°; estimated geolocation).
(Reish, 1968) comb. n.
Lagoon side of Engebi (= Enjebi) Island, Enewetak (= Eniwetok) Atoll, Ralik Chain, Marshall Islands, Pacific Ocean (11.658°, 162.235°; estimated geolocation).
(Montagu, 1813) comb. n.
Original type locality at Kingsbridge Estuary, Devon, England (50.263°, -03.765°; estimated geolocation). Neotype designated by Knight-Jones (1997: 314) from St. Anthony, Cornwall, England (50.15°, -5.2667°; original geolocation misplaced, corrected here to 50.152°, -5.006°).
(Claparède, 1870) [unreplaced junior secondary homonym]
Gulf of Naples, Mediterranean Sea (40.7°, 14.3°; estimated geolocation).
Branchiomma vigilans was described on the basis of three specimens from the Gulf of Naples, all of them found with their muddy tubes inserted among the dorsal chaetae of individuals of Aphrodita aculeata Linnaeus, 1758 (Claparède 1870). Afterwards the species was recorded on only a couple of occasions in the Western Mediterranean, first by Marion (1876), in the Gulf of Marseille and from 60–65 m (no habitat details), and later by Soulier (1903), who observed about ten specimens collected off Séte (Gulf of Aigues-Mortes) among the chaetae of A. aculeata specimens. Rioja (1923) attributed an empty sandy tube found among the dorsal chaetae of an A. aculeata collected in the region of Valencia to this species, but this record is very dubious, as not only was the worm not present but also the nature of the tube differed from that described by Claparède (1870). Moreover, no type material of B. vigilans is known to exist (Knight-Jones 1997, Giangrande and Licciano 2008, Tovar-Hernández and Carrera-Parra 2011) as Claparède normally did not deposit specimens in museums or collections (Fauchald 1989).
The species was transferred to Megalomma by Hartman (1959: 550), creating a junior secondary homonym of the tiger beetle Megalomma vigilans (Westwood, 1842) (see above), and has since remained a poorly known but valid taxon (Knight-Jones 1997, Tovar-Hernández and Salazar-Vallejo 2008). Giangrande and Licciano (2008) considered the species as being quite rare, probably due to its peculiar habitat, and in spite of stating that its real status needed confirmation, they also observed that it was likely a valid species. However, the species was subsequently omitted from the discussions on new Mediterranean species of Megalomma by Mikac et al. (2013) and Giangrande et al. (2015). The most recent reference to the species seems to be by Tovar-Hernández and Carrera-Parra (2011: 5), who wrote:
Megalomma vigilans (Claparède, 1870) was originally found as an epibiont of the sea mouse Aphrodita aculeata Linnaeus, 1758, in the Mediterranean Sea, however, no new records of this association exist. [...] In the case of M. vigilans, the description is poor, the type is lost and there are no additional records.
The described habitat of Megalomma vigilans is unusual, and there are no references of similar cases in the family Sabellidae. It is possible that the habitat is an artefact resulting from the collection process, and that the presence of the species on individuals of Aphrodita aculeata was the consequence of the rough treatment and mixing suffered by the biological material collected by grabs and trawls, or even during the processing of the samples. So, the presence of M. vigilans on A. aculeata could be a post-collection phenomenon, and not the natural habitat of the worm. It is difficult or even impossible to know if the records by Marion (1876) and Soulier (1903) refer to the same species as that collected and described by Claparède (1870) without studying the material, if still existing. There is a possibility that M. vigilans is not as uncommon as it seems, but that for some reason it has not been collected or recognised. For the time being, M. vigilans is here considered as a species inquirenda.
Syntypes collected at Aranuka Island, outside the coral reef (0.14°, 173.56°; estimated geolocation), and Tapeteuea (= Tabiteuea) Island, inside the lagoon (-1.5°, 175.0°; estimated geolocation), both at Gilbert Islands, Kiribati, Pacific Ocean.
According to Fitzhugh (2002), Megalomma pacificum Johansson, 1927 probably belongs to the genus Demonax Kinberg, 1867 (a name recently replaced by Parasabella Bush, 1905 due to homonymy; see Tovar-Hernández and Harris 2010). The fact that the holotype has dried out (Knight-Jones 1997) and that the species has a remote type locality have likely prevented a formal redescription. The species was not included in the Parasabella species list given by Tovar-Hernández and Harris (2010), but its possible inclusion in Parasabella has been implicitly accepted by subsequent authors (Capa and Murray 2009, Tovar-Hernández and Carrera-Parra 2011).
The authors would like to express their deep gratitude to the generous help of Miguel Ángel Alonso Zarazaga (MNCN, Madrid, Spain), who replied to our questions concerning some nomenclatural details and corrected the endings of the specific epithets of the new combinations, so they agreed in gender with the new generic name, and Andrew S. Y. Mackie (National Museum Wales, Cardiff, UK) for having found the time to make a critical reading of the first draft of the manuscript and correct its English while on vacations. Geoff Read and María Ana Tovar-Hernández are thanked for their very detailed revisions of the first version, which have improved the manuscript considerably, and Chris Glasby for his help as Editor. This work was financially supported by the European Community's Seventh Framework Programme (FP7/2007-2013), through the Assemble Grant Agreement no. 227799-ASSEMBLE to the project “Biodiversity of AnnelidaPolychaeta in the Ria Formosa coastal lagoon: a baseline study”, awarded to J.G. and developed at the CCMAR, University of Algarve (Faro, Portugal).
Gil J, Nishi E (2017) Nomenclatural checklist for Acromegalomma species (Annelida, Sabellidae), a nomen novum replacement for the junior homonym Megalomma Johansson, 1926. ZooKeys 677: 131–150. https://doi.org/10.3897/zookeys.677.12030