Trophoniella Caullery, 1944: 35 Fauchald 1977: 117 Trophoniella Hartman, 1959: 423 Therochaetella Hartman, 1967: 128 Therochaetella chilensis Hartman, 1967 Therochaetella Hartman, 1967 Therochaetella chilensis Hartman, 1967 Trophoniella Bradabyssa Hartman, 1967 Trophoniella Therochaetella Trophoniella Piromis Trophoniella Trophoniella Piromis Piromis Trophoniella Piromis Trophoniella Trophoniella T. americana ( Monro, 1928 ) T. ehlersi ( Day, 1973 ) T. grandis T. havaica ( Kinberg, 1867 ) T. incerta ( Augener, 1918 ) T. jareckiorum T. lindae T. minuta T. reishi T . rigida ( Caullery, 1944 ) T. rudis T. salazarae T. tumbensis (Hartmann-Schröder, 1962) T. avicularia Caullery, 1944 T. bastidai T. borealis T. capitata ( Nonato, 1966 ) T. chilensis ( Hartman, 1967 ) T. eliasi T. enigmatica T. fauveli T. fernandensis ( Augener, 1918 ) T. fiegei T. harrisae T. hospita T. indica ( Fauvel, 1928 ) T. orensanzi Revision of Trophoniella Hartman, 1959 (Polychaeta, Flabelligeridae) SALAzAR-Vallejo, Sergio I. Zoosystema 2012 2012-09-30 34 3 453 519 84VGN Hartman, 1959 Hartman 1959 [265,572,754,781] Polychaeta Flabelligeridae Trophoniella Animalia Terebellida 5 Annelida genus      Trophoniella Caullery, 1944: 35(unavailable). —  Fauchald 1977: 117.     Trophoniella Hartman, 1959: 423.     Therochaetella Hartman, 1967: 128. — Typespecies:  Therochaetella chilensis Hartman, 1967, by original designation.   TYPESPECIES. —  Trophoniella avicularia Caullery, 1944, by original designation ( Hartman 1959: 423).  DISTRIBUTION. — Species of the genus are found in tropical or subtropical regions, especially in sediments or mixed bottoms in shallow water (up to 200 m), but two species,  T. borealis( Hartman, 1965) n. comb.and  T. hospita( Fauchald, 1972) n. comb., were found in deeper water ( 500-600 m).  DIAGNOSIS. — Cephalic cage present.Tunic thick, mostly with sediment grains. Papillae arranged in longitudinal series, fragile, clavate, sometimes inconspicuous. Median chaetigers with multiarticulated or anchylosed neurochaetae, simple falcate or bidentate; posterior chaetigers with anchylosed neurochaetae, straight, tapering or subdistally expanded, tips entire, bidentate or bifid, never mucronate. Branchiae often abundant, short, cirriform filaments, on a tongue-shaped lobe; rarely, few filaments on a flat branchial plate.  REMARKS The chronic confusion of the generic definitions deserves to be explained. Ehlers (1887: 158-161)identified some specimens as  Siphonostomum cariboumGrube & Ørsted inGrube, 1859; he provided an extensive description and excellent illustrations for his specimens, which were followed by Saint-Joseph (1898: 367). Following Ehlers, Chamberlin (1919: 397)rightly proposed  Semiodera, with  Siphonostomum cariboumas its type species. However, the specimen that Ehlers illustrated differs from the holotypeof  S. cariboum(ZMUC 331), because this species has a well-developed dorsal shield over few anterior chaetigers, a very thin tunic, its body papillae are tiny without sediment cover, and its branchiae are few, arranged in 1-2 rows. Ehlers’ specimen had few or no large sediment particles, but no dorsal shield, a very thick tunic, body papillae are long, especially in notochaetal lobes, the branchiae are abundant and arranged on a tongue-shaped plate, and neurochaetae are anchylosed. Consequently, his specimen belongs to an undescribed species of  Trophoniella, which is described below as  T. lindae n. sp. The next confusion deals with  Trophoniella Caullery, 1944. Caullery proposed this name (“n. nom. gen.”, 1944: 35) to replace Trophonia Malmgren, 1867, that was introduced to replace  PherusaOken, 1815, used before for other taxa. The typespecies for  Pherusais  Amphitrite plumosaMüller, 1776, which had been proposed by Oken (1807)and confirmed by StØp-Bowitz (1948: 13). It must be emphasised that Caullery was following Malmgren’s generic concept, which was based upon the corresponding concept by: “M. Edw. 1829 sec. Agassiz.” But the original quote was: “Aud. et Edw. Litt. de la Fr. 1829 ( Agassiz 1848: 1096).” It has been shown that this name was introduced elsewhere and is a nomen nudum(Salazar-Vallejo 2011b: 166). Further, Caullery indicated that his genus was not following Milne-Edwards ( Caullery 1944: 35): “(= Trophonia Malmgren 1867, Levinsen 1883, LoBianco 1893, necAudouin et Edwards 1829)”. This is surprising because Malmgren (1867: 191)followed Milne-Edwards (1829), after Agassiz (1848); thus, Malmgren included two species in his proposed grouping,  Pherusa plumosa(Müller, 1776)and  P. glauca Malmgren, 1867, now called  Diplocirrus glaucus( Malmgren, 1867).  Trophoniellawas apparently proposed for a junior homonym: Trophonia Malmgren, 1867vs TrophoniaMilne- Edwards, 1836 ( ICZN 1999: art. 60.3), and in so doing, they would become objective synonyms ( ICZN 1999: art. 61.3.3 and art. 72.7). However, these generic concepts were addressed to different species or species groups. On the other hand, after ICZN (1999: art. 67.8), the typespecies for the original name and for the replacement name is the same, which would imply that  Trophoniella Caullery, 1944becomes a junior synonym of  Pherusa Oken, 1807, which would result in a worse situation because the species content is very different. Consequently, Trophonia Malmgren, 1867is regarded as a nomen nudumand  Trophoniellais regarded as a new name. Further,  Trophoniellaincluded three species after the original proposal, but the typespecies was not formally established and this would render the name unavailable ( ICZN 1999: art. 13.3). However, this has been solved by a subsequent designation of the typespecies ( ICZN 1999: art. 69.1). Hartman (1959: 423)recognised the genus as distinct and designated  T. avicularia Caullery, 1944as the typespecies; this was followed by Fauchald (1977: 117). This proposal made the name available and therefore it must be assigned to Hartman ( ICZN 1999: art. 50.1).  Caullery (1944)indicated that: “ T. aviculariarappelle  T. erucaClap.et  T. arenosaWebster[…] évidemment, il s’agit d’une espèce du même groupe, déjà distinguée par de Saint-Joseph.” This would be translated as: “  T. aviculariaresembles  T. erucaClap.and  T. arenosaWebster[…] certainly, it is a species of the same group, already recognised as such by Saint-Joseph.” Therefore,  Trophoniellacould be regarded as a close relative of  Piromis, but there are some important differences in the neurochaetae between them and hence  Trophoniellaincludes species having anchylosed neurochaetae with entire, bidentate or bifid tips. The two other species included in the genus,  T. intoshi Caullery, 1944and  T. sibogae Caullery, 1944do not belong to the genus because they have aristate neurochaetae; they should be transferred to  BradaStimpson, 1854. On the contrary,  Brada rigida Caullery, 1944that is provided with anchylosed neurochaetae should be transferred to  Trophoniella.Likewise several other species currently included in different genera but having anchylosed neurohooks and branchial filaments in a tongue-shaped branchial plate are herein regarded as belonging in  Trophoniellaas well.  Fauchald (1977: 117)regarded  Semioderaas a junior synonym of  Piromis, perhaps by following Ehlers’ description and illustrations, and stated that there were 10 species in the genus, all provided with a tongue-shaped branchial plate that could be entire or bifid, and uni- or bidentate neurohooks. However, Day (1973: 108)had redefined  Piromisto include only those species provided with bidentate neurohooks, and he regarded those species with entire neurohooks as members of  Pherusa.  Piromis arenosus Kinberg, 1867, the typespecies for the genus, has bidentate neurohooks with long distal articles ( Hartman 1949: 117, pl. 15, fig. 7), and a single tongue-like branchial plate ( Day 1967: 664, fig. 32.4c). These features are diagnostic for species belonging to that genus as confirmed elsewhere (Salazar-Vallejo 2011a). Therefore, the species formerly included in  Piromisbecause they have a tongue-like branchial plate, but are provided with neurohooks having very short, anchylosed articles in median or posterior chaetigers, should be transferred to  Trophoniella, which must be redefined accordingly.    Therochaetella Hartman, 1967(p. 128), which was proposed for  Therochaetella chilensis Hartman, 1967(p. 128), is a subjective junior synonym of  Trophoniella.The paratypes(USNM 55551) include an undescribed species of  Bradabyssa Hartman, 1967, which prompted the inclusion of an “elongated oral tube” as a diagnostic feature ( Hartman 1967: 120, key), whereas the illustration letterings were transposed. The holotype, however, is a member of  Trophoniella, with a low tongue-shaped branchial lobe. The presence of bidentate neurochaetae is stressed in the generic diagnosis for  Therochaetellaand in the type-species description, but the only illustration (pl. 37, fig. D, from chaetiger 25) depicts an unidentate hook. Further, the lettering of the figure was transposed; thus, what is indicated to be C should be D, and vice versa. Consequently, the species is herein restricted, redefined and transferred to  Trophoniella. Therefore,  Piromisand  Trophoniellashare several features; both have tunics with variable amounts of sediment particles, especially over the dorsal surface, and longitudinal rows of elongated single papillae along the body.However,  Trophoniellamust be separated from  Piromison the basis of having differently constructed neurohooks, especially in posterior chaetigers; thus, whereas in  Piromisthey have at least one distal long article, that articulates with the previous one, in  Trophoniella, as in some other flabelligerid genera, neurohooks have short or very short anchylosed articles, mostly fused to the next. This modification implies that some species must be transferred from elsewhere, including  Piromis, to  Trophoniella. Within  Trophoniella, besides the differences on the relative cover of sediment grains on the body wall, there are two different patterns on the start of anchylosed neurohooks. They might start before chaetiger 10, or they can start by chaetiger 15, but usually beyond it. The species having anchylosed or falcate neurohooks from anterior chaetigers are  T. americana( Monro, 1928) n. comb.from Galápagos and Panama,  T. ehlersi( Day, 1973) n. comb.from the eastern United States,  T. grandis(Blanchard in Gay, 1849) n. comb. from central Chile,  T. havaica( Kinberg, 1867) n. comb.from Hawaii,  T. incerta( Augener, 1918) n. comb.from West Africa,  T. jareckiorum n. sp.from the Caribbean Sea,  T. lindae n. sp.from the Caribbean Sea,  T. minuta(Blanchard in Gay, 1849) n. comb. from Chile,  T. reishi n. sp.from the Gulf of California,  T. rigida( Caullery, 1944) n. comb.from Indonesia,  T. rudis(Grube & Müller in Grube, 1877) n. comb. from southern Brazil,  T. salazarae n. sp.from western Mexicoand  T. tumbensis(Hartmann-Schröder, 1962) n. comb.from Chile. All other species, including the typespecies, have anchylosed neurohooks beginning on more posterior chaetigers. These are  T. avicularia Caullery, 1944from Indonesia,  T. bastidai n. sp.from western Mexico,  T. borealis n. comb.from northeastern South America,  T. capitata( Nonato, 1966) n. comb.from Brazil,  T. chilensis( Hartman, 1967) n. comb.from Chile,  T. eliasi n. sp.from central Argentina,  T. enigmatica n. sp.from the Mediterranean Sea,  T. fauveli n. sp.from the Mediterranean Egyptian coast,  T. fernandensis( Augener, 1918)reinstated, n. comb. from northwestern Africa,  T. fiegei n. sp.from the Persian Gulf,  T. harrisae n. sp.from southern California,  T. hospita n. comb.from the Gulf of California,  T. indica( Fauvel, 1928) n. comb., n. stat.from the Bay of Bengal and  T. orensanzi n. sp.from Uruguay.