Trophoniella Hartman, 1959

SALAzAR-Vallejo, Sergio I., 2012, Revision of Trophoniella Hartman, 1959 (Polychaeta, Flabelligeridae), Zoosystema 34 (3), pp. 453-519 : 453-519

Genus Trophoniella Hartman, 1959

Trophoniella Caullery, 1944: 35 (unavailable). — Fauchald 1977: 117.

Trophoniella Hartman, 1959: 423 .

Therochaetella Hartman, 1967: 128 . — Type species: Therochaetella chilensis Hartman, 1967 , by original designation.

TYPE SPECIES. — 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 cariboum Grube & Ørsted in Grube, 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 cariboum as its type species. However, the specimen that Ehlers illustrated differs from the holotype of 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 Pherusa Oken, 1815 , used before for other taxa. The type species for Pherusa is Amphitrite plumosa Müller, 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, nec Audouin 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) . Trophoniella was apparently proposed for a junior homonym: Trophonia Malmgren, 1867 vs Trophonia Milne- 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 type species for the original name and for the replacement name is the same, which would imply that Trophoniella Caullery, 1944 becomes a junior synonym of Pherusa Oken, 1807 , which would result in a worse situation because the species content is very different. Consequently, Trophonia Malmgren, 1867 is regarded as a nomen nudum and Trophoniella is regarded as a new name.

Further, Trophoniella included three species after the original proposal, but the type species 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 type species ( ICZN 1999: art. 69.1). Hartman (1959: 423) recognised the genus as distinct and designated T. avicularia Caullery, 1944 as the type species; 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. avicularia rappelle T. eruca Clap. et T. arenosa Webster […] évidemment, il s’agit d’une espèce du même groupe, déjà distinguée par de Saint-Joseph.” This would be translated as: “ T. avicularia resembles T. eruca Clap. and T. arenosa Webster […] certainly, it is a species of the same group, already recognised as such by Saint-Joseph.” Therefore, Trophoniella could be regarded as a close relative of Piromis , but there are some important differences in the neurochaetae between them and hence Trophoniella includes species having anchylosed neurochaetae with entire, bidentate or bifid tips. The two other species included in the genus, T. intoshi Caullery, 1944 and T. sibogae Caullery, 1944 do not belong to the genus because they have aristate neurochaetae; they should be transferred to Brada Stimpson, 1854 . On the contrary, Brada rigida Caullery, 1944 that 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 Trophoniella as well.

Fauchald (1977: 117) regarded Semiodera as 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 Piromis to 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 type species 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 Piromis because 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 Therochaetella and 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, Piromis and Trophoniella share 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, Trophoniella must be separated from Piromis on the basis of having differently constructed neurohooks, especially in posterior chaetigers; thus, whereas in Piromis they 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 Mexico and T. tumbensis (Hartmann-Schröder, 1962) n. comb. from Chile.

All other species, including the type species, have anchylosed neurohooks beginning on more posterior chaetigers. These are T. avicularia Caullery, 1944 from 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.