Trophoniella avicularia Caullery, 1944

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

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Trophoniella avicularia Caullery, 1944


Trophoniella avicularia Caullery, 1944

( Fig. 1 View FIG )

Trophoniella avicularia Caullery, 1944: 35 , 36, fig. 27. — Bleeker & Van der Spoel 1992: 163.

Pherusa eruca indica – Al-Hakim & Glasby 2004: 38 (non Fauvel 1928).

TYPE MATERIAL. — Indonesia. Holotype of Trophoniella avicularia (ZMA 1598) , Irian Jaya, R/V Siboga expedition, stn 162, between Loslos and Broken Island, Salawatti, W coast, 18 m, dredge, sand, shells, 18.VIII.1899.

ADDITIONAL MATERIAL. — Indonesia. 1 specimen ( ZMB 123) damaged, in 2 pieces, stn NB20B (03°34.92’N, 107°58.54’E), off Natuna Barat, 105 m, 3.VIII.2001, I. Al-Hakim (32 (19 + 13) mm long, 2.5 mm wide, cephalic cage 2.5 mm long, 79 (38 + 41) chaetigers).

DISTRIBUTION. — Only known from two localities in Indonesia, from shallow water to 105 m depth.


Holotype (ZMA 1598) complete. Tunic thick, heavily covered by sand, shell fragments, forams and other large particles dorsally and ventrally ( Fig. 1A, B View FIG ); posterior end grayish, without sediment cover ( Fig. 1C View FIG ). Body cylindrical, tapering posteriorly, 29 mm long, 3.5 mm wide, cephalic cage 5 mm long, 49 chaetigers. Body papillae long, capitate, sparse, arranged in longitudinal rows, five dorsally, three ventrally; difficult to follow because of the size of foreign particles.

Anterior end not observed; holotype not dissected. Cephalic cage damaged, chaetae as long as 1⁄₆ body length or 1.3 times longer than body width. Chaetigers 1-3 involved in the cephalic cage; chaetae of chaetiger 1 arranged in short dorsal and ventral series; others in short ventrolateral series. Chaetiger 1 with two chaetae per bundle, one neurochaeta thicker; chaetiger 2 with two chaetae per bundle, neurochaetae of about the same thickness; chaetiger 3 with three chaetae per bundle.

Anterior dorsal margin of first chaetiger papillated, without projection. Anterior chaetigers without especially long papillae. Chaetigers 1-3 enlarging posteriorly. Chaetal transition from cephalic cage to body chaetae gradual; anchylosed, bidentate neurohooks starting by chaetiger 18. Gonopodial lobes not seen.

Parapodia poorly developed ( Fig. 1D, E View FIG ); chaetae emerging from the body wall. Parapodia lateral; median neuropodia ventrolateral. Noto- and neuropodia short lobes with 4-6 papillae per bundle. Noto- and neuropodia distant to each other.

Median notochaetae arranged in short longitudinal series, about 6-8 (3-5 remains) per bundle, as long as 1⁄₃ body width; all notochaetae multiarticulated capillaries, articles short throughout the chaetae ( Fig. 1D View FIG , insert), slightly longer near the tip, tips falcate, most broken. Neurochaetae multiarticulated neurohooks in chaetigers 1-17; each straight, with short articles, becoming slightly longer near the falcate tip; by chaetiger 15, smaller hooks together with multiarticulated falcate capillaries. By chaetiger 20, all neurohooks anchylosed, curved distally, darker, medially widened, subdistally expanded, with short rings continued to the subdistal hyaline region, tip bidentate, accessory tooth wide, not passing fang ( Fig. 1E View FIG , insert); neurohooks arranged in transverse series, 4-5 per bundle.

Posterior end ( Fig. 1C View FIG ) mostly without sediment (last nine chaetigers at least), tapering to a blunt cone; pygidium with anus terminal; no anal cirri.


Caullery (1944) overlooked the first chaetiger when he described T. avicularia , because of its very small size and because chaetae were broken.As stated above, he thought his species was clearly associated with T. eruca Claparède, 1868 because both have bidentate neurohooks. He overlooked Fauvel (1928: 7), who had provided a short description and illustrations for his S. eruca indica Fauvel, 1928 , noticing that his new variety was closely allied to Stylarioides eruca but differing in having longer, thinner neurochaetae. This implies that Fauvel noticed that parapodial lobes are pointed and directed forwards, as it is the case in the supposedly parental form. However, their neurochaetae are markedly different because they have anchylosed, short articles restricted to the base or up to the median chaetal region, and hooks can be distally expanded, hyaline, whereas in Piromis the neurohooks are multiarticulated, having longer articles, rarely distally expanded, and rings are visible up to the subdistal chaetal region.

Trophoniella avicularia is closely allied to T. indica n. comb., n. stat. (see below), but they differ in the relative development of the notopodial lobes in anterior chaetigers, and in the distal chaetal article. In T. avicularia , the notopodial lobes are reduced, not forming any prominent lobe, whereas in T. indica n. comb., n. stat. they are well delineated and pointing forward. Further, the bidentate neurohooks have different proportions in teeth size; in T. avicularia the accessory tooth is short, never passing the fang, whereas in T. indica n. comb., n. stat. they are larger, often passing the fang. The other specimen (ZMB 123) is regarded as belonging to the same species with hesitation, because the neurohooks are straight and subdistally swollen, but because it is damaged, its recognition as a distinct, separate species should be postponed.

Trophoniella americana ( Monro, 1928) n. comb. ( Fig. 2 View FIG )

Stylarioides capensis americana Monro, 1928: 96 , 97, fig. 16; 1933: 1057, 1058, textfig. 6 partim.

Piromis americana – Hartman 1961: 123. — Fauchald 1972: 415 partim.

Semiodera glabra Treadwell, 1928: 479 , 480, fig. 179: 62-65.

TYPE MATERIAL. — Eastern tropical Pacific. Lectotype of Stylarioides capensis americana ( BMNH 1933.7.10.144), by present designation, cut in 2 pieces, anterior fragment ventrally dissected, labeled “Balboa, Galapagos”, C. Crossland; 4 paralectotypes of S. c. americana ( BMNH 1933.7.10.138/9), damaged, 1 broken in 2 pieces, off Balboa, Panama, 1923/4, C. Crossland (19.0- 24.5 mm long, 1.5-2.5 mm wide, cephalic cage 3-4 mm long, 44-56 chaetigers, first neurohooks from chaetiger 5).

Holotype and 2 paratypes of Semiodera glabra ( AMNH 3520 View Materials ), 1 complete paratype, off Isla Española (01°22’59”S, 89°39’00”W), Galápagos Islands, 5 m, 25.IV.1925 (holotype 24 mm long, 4.0 mm wide, cephalic cage 4.5 mm long, 49 chaetigers; larger paratype 7 mm long, 1.2 mm wide, cephalic cage 2.0 mm long, 40 chaetigers) GoogleMaps . ADDITIONAL MATERIAL. — Eastern tropical Pacific, Gulf of California. 1 specimen ( LACM-AHF 2484), E of La Paz, Baja California Sur, 21.III.1940, E. Ricketts (37 mm long, 4 mm wide, cephalic cage 5 mm long, 54 chaetigers). — 1 specimen ( ECOSUR), among mussel beds, Bahía de Los Angeles, Baja California, 2 m depth, 27.III.1985, E. & C. Aguirre (16 mm long, 3.5 mm wide, cephalic cage 4 mm long, 55 chaetigers). — 1 specimen ( ECOSUR), Playa El Caimancito, La Paz, Baja California Sur, mixed bottom (rocks and sand), 1 m depth, 8.X.1987, J. A. de León & S. I. Salazar-Vallejo (33 mm long, 5 mm wide, cephalic cage 5.5 mm long, 60 chaetigers).

Western Mexico. 1 specimen ( ECOSUR), Islas Marietas, Nayarit, mixed bottom (rocks and sand), 1 m depth, 27.XI.2004, P. Salazar & B. Yáñez (6 mm long, 2 mm wide, cephalic cage 2.5 mm long, 48 chaetigers). — 1 specimen ( ECOSUR), Sayulita, Nayarit, mixed bottom, intertidal, 28.XI.2004, P. Salazar & B. Yáñez (54 mm long, 4 mm wide, cephalic cage 5 mm long, 91 chaetigers). — 1 specimen ( ECOSUR), Playa Manzanilla, Nayarit, mixed bottom (rocks and sand), 1 m depth, 29.XI.2004, P. Salazar & B. Yáñez (19 mm long, 3 mm wide, cephalic cage 4.5 mm long, 59 chaetigers). — 1 specimen ( ECOSUR), broken in 2, Muelle Balleto, Islas Marías, Nayarit, mixed bottom (rocks and sand), 1 m depth, 2.XI.1979, S. I. Salazar-Vallejo (13 mm long, 2 mm wide, cephalic cage 3 mm long, 55 chaetigers).

Panama. 6 specimens ( ECOSUR), juveniles, Club de Yates , 1.VI.2002, L. Harris & S. I. Salazar-Vallejo (4-9 mm long, 1.0- 1.5 mm wide, cephalic cage 1.5-3.0 mm long, 32-49 chaetigers; first neurohooks in chaetiger 5; very few sediment particles). — 2 specimens ( USNM 66368 View Materials ), partially dried-out, Paitilla Beach, 28.X.1970, A. A. Reimer .

Ecuador. 2 specimens ( LACM-AHF 2485 ), a mature ♀ and a small juvenile with sediment grains dorsally, Manta Bay , R / V Velero III, stn 400 (00°56’30”S, 80°44’18”W), shore, rock and sand, 19.I.1935 (♀, incomplete posteriorly, 55 mm long, 4.5 mm wide, cephalic cage 5 mm long, 69 chaetigers; oocytes 100 µm in diameter; juvenile 9 mm long, 1.5 mm wide, cephalic cage 1.5 mm, 39 chaetigers; dorsal and chaetal lobes papillae larger than in adults). — 1 specimen ( USNM 1132100 View Materials ) apparently fixed in alcohol, Salinas, no further data, 12.IX.1926 (40 mm long, 5.5 mm wide, cephalic cage 5 mm long, 60 chaetigers; regenerating the last 20 chaetigers) GoogleMaps .

LECTOTYPE LOCALITY. — The lectotype has a label indicating “Balboa, Galápagos ”. This label is confusing because “Balboa” is on the Panamian Pacific coast whereas “ Galápagos ” indicates the archipelago, but because the species distribution includes both localities, no restriction is needed nor easy to discern.

DISTRIBUTION. — Gulf of California to Ecuador and the Galápagos Islands, in shallow subtidal depths. The records in California ( Hartman 1969: 305) are questionable.


Lectotype of S. c. americana ( BMNH 1933.7.10.144), cut in 2 pieces ( Fig. 2A View FIG ), anterior fragment ventrally dissected. Body cylindrical, posteriorly tapered, body wall pinkish or pale, 77 mm long, 5.5 mm wide, cephalic cage 9 mm long, 93 chaetigers.Tunic thick, papillated, with sediment grains dorsally, extending laterally; ventral papillae tiny, abundant, larger papillae in longitudinal rows, three dorsally, four ventrally.

Cephalic hood exposed in holotype, short, margin smooth ( Fig.2B View FIG ). Prostomium low cone, eyes faded; caruncle well developed, separating the branchial plate in two halves. Palps lost, palp keels low, round- ed. Branchiae cirriform, sessile on a tongue-like protuberance; arranged in three rows per side, with many filaments each; branchial plate distally fused in juveniles. Lectotype ( BMNH 1933.7.10.144) with distally notched branchial plate ( Fig. 2C View FIG ); most filaments lost; size relationship with palps unknown.

Cephalic cage chaetae as long as 1/9 body length, about twice as long as body width (larger or smaller specimens). Chaetigers 1-2 involved in the cephalic cage; chaetae arranged in short series, each about the body corners in the smaller syntype, dorsolateral in the larger one, 2-3 noto- and 4-5 neurochaetae.

Anterior dorsal margin of first chaetiger with a papillated margin, partially damaged; chaetigers 1-4 with long notopodial papillae. Chaetigers 1-3 of about the same length. Chaetal transition from cephalic cage to body chaetae abrupt. Chaetiger 4 with transitional neurohooks; falcate anchylosed neurohooks from chaetiger 5. Gonopodial lobes in chaetiger 5, small, yellowish (lacking in the smaller syntype, no paralectotype).

Parapodia better developed in chaetigers 1-4, posterior ones less developed, placed over the body corners.Median neuropodia ventrolateral.Notopodia as a short conical lobe, one short and one long postchaetal papillae, becoming longer in posterior chaetigers. Neuropodia flat transversal lobe, with two longer papillae, one superior, and one postchaetal ( Fig. 2D View FIG ).

Median notochaetae arranged in longitudinal short series, 4-5 per bundle, as long as 1⁄₃ body width; all multiarticulated capillaries, articles short basally, medium-sized to long medially, long distally ( Fig. 2E View FIG ). Smaller paralectotype with neurochaetae multiarticulated capillaries in chaetiger 1, chaetigers 2-4 with transition hooks, each with falcate tips, becoming shorter, distally widened. Anchylosed neurohooks entire, from chaetiger 5, arranged in transverse series, 3-4 anteriorly, 5-6 medially ( Fig. 2F View FIG ), 2-4 posteriorly, thinner, each subdistally straight, with a subdistal oblique dark spot, short articles feebly developed, continued to the median region.

Posterior end short, conical, with long papillae; pygidium with anus terminal, no anal cirri.


Trophoniella americana n. comb. is a senior synonym of T. glabra ( Treadwell, 1928) . The former was published on 28 July 1928, as indicated in the reprint’s last page, whereas the second was published on 5 December 1928 (Steve Johnson pers. comm.). The printed number code was not easily identified as the publication date, until several volumes of the same journal became available online; the number code was then confirmed as the publication date. Before that the priority was not settled and some specimens identified as T. glabra should be relabeled as T. americana n. comb.

Monro (1928) described Stylarioides capensis americana using several specimens collected in both Pacific Panama and Galápagos, whereas Treadwell (1928) described Semiodera glabra from Isla Española, Galápagos Islands and Ecuador. However, this latter name was not used again, mainly due to a confusion about the type locality that Treadwell had indicated; thus, he stated as the type locality the coordinates for one island in Galápagos ( Treadwell 1928: 480), but in a later paper, he stated the type material was coming from Haiti ( Treadwell 1929: 9; Hartman 1956: 294), and that it was identical with Stylarioides dubius Treadwell, 1929 . The confusion stems from the similarity between the two Spanish names: Isla Española, also known as Hood Island in Galápagos, and Hispaniola, the island now separated into Haiti and the Dominican Republic.

On the other hand, Monro (1928: 97) proposed Stylarioides capensis americana because he did not find any “trace in the dorsal chaetae of the small terminal hook described by both McIntosh and Gravier”. He later redefined the new variety, after finding the falcate notochaetae, and then he diagnosed it ( Monro 1933: 1058) by “the absence of the bold, transverse, distal striae [= articulation] on the ventral hooks […] described and figured by McIntosh” and by having neurospines from chaetigers 6-7. Further, Monro (1933: 1057, 1058) restricted the definition of his new variety by using two basic features: the lack of striae (articles) in neurospines, which defines Piromis capensis ( McIntosh, 1885) , which is in turn a junior synonym of P. arenosus Kinberg, 1867 , and the start of neurospines by chaetigers 6-7. His syntypes, albeit damaged, have anchylosed neurospines (starting in chaetiger 5), such that they cannot fit Piromis but rather belong into Trophoniella , as proposed here.

Treadwell (1928: 479) indicated that “the type which is more than twice as large as any other in the collection is 30 mm long”. All the specimens were together in the same vial, but this does not invalidate regarding one of them as the holotype ( ICZN 1999: art. 73.1.1). Further, there are other materials which could have been regarded as paratypes; they were not examined because Treadwell stated that they were even smaller. Concerning the suspected synonymy with Stylarioides dubius , as stated by Treadwell, it has to be rejected; S. dubius is provided with a dorsal shield, has a very thin tunic and short papillae in transverse rows. Hence, it belongs in Semiodera , as will be shown in a forthcoming contribution. Hartman (1956: 294, 295) noticed the change in the type locality mentioned above, indicating that it could be the same as S. cariboum , and that it could also include S. c. americana . In fact, Monro included different forms under the same name and some former syntypes have been subsequently separated as members of other species (see below), whereas his name has been restricted to those forms with sediment particles over the dorsal and lateral surfaces, and having anchylosed neurohooks from early chaetigers (5-7).

Trophoniella americana n. comb. is closely allied to T. tumbensis n. comb. because their tunics have large sediment particles over the dorsal surface and often extending laterally. They differ in two main features; T. americana n. comb. has fewer notochaetae (3-5) than T. tumbensis n. comb. (8-10), and its body wall is pale or pinkisk whereas it is grayish in T. tumbensis n. comb.

The syntypes include some specimens that belong to T. grandis n. comb., and therefore, in order to clarify the delineation for the species, one lectotype has been selected from the type series ( ICZN 1999: art. 74).


Museum für Naturkunde Berlin (Zoological Collections)


El Colegio de la Frontera Sur (Mexico)














Trophoniella avicularia Caullery, 1944

SALAzAR-Vallejo, Sergio I. 2012

Piromis americana

FAUCHALD K. 1972: 415
HARTMAN O. 1961: 123

Trophoniella avicularia

CAULLERY M. 1944: 35

Stylarioides capensis americana

MONRO C. C. A. 1928: 96

Semiodera glabra

TREADWELL A. L. 1928: 479