Trophoniella harrisae, SALAzAR-Vallejo, 2012

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

Trophoniella harrisae n. sp.

( Fig. 14 )

Piromis hospitis – Blake 2000: 12, 13, fig. 1.4 (non Fauchald 1972).

TYPE MATERIAL. — Northeastern Pacific Ocean , California. Holotype ( LACM-AHF 2486 ), off White Point, R / V Velero IV, stn not in log book, 93 m, 10.IV.1957. Paratypes: 3 anterior fragments and some posterior ones ( LACM-AHF 2487 ), 2 miles off Point Fermin Light, R / V Velero IV, stn 5502 (33°41’30”N, 118°19’50”W), 49 fathoms, fine, gray, silty sand, 16.XII.1957 (15-50 mm long, 3.5-8.0 mm wide, cephalic cage 6-7 mm long, 28-53 chaetigers, dorsal tubercles in chaetigers 2-end of fragment; first neurohooks from chaetigers 22-27). — 2 complete and another one almost complete specimens ( LACM-AHF 2488 ), 3.7 miles off San Clemente Pier, R / V Velero IV, stn 5632 (33°22’50”N, 117°39’00”W), 32 fathoms (tag: 213 feet), fine green sand, 22.II.1958 (completes 75-76 mm long, 6-7 mm wide, cephalic cage 5.5-6.0 mm long, 96-107 chaetigers, dorsal tubercles in chaetigers 2-90 [95]; first neurohooks from chaetigers 27- 28; the other 63 mm long, 7 mm wide, cephalic cage 7 mm long, 86 chaetigers, dorsal tubercles in chaetigers 2-85; first neurohooks from chaetiger 24). — 1 anterior fragment ( LACM-AHF 2489 ), 2.75 miles off Point Fermin Light, R / V Velero IV, stn 5664 (33°43’10”N, 118°20’40”W), 11 fathoms, medium gray to black sand, 21.III.1958 (58 mm long, 6 mm wide, cephalic cage 6.5 mm long, 64 chaetigers, dorsal tubercles in chaetigers 2-37 [tunic removed from dorsum of chaetiger 38]; first neurohooks from chaetiger 20). — 2 anterior fragments ( LACM-AHF 2490 ), 10.3 miles off Santa Monica Pier Light, R / V Velero IV, stn 5975 (34°00’28”N, 118°42’28”W), 29 fathoms, green mud, 22.XI.1958 (largest dissected; 51 mm long, 7 mm wide, cephalic cage 5 mm long, 43 chaetigers; dorsal lappets in chaetigers 2-43; first neurohooks from chaetiger 25) .

ADDITIONAL MATERIAL. — Northeastern Pacific Ocean, California. Anterior fragment ( CAS 133248), Moss Landing Monterey Bay Study, 36°52.3’N, 121°59.8’W, 63 m, 13.X.1971. — Anterior fragment ( CAS 133251), Moss Landing Monterey Bay Study, 36°50.8’N, 121°53.6’W, 62.5 m, 24.XI.1971.

TYPE LOCALITY. — Off White Point, California, in sediments at 93 m depth.

DISTRIBUTION. — Central to southern California sediments in 22-100 m depth.

ETYMOLOGY. — This species is gratefully being named after Leslie H. Harris, who has worked a lot on Californian polychaete fauna, have supported my research activities during many years, and especially because she noticed that there were some relevant differences between this and similar species, paving the road for this description.

DESCRIPTION

Holotype complete, brownish pink ( Fig. 14A ), broken into two pieces, most chaetae broken; body club-shaped, tapering posteriorly, 147 mm long, 10 mm wide, cephalic cage 9 mm long, 141 chaetigers.Tunic papillated, with fine sediment particles dorsally, almost no sediment particles ventrally; posterior region almost without sediment ( Fig. 14C ). Papillae arranged in longitudinal rows, two dorsal series of larger papillae, four ventral ones with smaller papillae. Dorsal papillae forming large, mammiliform, paired dorsal tubercles, decreasing in size posteriorly, present to chaetiger 40.

Anterior end dissected from one paratype (LACM- AHF 2490). Cephalic hood not exposed, smooth. Prostomium low, eyes blackish, large. Caruncle well developed, running through the tongue-like branchial plate, but not reaching the distal margin, lateral ridges black. Palps black, deeply corrugated; palp keels rounded, slightly pigmented. Lips black; lateral and ventral lips fused, retracted; dorsal lip projected as a small rounded lobe. Branchiae cirriform, pale, sessile on a short, tongue-like protuberance, separated in two lateral groups, each with filaments arranged in about 12 concentric rows, over 100 filaments per group ( Fig. 14D ). Branchiae slightly shorter than palps. Nephridial lobes double, dehiscent, stemming from the basal inner corner of branchial groups.

Cephalic cage chaetae about 1⁄₁₅ body length, or about as long as body width ( Fig. 14B ). Chaetigers 1-3 involved in the cephalic cage (damaged); chaetae arranged in short series, chaetiger 1 dorsolateral, chaetigers 2-3 lateral. Chaetiger 1 with about 12 noto- and 10 neurochaetae, chaetigers 2-3 with about six noto- and eight neurochaetae each. Anterior dorsal margin of first chaetiger with a median multifid lobe, projected anteriorly. Anterior chaetigers with long, notopodial papillae. Chaetigers 1-3 progressively longer.Chaetal transition from cephalic cage to body chaetae abrupt; bidentate neurohooks present from chaetiger 5, each with short articles basally, medium-sized articles medially and distally; by chaetiger 20 anchylosed, bidentate neurohooks. Gonopodial lobes not seen.

Parapodia poorly developed; chaetigers 1-9 with large, globose notopodial lobe, decreasing posteriorly; in others, chaetae emerging from the body wall. Parapodia lateral; median neuropodia ventrolateral. Noto- and neuropodia with one small prechaetal papilla and three larger postchaetal papillae. Noto- and neuropodia distant to each other.

Median notochaetae arranged in short transverse series; 8-9 per bundle, as long as ¼ body width (after the swollen anterior region); all notochaetae multiarticulated capillaries, each with short articles basally, medium-sized medially, longer distally ( Fig. 14E ). Neurochaetae multiarticulated capillaries in chaetigers 1-4. Bidentate neurohooks with medium-sized articles in chaetigers 5-19 ( Fig. 14F ), being gradually replaced by anchylosed bidentate neurohooks, arranged in an oblique series, each with five per bundle ( Fig. 14G ); each neurohook with short rings continued to the subdistal straight region, blade bifid, main fang and accessory tooth almost parallel, accessory tooth thinner, sharper, shorter. Posterior region almost without sediment particles ( Fig. 14C ), tapering to a blunt cone; pygidium with large anus, as wide as pygidium, without anal cirri.

REMARKS

Trophoniella harrisae n. sp. resembles T. hospita n. comb. and T. orensanzi n. sp. because of the presence of large dorsal lappets along anterior chaetigers. They differ because in T. harrisae n. sp. the dorsal tubercles extend along about 40 chaetigers, whereas in T. hospita n. comb. they extend along about 25 chaetigers, and only along 10 chaetigers in T. orensanzi n. sp. Most of the specimens examined were previously identified as Piromis capulata (Moore, 1909) ; they differ from P. capulata , which is an intertidal and shallow-water form, in the type of neurochaetae which separates them in different genera, and about the development of paired dorsal lappets which is much reduced in P. capulata (see Salazar-Vallejo 2011a: 14).

Trophoniella havaica ( Kinberg, 1867) n. comb. ( Fig. 15 )

Chloraema havaica Kinberg, 1867: 337 , 338; 1910: 68, pl. 26, fig. 1.

Stylarioides havaica – Hartman 1949: 118, pl. 15, figs 5, 6.

Pherusa havaica – Hartman 1966: 228. — Bailey-Brock & Hartman 1987: 392, fig. 3.2.167.

TYPE MATERIAL. — North Pacific Ocean. Holotype ( SMNH 508 ), R/ V Eugenie expedition, 1851 - 1853, Oahu (21°19’N, 157°52’W), Honolulu, Hawaii, no further data.

DISTRIBUTION. — The species has been found only in the type locality, Oahu, Hawaii.

DESCRIPTION

Holotype broken in three pieces, anterior and posterior ends and a median segment ( Fig. 15A ).

Body fusiform, tapering posteriorly. Anterior fragment 5 mm long, 5 mm wide, cephalic cage 3 mm long, nine chaetigers; posterior fragment 8 mm long, 5 mm wide, 19 chaetigers (last five in regeneration). Tunic thick, finely papillated, without sediment particles. Papillae abundant, most thin, fragile, capitate or digitate; larger papillae clavate, placed along chaetal lobes and arranged in longitudinal rows, two dorsal, one lateral and four ventral ones. Anterior end not exposed; not dissected to avoid further damage. Cephalic cage chaetae as long as ³⁄₄ body width. Chaetigers 1-3 involved in the cephalic cage, chaetae arranged in short series with 3-4 noto- and 3-4 neurochaetae. Anterior dorsal margin of first chaetiger with a trifid lobe, breaking apart, tines twisted ( Fig. 15B ). Anterior chaetigers with long papillae. Chaetigers 1-3 progressively longer.Chaetal transition from cephalic cage chaetae to body chaetae abrupt, falcate anchylosed neurohooks starting in chaetiger 5 ( Fig.15C ). Gonopodial lobes not visible.

Parapodia lateral, poorly developed; chaetae emerging from the body wall ( Fig. 15E ). Median neuropodia ventrolateral. Noto- and neuropodia not forming long lobes; anterior parapodia with few longer papillae. Median notopodia with 2 - 3 infrachaetal smaller papillae and one large median chaetal lobe papilla, sometimes missing, each markedly expanded distally. Neuropodia with 1-2 infrachaetal and 1-2 suprachaetal papillae.

Median notochaetae arranged in short transverse series; 4 - 5 notochaetae per bundle, about as long as 1⁄₅ body width; all notochaetae multiarticulated capillaries, each with articles short basally and medially, long distally, but not in a regular pattern, a longer article being placed between a series of smaller ones ( Fig. 15F ). Neurochaetae multiarticulated capillaries in chaetigers 1 - 4, all articles short, tips eroded. Anchylosed, darker neurohooks from chaetiger 5, arranged in transverse series (oblique in posterior chaetigers), 3-4 in anterior and 4-5 in posterior chaetigers, barely exposed, feebly developed rings basally, not visible in the exposed part, subdistally straight, fang curved, entire ( Fig. 15G ).

Posterior end regenerating, tapered to a blunt round tip ( Fig. 15D ), with a middorsal longitudinal furrow; pygidium with anus terminal, without anal cirri.

REMARKS

Trophoniella havaica n. comb. resembles other species having anchylosed neurohooks from an anterior chaetiger and lacking sediment particles on the body, such as T. minuta n. comb., and two newly described species: T. jareckiorum n. sp. from the Antilles and T. salazarae n. sp. from the Mexican Pacific coast. The main differences are the type of parapodial papillae and the type or articulation in notochaetae; thus, T. havaica n. comb. has markedly capitate parapodial papillae, each with a marked distal expansion, whereas the other species have digitate or slightly capitate papillae. T. havaica n. comb. is also unique in having an irregular pattern of articulation, having a longer article between a series of smaller ones. The other species have a gradual change in article size throughout the chaeta.

Trophoniella hospita ( Fauchald, 1972) n. comb. ( Figs 14A ; 16 )

Piromis hospitis Fauchald, 1972: 229-231 , pl. 48, figs a-e (non P. hospitis – Blake 2000: see Trophoniella harrisae n. sp.).

TYPE MATERIAL. — Eastern tropical Pacific, Gulf of California. Holotype ( LACM-AHF 1178 ) and paratypes ( LACM-AHF 1214 ), off SW Punta Arena, Carmen Island , R / V Velero IV, stn 1746 (25°48’50”N, 111°15’00”W → 25°49’40”N, 111°15’30”W), 95-115 fathoms, sand, mud, pebbles, 18.III.1949 (paratypes 1 posterior fragment and 9 anterior fragments: 17-33 mm long, 4-5 mm wide, cephalic cage 10-13 mm long, 20-31 chaetigers, dorsal lappets present in chaetigers 2-22; first anchylosed neurohooks in chaetigers 22-28).

ADDITIONAL MATERIAL. — Eastern tropical Pacific. Anterior fragment ( LACM-AHF 2491 ), Dewey Channel, opposite San Eugenio Point, R / V Velero III, stn 1260 (27°49’50”N, 115°06’05”W → 27°49’35”N, 115°06’00” W), 23-26 fathoms, coralline rock, 27.II.1941 (68 mm long, 4 mm wide, cephalic cage 5 mm long, 68 chaetigers; dorsal lappets in chaetigers 2-25). — Anterior fragment ( LACM-AHF 2492 ), 1 mile off Perico Point, Carmen Island, Gulf of California, R/ V Velero IV, stn 1758 (25°57’30”N, 111°06’07”W → 25°57’31”N, 111°05’54”W), 15-18 fathoms, coarse sand, 21.III.1949 (52 mm long, 5 mm wide, cephalic cage 11 mm long, 40 chaetigers; dorsal lappets in chaetigers 2-22) .

DISTRIBUTION. — Gulf of California, western Mexico, in mixed bottoms at 30-230 m depth.

DESCRIPTION

Holotype (LACM-AHF 1178), anterior fragment, slightly damaged ( Fig. 14A ).Tunic papillated, with fine sediment particles, especially dorsally. Body club shaped, tapering posteriorly, stiff, 40 mm long, 7 mm wide, cephalic cage 14 mm long, 30 chaetigers. Body papillae long, capitate, arranged in longitudinal rows, two dorsally, four ventrally; dorsal papillae forming triangular, blunt lappets ( Fig. 16A, C ), directed anteriorly, present to chaetiger 26, largest in chaetigers 2-10, diminishing in size posteriorly; ventral papillae forming large, rounded lobes only in chaetigers 1-5, posterior ones smaller.

Anterior end slightly everted in holotype ( Fig.16B ). one paratype dissected to study the anterior end. Prostomium low, eyes large, brownish (posterior eyes not seen). Caruncle well developed, running through the tongue-like branchial plate. Palps very large; palp keels reduced, slightly pigmented. All lips invaginated, probably due to contraction during preservation. Branchiae cirriform, thin, sessile on a tongue-like protuberance, arranged in two lateral groups, filaments arranged in rows, more than 100 filaments per group; most shorter than palps. Nephridial lobes not seen.

Cephalic cage chaetae twice as long as body width. Chaetigers 1-3 involved in the cephalic cage; chaetae arranged in short series, dorsolateral in chaetiger 1, lateral in chaetigers 2-3; 9-10 noto- 7-8 neurochaetae per bundle, carrying peduncled epizoic organisms (hydroids?). Anterior dorsal margin of first chaetiger with a median, pentafid lobe, projected anteriorly, slightly eroded. Anterior chaetigers (1-8) with longer notopodial papillae forming notopodial lobes and dorsal tubercles, directed anteriorly, diminishing in size posteriorly. Chaetigers 1-3 of about the same size. Chaetal transition from cephalic cage to body chaetae gradual; bidentate neurohooks resembling anterior neurochaetae present from chaetiger 5; anchylosed neurohooks present from chaetiger 25. Gonopodial lobes not seen.

Parapodia better developed in anterior chaetigers; median and posterior ones with elongated notopodial papillae, chaetae emerging from the body wall. Parapodia lateral; median neuropodia ventrolateral. Noto- and neuropodia with one prechaetal papilla,one large postchaetal, and 2(-3) inferior smaller papillae.

Median notochaetae arranged in short oblique series (becoming transverse, U-shaped by chaetiger 20), 7-8 per bundle (up to 10 by chaetiger 30 in other specimens), as long as 1⁄₃ body width; all notochaetae multiarticulated capillaries, each with articles short basally, becoming long medially, longer distally ( Fig.16D , inserts).Neurochaetae multiarticulated capillaries in chaetigers 1-4; multiarticulate bidentate hooks from chaetiger 5, each with short, basal articles and long articles medially and distally, arranged in transverse series, 4-5 per neuropodium ( Fig.16E ). Anchylosed bidentate neurohooks starting by chaetiger 25 (by chaetiger 28 in other specimens); each with short, anchylosed articles present to the subdistal straight region, blade hyaline (with fine oblique striae), bidentate but accessory tooth often eroded ( Fig. 16F ). Posterior chaetigers (observed in paratypes) having most neurohooks with eroded tips, looking blunt, but a few retaining the bidentate tip. Posterior end (observed in a single posterior end in paratype lot) subdistally swollen, sediment particles restricted to the middorsal line (other posterior regions with sediment covering the whole dorsal surface); pygidium with anus dorsoterminal, no anal cirri.

REMARKS

Trophoniella hospita n. comb. resembles T. harrisae n. sp. by having dorsal paired lappets along several anterior chaetigers. They differ because these lappets are more foliose and basally narrower in T. hospita n. comb., whereas they are thicker and wider basally in T.harrisae n. sp., and because they extend through a different number of chaetigers or body length: in T. hospita n. comb. they frequently reach up to chaetiger 20, whereas in T.harrisae n. sp. they reach up to chaetiger 40. The original description regarded the posterior hooks as entire, but the tips might be eroded, as indicated by Blake (2000), and herein confirmed.

Trophoniella incerta ( Augener, 1918) n. comb. ( Fig. 17 )

Stylarioides incertus Augener, 1918: 442-444 , pl. 6, figs 156, 157, pl. 7, figs 235, 236, textfig. 65.

TYPE MATERIAL. — Lost; originally described from “West Africa”, without further details.

DISTRIBUTION. — Western Africa, probably in shallow water.

DIAGNOSIS. — Body cylindrical, slightly tapering posteriorly ( Fig. 17A ). Tunic without sediment cover; papillae minute ( Fig. 17B ). Cephalic cage chaetae slightly longer than body width. Notopodia ( Fig. 17C ), and neuropodia ( Fig. 17D ) with two long, capitate papillae. Median notochaetae with articles short basally, long medially, distally medium-sized, 7-9 per bundle ( Fig. 17E ). Anchylosed neurohooks from chaetiger 6, each with short rings present to the subdistal, straight region; median chaetigers with bidentate neurohooks ( Fig. 17F ), posterior chaetigers with blunt, unidentate tips ( Fig. 17G ).

REMARKS

Hartman (1953: 51) thought Trophoniella incerta n. comb. could be the same as Piromis arenosus because neurohooks are bifid. However, the bifid neurohook that Augener illustrated (his fig. 65b, herein redrawn as Fig. 17F ) is anchylosed, with very short rings, not provided with long articles as in Piromis . The species has apparently not been collected again and regretfully, there was no clear indication about the type locality.

Figures

FIG. 14.— Trophoniella harrisae n. sp.: A, holotypes of T.harrisae n. sp. (left,LACM-AHF 2486) and T.hospita (Fauchald,1972) n. comb. (right, LACM-AHF 1178), dorsal view; B, paratype of T. harrisae n. sp. (LACM-AHF 2488), anterior end, lateral view; C, paratype (LACM-AHF 2490), posterior end, lateral view; D, same, head dissected; palps and branchiae detached; E, paratype (LACM-AHF 2478), chaetiger 13, notochaetae (insert: basal, medial and distal regions); F, same, chaetiger 10, neurochaetae (insert: chaetal tip); G, same, chaetiger 27, neurochaetae (inserts: chaetal tips). Scale bars: A, 3.8 mm; B, 2.6 mm; C, 0.7 mm; D, 0.8 mm; E, 180 µm; F, 150 µm; G, 160 µm.

FIG. 15. — Trophoniella havaica (Kinberg, 1867) n. comb., holotype (SMNH 508): A, anterior and posterior fragments, dorsal view; B, anterior end, dorsal view; C, anterior end, ventral view; D, posterior end, ventral view; E, parapodium from median chaetiger; F, same, notochaetal basal,medial and distal regions;G, same, neurohooks.Scale bars:A, 1 mm; B, 0.4 mm; C, 0.33 mm; D, 0.8 mm; E, 0.5 mm; F, 30 µm; G, 90 µm.

FIG. 16. — Trophoniella hospita (Fauchald, 1972) n. comb.: A, holotype (LACM-AHF 1178), anterior end, dorsal view;B, same, anterior end, ventral view; C, non-type specimen (LACM-AHF 2492), anterior end, dorsal view; D, same, chaetiger 18, notochaetae (inserts: basal, medial and distal notochaetal regions); E, same, chaetiger 20, neurochaetae (inserts: medial and two enlargements of the same chaetal tip); F, same, chaetiger 30, neurochaetae (inserts: two chaetal tips). Scale bars: A, B, 1.8 mm; C, 1 mm; D, 375 µm; E, 360 µm; F, 225 µm.

FIG. 17.— Trophoniella incerta (Augener,1918) n. comb.,redrawn from Augener 1918:A, lateral view; B, anterior end,lateral view,most branchiae lost; C, median notopodium; D, median neuropodium; E, notochaeta from median chaetiger; F, neurohook from median chaetiger; G, neurohook from posterior chaetiger.