Fauveliopsis glabra (Hartman in Hartman & Barnard, 1960 )

Salazar-Vallejo, Sergio I., Zhadan, Anna E. & Rizzo, Alexandra E., 2019, Revision of Fauveliopsidae Hartman, 1971 (Annelida, Sedentaria), Zootaxa 4637 (1), pp. 1-67 : 17-21

Fauveliopsis glabra (Hartman in Hartman & Barnard, 1960)

Figures 7–9

Brada glabra Hartman in Hartman & Barnard, 1960: 129–130 , Pl. 14, Figs. 1 , 2 .

Fauveliopsis glabra: Hartman 1969: 283–284 (n. comb.); Fauchald & Hancock 1981: 38 (partim); Imajima 2009: 123–125, Fig. 40; Blake & Petersen 2000: 35–38, Figs. 3.1–3.3, frontispiece (redescr.).

Fauveliopsis rugosa Fauchald, 1972b: 220–222 , Pl. 45, Figs. a–e (partim).

TL: San Nicolas Basin, and Santa Catalina Basins, California, 1630.5 m (holotype). D: Off Southern California to Jalisco, México ( Fauchald 1972b:219–220), in 500–4997 m.

Type material. Northeastern Pacific Ocean, California. Holotype ( LACM 525 ), San Nicolas Basin, R / V Velero IV, Sta. 6341 (32°51’00” N, 119°01’12” W), 1630.5 m, olive green silty clay, glass sponge, Campbell grab, 15 Aug. 1959 (holotype bent laterally, body wall broken in anterior and near posterior ends, by dissecting parapodia; body 4.5 mm long, 0.6 mm wide, 33 chaetigers).

Additional material. Northeastern Pacific Ocean, Oregon. One specimen ( LACM 7593), Cascadia Abyssal Plain, Sta. AD74, NAD 14 (no coordinates), 1400 m, 20 Feb. 1964 (6 mm long, 0.9 mm wide, 36 chaetigers; GP on right side, posterior margin of chaetiger 10; oocytes not seen). One specimen ( USNM 74889), SW mouth of Columbia River, R/V Commando, Sta. unnumb. (45°43’ N, 125°13’ W), 1893 m, 28 May 1964, M.S. Alton, coll. (11 mm long, 1.5 mm wide, 35 chaetigers; GP in posterior margin of chaetiger 11). One specimen ( USNM 74890), SW mouth of Columbia River, R/V Commando, Sta. unnumb. (45°45’ N, 125°09’ W), 1620 m, 29 May 1964, M.S. Alton, coll. (11.5 mm long, 1 mm wide, 38 chaetigers; GP in the posterior margin of chaetiger 11). Northeastern Pacific Ocean, California. Two specimens ( LACM 7611), in Cadulus shells (and two Cadulus specimens in same vial), R/V Velero IV, Sta. 2337 (33°32’ N, 118°10’ W), 311 m, grey green mud, orange-peel grab, 1 Jul. 1953 (6.2–6.5 mm long, 0.9–1.1 mm wide, 34–37 chaetigers; GP posterior margin of chaetiger 10 or 11; oocytes not seen; inside of shell with loose sediment, not cemented particles). Four specimens ( LACM 7612), in Cadulus shells, R/V Velero IV, Sta. 6818 (33°32’53” N, 118°30’57” W), 344 m, green sandy mud, Campbell grab, 27 Jan. 1960 (3.5–6.0 mm long, 0.5–1.5 mm wide, 30–36 chaetigers; GP chaetiger 11 or 12; oocytes not seen). Six specimens ( LACM 7610), and one small paranoid, R/V Velero IV, Sta. 6834 (32°39’33” N, 119°01’24” W), 586 m, green sandy mud, Campbell grab, 29 Jan. 1960 (1.8–8.0 mm long, 0.3–2.0 mm wide, 23–37 chaetigers; GP chaetiger 11 only in largest specimen; oocytes not seen; in Cadulus or chitinoid tubes, probably a secondary shelter). Five specimens ( LACM 7613), in tubular structures, probably foraminiferan tests ( Psammosiphonella sp), R/V Velero IV, Sta. 6850 (32°29’48” N, 117°22’58” W), 2470 m, green mud, Campbell grab, 1 Feb. 1960 (5.0– 11.8 mm long, 1.3 mm wide, 37–39 chaetigers; GP chaetiger 12 or 13; oocytes not seen). Western Mexico. One specimen ( LACM 7589), off Natividad Island, R/V Velero IV, Sta. 7234 (27°38’00” N, 115°16’16” W), 844 m, mud and sand, piston corer, 2 Jan. 1961 (6.9 mm long, 1.1 mm wide, 34 chaetigers; GP single, on posterior margin of chaetiger 10, just before chaetae of chaetiger 11; oocytes 100 µm, in swollen posterior region).

Diagnosis. Fauveliopsis with 23–37 chaetigers ( Fig. 7A ). First notopodia with one large acicular and one capillary, first neuropodia with two aciculars and two capillaries per bundle; anterior and median parapodia with one large acicular and one capillary per ramus ( Fig. 7B ); posterior notopodia with one acicular and one capillary, neuropodia with 2–3 neuroaciculars and 2–3 capillaries. Chaetal formula: 2A/2A (ant.), 1A1–2c/1–2c1A (med.), 2–3A2c/2–3A2c (post.). Interramal papillae globose, sessile in median chaetigers, becoming longer and migrating dorsally in posterior chaetigers, being placed closer to notoaciculars. GP mostly single, on the right side of chaetiger 10 or 11, before chaetae of chaetiger 11 or 12 ( Fig. 7A, B ). In scaphopod shells, Cadulus ( Fig. 7D, E ), foraminiferan tests ( Psammosiphonella ), or tubular polymetallic structures.

Tubes. Tubes are subcylindrical with smooth or granulose surfaces ( Fig. 8A ); some are progressively wider, other have some enlargements basally or medially, with different pigmentation and surface patterns ( Fig. 8B, C ), but a distinctive inner mucous layer with fine sediment particles is always present ( Fig. 8 C–F). The tube wall is variable; sometimes a darker area is smooth and another, variably colored is rugose. The wall structure is also variable, sometimes with ill-defined rings ( Fig. 8C ) whereas other tubes show a distinctive ring either in the inner core as a variable width ring ( Fig. 8D ), or as a series of distinctive thin rings, being darker either externally ( Fig. 8E ), or internally ( Fig. 8F ).

Tube wall has a thin, brownish outer layer ( Fig. 9A ), but the inner structure is variable, with a paler inner layer and an irregular combination of darker layers, each with variable thickness, and sometimes with some paler round spots within darker layers ( Fig. 9B ) or darker layered or punctuated areas ( Fig. 9C, D ) indicating variable growth or mineralization patterns.

Remarks. Fauveliopsis glabra belongs in the group of species having chaetigers with three or more chaetae along medial and posterior regions, together with F. scabra and F. olgae . However, as indicated in the key above, F. glabra is the only one having smooth integument and its genital papillae are before chaetal lobe of chaetiger 11. The two other species have rugose integument and genital papillae on chaetiger 8, or none at all.

Fauveliopsis scabra was described from off California. Riser (1987) and Blake & Petersen (2000) indicated that body ends were reverted in the description. The latter authors described the species with specimens from northern localities. It should be indicated that some specimens had a ventral posterior modified region, as shown in Blake & Petersen’s figure 3.1, or in figure 1F in Petersen (2000), but because its presence is not confirmed in all specimens of similar size, it is herein regarded as a preservation artifact, and not a place of attachment as indicated by Petersen (2000).

North Atlantic records of Brada glabra by Hartman (1965:173–174) and Bellan (1969:46), or F. glabra by Hartman & Fauchald (1971:116) and López (2011:2–4, Figs 1 , 2 ) deserve confirmation. It was indicated that they have 21 chaetigers (last paragraph in B. brevis remarks, Hartman 1965:173), and later the number was modified to 23–28 and up “to 36 in larger specimens”, which differ from the about 33 chaetigers for the southern California species. Hartman & Fauchald (1971:116) indicated that there can be 20–47 chaetigers, 1–3 chaetae per bundle, and that interramal papillae are subspherical, placed closer to notopodia than to neuropodia. For the depth distribution, they indicated it can thrive in 530–5023 m depth. A hand-written label by Hartman (undated) could help clarify this confusion; the label for some non-type specimens reads: “ Fauveliopsis scabra n. sp. H (as the initial for Hartman) 1967 = Brada glabra Hartman, 1965 not 1960”. Consequently, some North Atlantic records should be referred to F. scabra , but the Northeastern Atlantic records might belong to F. adriatica Katzmann & Laubier, 1974 if they are different species, as indicated above.

This species has been reported as occupying tubular agglutinated foraminiferans such as Psammosiphonella , or Amphitremoidea Eisenack, 1937. Like other members of the group, the tests have a granulose surface and wall, and the inner tunnels are variable in Psammosiphonella ( Barker 1960, Pl. 20, Figs 1–9 ), and subcylindrical in Amphitremoidea ( Barker 1960, Pl. 20, Figs 15 –23 ). Some species of Hyperammina Brady, 1878 have smooth walls and can be slightly expanded basally ( Sen Gupta et al. 2009, Pl. 82, Figs 1–2 ).

The surface and inner tube wall features resemble those found in polymetallic nodules and crusts, as reviewed elsewhere ( Wang & Müller 2009, Kuhn et al. 2017), rather than those observed in tubular foraminiferans. It would be interesting to evaluate the role of the fauveliopsids in promoting or enhancing the mineralization processes, but that is beyond the purpose of our research activities.

Figures

FIGURE 1. Fauveliopsis adriatica Katzmann & Laubier, 1974, non-type specimens.A. Seven specimens, one in a scaphopod, three others in gastropods, and three others removed from shells (Ca: Cadulus, Ol: Olivella, Se: Seguenzia, Vi: vitrinellid, fragment), (LACM 7527). B. Another specimen (LACM 7525) preserved inside a scaphopod, methyl green stained (inset: genital papillae before chaetiger 11). C. Same, anterior end, dorsal view. D.Another specimen (LACM 7525), mature female, preserved within a gastropod shell. Scale bars: A: 1.6 mm, B: 0.9 mm, C: 140 µm, D: 0.6 mm.

FIGURE 2. Fauveliopsis armata Fauchald & Hancock, 1981, paratypes (LACM 1161). A. Methyl green stained, dorsal view, stereoscope. B. Same, dorsal view, compound microscope. C. Same, right side of anterior end. D. Same, posterior end, dorsal view. Scale bars: A, B: 250 µm, C: 60 µm, D: 80 µm.

FIGURE 7. Fauveliopsis glabra (Hartman in Hartman & Barnard, 1960), non-type specimens (LACM 7610).A. Largest specimen, dorsal view (arrow points to genital papilla). B. Same, close-up of chaetigers 9–13, right lateral view showing genital papilla (GP) over posterior margin of chaetiger 10. C. Three specimens of different size, larger ones in ventral view, smallest one in oblique dorsal view; largest specimen has a ventral region distorted (‘shield’). D. Anterior region of another specimen, left lateral view, exposed after fracture of scaphopod shell. E.Another specimen, in dorsal view, after shell was broken and sediment removed. Scale bars: A: 1.3 mm, B: 0.3 mm, C: 0.8 mm, D: 0.4 mm, E: 0.7 mm.

FIGURE 8. Fauveliopsis glabra (Hartman in Hartman & Barnard, 1960), non-types (LACM 7613), tubes. A. Tube fragments and one transverse section. B. Eight-shaped tube, opening directed upwards. C. Medial and basal fragments showing complex inner features, worms surrounded by fine sediment. D. Cross section of a medial fragment, inner cover is mucous with fine particles, median brownish core of variable width, outer tube regions more homogeneous. E. Basal fragment, outer cover darker than outer layer, inner with fine sediment particles (broken for figure 10). F. Basal fragment with darker cover inside, fine sediment particles covering tube lumen. Scale bars: A, B: 2 mm, C: 1.3 mm, D: 0.3 mm, E, F: 0.7 mm.

FIGURE 9. Fauveliopsis glabra (Hartman in Hartman & Barnard, 1960), non-types (LACM Sta. 6850), tubes 2. A. Piece broken longitudinally to show tube wall structure. B. Close-up of fragment in lower left part of A. C. Close-up of fragment in lower right part of A. D. Same, close-up of basal region. Scale bars: A: 0.9 mm, B: 0.6 mm, C: 0.5 mm, D: 0.3 mm.

FIGURE 15. Fauveliopsis magna Fauchald & Hancock, 1981, holotype (LACM 1149). A. Slightly oblique lateral view. B. Anterior region, oblique left, lateral view, methyl green stained (arrow points to genital papilla). C. Posterior region, left lateral view. Scale bars: A: 1.2 mm, B: 0.3 mm, C: 0.4 mm.

FIGURE 16. Fauveliopsis olgae Hartmann-Schröder, 1983, holotype (BMNH 1982.37). A. Dorsal view. B. Ventral view. C. Same, anterior region. D. Anterior region, right lateral view. Scale bars: A, B: 0.5 mm, C, D: 0.2 mm.

FIGURE 17. Fauveliopsis olgae Hartmann-Schröder, 1983, paratype (ZMH P-17388).A. Dorsal view. B. Right lateral view. C. Anterior region, ventral view. D. Posterior region, ventral view. E. Tubular foraminiferan fragment, external surface granulose, inner surface covered by mud. Scale bars: A, B: 0.5 mm, C, D: 0.3 mm, E: 1.8 mm.

FIGURE 18. Fauveliopsis olgae Hartmann-Schröder, 1983, non-type specimens (MNHN 901c).A. Specimens of different size, including the largest one in dorsal view. B. Small specimen, dorsal view. C. Same, ventral view. D. Medium-sized specimen, ventral view. E. Same, dorsal view, asterisk indicates genital papilla (inset: close-up of genital papilla). F. Largest specimen, anterior region, dorsal view. G: Same, ventral view. Scale bars: A: 0.6 mm, B, C: 0.3 mm, D–F: 0.5 mm, G: 1.1 mm.

FIGURE 19. Fauveliopsis rugosa Fauchald, 1972b, holotype (LACM 990). A. Anterior end fragment, oblique dorsal view. B. Same, oblique ventral view. C. Anterior region, oblique lateral view. D. Last chaetiger (inset: close up of genital papilla; arrows point to genital papillae). Scale bars: A: 0.8 mm, B: 0.7 mm, C: 110 µm, D: 90 µm.

FIGURE 20. Fauveliopsis rugosa Fauchald, 1972b, non-type specimen (Scamit 85203): A. Anterior region, right lateral view, genital papilla in chaetiger 12. B. Same, anterior region, ventral view (asterisk indicates segments enlarged in C). C. Same, chaetigers 12–13, ventral view. D. Same, chaetigers 11–13, right lateral view, showing genital papilla. E. Posterior end, right lateral view. F. Posterior region. G. Close-up of collapsed pygidium taking a subpentagonal shape. Scale bars: A: 150 µm, B: 60 µm, C: 150 µm, D: 140 µm, E: 120 µm, F: 0.6 mm, G; 70 µm.

FIGURE 21. Fauveliopsis scabra Hartman & Fauchald, 1971, holotype (LACM 900). A. Oblique, distorted, ventral view, anterior end slightly exposed. B. Anterior region, ventral view. C. Posterior region, left lateral view. D. Paratypes (LACM 901), larger one broken posteriorly, smaller one complete, oblique lateral view. E. Smaller paratype, anterior region, right lateral view (inset: chaetiger 8 with genital papilla and chaetae of chaetiger 9). F. Same, posterior region, right lateral view. Scale bars: A, D: 0.6 mm, B: 110 µm, C: 220 µm, E: 150 µm, F: 100 µm.

FIGURE 22. Fauveliopsis scabra Hartman & Fauchald, 1971, non-type specimen (LACM 7555). A. Complete specimen removed from gastropod shell, integument damaged, anterior region above. B. Another specimen, anterior region, ventral view (inset: basal regions of acicular and capillary chaeta). C. Same, head, frontal view showing smooth prostomium (Pr) and ciliated peristomium (Pe) and nuchal organs (NO). D. Specimen in A, posterior end, dorsal view. Scale bars: A: 0.55 mm, B: 0.35 mm (inset: 40 µm), C: 0.25 mm, D: 90 µm.

FIGURE 23. Laubieriopsis arenicola (Riser, 1987), holotype (USNM 97494). A. Anterior region, dorsal view. B. Posterior region, dorsal view. C. Paratype (USNM 97495), oblique lateral view. Scale bars: A: 140 µm, B: 160 µm, C: 150 µm.