Flabelliderma papillosa ( Essenberg, 1922 ) Salazar-Vallejo, 2007

Salazar-Vallejo, Sergio I., 2007, Revision of Flabelliderma Hartman, 1969 (Polychaeta: Flabelligeridae), Journal of Natural History 41 (33 - 36), pp. 2037-2061 : 2042-2047

Flabelliderma papillosa ( Essenberg, 1922) n. comb., re-instated

( Figure 1 )

Stylarioides papillosa Essenberg 1922, p 379 –381, Figures 1 –8.

Flabelligera essenbergae Hartman 1961, p 118 –120, Plate 22, Figures 1 , 2 , Plate 23, Figures 1–4 (partim).

Flabelliderma essenbergae: Light 1978, pp 685–686 , Figures 1–4 (partim). Type material

Eastern Pacific Ocean: neotype ( LACM-AHF 2187 ) and many neoparatypes ( LACM-AHF, MNHN, ECOSUR) collected in kelp holdfasts, off Goleta sewer outfall pipe (32 ° 24.69N, 119 ° 44.59W), Santa Barbara , California, 12 m, 18 February 2004, S. Anderson, coll.

Additional material

Two specimens (CAS-12) collected off Point Lobos (36.52 ° N, 121.95 ° W), Whaler’s Cove , Monterey , California, 20 m depth, 6 May 1972, sand, scuba-diving, A. J. Ferreira, coll.

Description

Neotype ( LACM-AHF 2187) complete, pale, fusiform, delicate, slightly damaged ( Figure 1A ). Body densely papillated; papillae forming long clavate sediment tubercles dorsally (12–14 per segment), and laterally ( Figure 1B ); smaller ventrally. Papillae long, covering noto- and neurochaetae, with adherent sediment particles. It is 13 mm long, 4 mm wide (including notopodia), cephalic cage 1.5 mm long, 27 chaetigers.

Anterior end and branchial structure based on neoparatypes. Prostomium high cone with four dark-reddish large eyes. Caruncle well developed, lateral keels elevated, thin, median one wider, swollen. Palps long, tips eroded; palp bases rounded, small. Lips damaged. Branchiae mostly lost; each lateral group with about 40 scars. Nephridial lobes long, one lost, as long as palps, placed towards the dorsal margin, separated from the branchiae ( Figure 1C ).

Cephalic cage chaetae as long as one-ninth body length, densely covered by papillae, difficult to count. Anterior dorsal margin of first chaetiger with long papillae (mostly damaged). Anterior chaetigers without especially long papillae, with large vesicles. Chaetigers 1–3 of about the same length. Chaetal transition from cephalic cage to body chaetae abrupt, neurohooks present from chaetiger 2. Gonopodial lobes not seen.

Parapodia well developed, placed on the body corners ( Figure 1D ); median neuropodia ventrolateral. Notopodial lobes ovoid with rough surface. Dorsal sediment tubercles about as long as notopodial lobes, mostly of a single size; soft, made by loosely packed large globular papillae; most papillae ovoid, with a short nipple-like distal projection. Some large vesicular papillae separated from the lobes. Neuropodia shorter lobes, masked by elongate papillae almost completely covering the neurohooks.

Median notochaetae arranged in a short transversal line; four to five multiarticulated capillaries per bundle, as long as two-thirds body width, with short articles basally and medially, longer distally. Neurochaetae multiarticulated hooks from chaetiger 2, mostly a single hook per ramus. Handle articulation with three longer articles medially, becoming progressively longer, distal articles shorter. Crest darker, tapering, acute, slightly curved distally ( Figure 1E ). Posterior end tapering; pygidium with anus terminal (as seen in neoparatypes), without anal cirri, or pigment, rarely pale brownish.

Discussion

Essenberg (1922) did not designate type material and his single specimen is lost. When Hartman (1961) changed the original name and redescribed the species, she failed to designate any of her three specimens as types, but her materials belong elsewhere (see below). Later, Light (1978) regarded two of Hartman’s specimens as syntypes, which cannot be the case because they were not employed for the original description, but did not recognize any of them as neotypes. He further expanded the species definition by introducing some rather different specimens under the same species name. Thus, in order to avoid any instability in the usage of the species name, a neotype and neoparatypes are herein designated. The specimens come from about 0.3 ° north from the type locality (San Diego, California), and were found in the same type of environment (subtidal, eelgrass roots). The other specimens indicate that size can be 8–26 mm long, with 25–27 chaetigers.

In introducing Flabelligera essenbergae as a new name, Hartman (1961, p 338) stated that ‘‘The specific name is preoccupied ( Hartman 1959, p 416) and here replaced with one in honor of the first describer’’. However, she was not aware that her materials differ from the original ones in at least one basic and relevant feature: the dorsal tubercles are different. In F. papillosa they are ‘‘long, finger-shaped, attached to the body by a narrow neck’’ ( Essenberg 1922, p 379), while Hartman (1961) provided no details of the dorsal papillae, but her illustrations show they are larger, thicker, less abundant, and with a very wide base. Therefore, F. essenbergae becomes a junior subjective synonym, because it was introduced as a replacement name, and a new name is required for the other form (see below).

Flabelliderma papillosa n. comb. is closely allied to F. lighti n. sp. by having dorsal and lateral clavate tubercles. They differ in the relative size of dorsal sediment tubercles in comparison with the notopodial lobes, and in the relative number of dorsal sediment tubercles per transverse row on each segment. In F. papillosa the dorsal sediment tubercles are larger, as long as notopodial lobes, and they are less abundant than in F. lighti n. sp. On the other hand, F. papillosa resembles F. claparedei ( de Saint-Joseph, 1898) n. comb. because they have dorsal clavate sediment tubercles; however, they differ in the relative size of their tubercles. Thus, they are of a single size in F. papillosa while they are of two different sizes in F. claparedi .

Neotype locality

The neotype locality is Santa Barbara , California, in subtidal kelp holdfasts ( ICZN 1999 , Art. 76.3).

Distribution

From Monterey to San Diego, California, in subtidal kelp holdfasts, seagrass root masses, and in shallow sandy bottoms (20 m).

Flabelliderma berkeleyorum n. sp.

( Figure 2 )

Flabelligera affinis: Berkeley and Berkeley 1960, p 792 –793 (non Sars, 1829).

Type material

Eastern Pacific Ocean: holotype (USNM-40475) and four paratypes (USNMunnumbered) collected in Friday Harbor , Washington, swimming and spawning at night light off dock, 16 July 1956, B. L. Fernald, coll. (paratypes with sediment tubercles eroded; they are 10.5–14.0 mm long, 1.0– 1.5 mm wide, cephalic cage 2 mm long, 28–30 chaetigers) .

Description

Holotype (USNM-40475) complete, pale, clavate, truncate anteriorly, tapering posteriorly, with most dorsal sediment tubercles eroded ( Figure 2A, B ). Body densely papillated; papillae small, capitate, forming rounded sediment tubercles dorsally arranged in three to four transverse rows, each with about 12–14 papillae per row, reaching the interramal region but venter apparently deprived of papillae. It is 15 mm long, 1.8 mm wide (including notopodia 4 mm), cephalic cage 2 mm long, 28 chaetigers.

Anterior end observed in one paratype. Prostomium a low cone, with four fading eyes. Caruncle well developed. Palps white, thick, slightly longer than branchial filaments; palp bases low, rounded. Lateral and dorsal lips fused, laterals massive, ventral lip reduced. Branchial groups with filaments arranged in 15 rows, each with two to five filaments, about 40 filaments per group; longest shorter than palps. Nephridial lobes large, base rounded, tips rounded, placed above the level of the eyes ( Figure 2C ).

Cephalic cage chaetae as long as one-seventh body length or as long as body width; about 70 per fascicle. Anterior dorsal margin of first chaetiger papillated. Anterior chaetigers without especially long papillae. Chaetigers 1–3 slightly decreasing in size posteriorly. Chaetal transition to body chaetae abrupt; neurohooks present from chaetiger 2. Gonopodial lobes not seen in holotype (one paratype with whitish short, rounded lobes in chaetigers 5–6).

Parapodia well developed, lateral; median neuropodia ventrolateral. Notopodial lobes cylindrical, thin, soft, surface regularly slightly nodulose, made by closely packed globular papillae ( Figure 2D, E ). Dorsal sediment tubercles rounded, smaller than notopodial lobes, all of about the same size. Neuropodia projected lobes, carrying lageniform papillae but no globular papillae.

Median notochaetae arranged in a short transverse line; multiarticulated capillaries with short articles basally, longer medially and distally; about five notochaetae per bundle; chaetae about two-thirds as long as notopodial lobe or about two-thirds as long as body width. Neurochaetae multiarticulated hooks from chaetiger 2, one per ramus. Handle articulation with short articles basally, three to four longer ones medially, then progressively shorter. Crest darker, subbasally wider, blunt, markedly curved distally ( Figure 2F ). Posterior end as a rounded lobe; pygidium with anus pale, terminal, no anal cirri.

Discussion

Flabelliderma berkeleyorum n. sp. resembles F. ockeri by having short sediment dorsal tubercles. They differ especially in the relative size of the tubercles, being larger, more or less rectangular in F. ockeri while they are short, rounded in F. berkeleyorum .

This species was caught because they were swimming and were attracted to a light ( Berkeley and Berkeley 1960). Swimming or swarming members of Flabelligera have been recorded on at least three other occasions. The first record was made by Sorby (1906, p 437; McIntosh 1915, p 112); he found that F. affinis was swimming abundantly during several years and that this abundance lasted for a few years. Sorby thought that while swimming, the adults may release their gametes. The second record was made by Gravier and Dantan (1928, p 159–160); they collected some specimens with a light trap in February and June in the Mediterranean but they were small specimens. In the third report, Herpin (1929, p 86–87) noticed many small specimens without sexual products, but in July, he found large males which released sperm when placed in flasks with sea water, and concluded that swarming may be made by just a few individuals. The fourth record was made by Berkeley and Berkeley (1960, p 792–793); they noticed five Flabelligera specimens collected with a light in Friday Harbor. They indicated that they were sexually mature and spawning; about their specific identity, they found it closer to F. affinis than to F. infundibularis , which is common in the region. However, although there are some morphological differences that might be related to reproductive transformation or epitoky, this is unlikely since there are no different chaetae and at least in the holotype there remain many dorsal sediment tubercles. Further, there were no ova in the specimens; thus, either they had spawned completely or the spherules found were rather detached dorsal sediment tubercles. It is noteworthy that Pettibone (1954, p 289) noticed that when the specimens appear in the water column, they lack any transparent tunic. This might indicate that they could be members of Flabelliderma , rather than Flabelligera , as was the case for these specimens from Friday Harbor.

Etymology

This species is named to honour the life and work of the late Edith and Cyril Berkeley. They made many relevant publications on Canadian polychaetes, especially on those living along the Pacific coast, and also made some contributions to the study of more southern polychaetes, including those living in Mexican waters.

Type locality

Friday Harbor, Washington.

Distribution

Northeastern Pacific coasts but apparently restricted to the type locality, although it might have been confused with the other common species there, Flabelligera infundibularis Johnson, 1901 .

Figures

Figure 1. Flabelliderma papillosa (Essenberg, 1922) n. comb. (A) Neotype (LACM-AHF 2187) in dorsal view; (B) neoparatype (LACM-AHF), anterior end in dorsal view; (C) another paraneotype, eversible anterior end in frontal view; branchiae and palps removed; (D) same, a complete median chaetiger in frontal view; (E) tip of neurohook from a median chaetiger. Scale bars: 1.5 mm (A); 1.0 mm (B); 0.5 mm (C, D); 90 Mm (E).

Figure 2. Flabelliderma berkeleyorum n. sp. (A) Holotype (USNM-40475) in dorsal view; (B) same, anterior end in dorsal view; (C) paratype, eversible anterior end in frontal view (palps and branchiae lost); (D) holotype, close-up of some median chaetigers, in dorsal view, showing the dorsal tubercles and notopodial lobes; (E) paratype, median parapodia showing the regularly constricted notopodial lobes; (F) paratype, tip of neurohook from a median parapodia. Scale bars: 1 mm (A); 0.4 mm (B); 0.25 mm (C, E); 0.33 mm (D); 40 Mm (F).