Notaulax californica (Treadwell, 1906)

Tovar-Hernandez, Maria Ana Ana, Garcia-Garza, Maria Elena & de Leon-Gonzalez, Jesus Angel, 2020, Sclerozoan and fouling sabellid worms (Annelida: Sabellidae) from Mexico with the establishment of two new species, Biodiversity Data Journal 8, pp. 57471-57471 : 57471

Notaulax californica (Treadwell, 1906)

Potamilla californica Treadwell, 1906 ( Treadwell 1906): 1178.

Hypsicomus sp.- Hartman 1942: 133 (holotype of P. californica ).

Hypsicomus californicus .- Hartman 1956: 258, 262, 270; Hartman 1969: 701-702 (holotype of P. californica ).

Notaulax californica .- Perkins 1984: 342-343, fig. 31 (holotype of P. californica ).-- Yáñez-Rivera et al. 2020: 24-26, Figs. 12A-C.

Materials

Type status: Other material. Occurrence: catalogNumber: UANL 0335 ; recordedBy: Jesús Angel de León-González; individualCount: 4; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Baja California Sur; municipality: Mulegé; locality: Bahia Concepcion , Playa Santispac ; verbatimLatitude: 26°45 ’52.58” N; verbatimLongitude: 111°53 ’14.88” W; Identification: identifiedBy: Jesús Angel de León-González; Event: samplingProtocol: By hand; eventDate: May 20, 1985; year: 1985; month: 7; day: 20; habitat: Dead coral; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09 Type status: Other material. Occurrence: catalogNumber: UANL 0336 ; recordedBy: Jesús Angel de León-González; individualCount: 14; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Baja California Sur; municipality: Mulegé; locality: Bahia Concepcion , Playa Requeson ; verbatimLatitude: 26°38 ’15.01” N; verbatimLongitude: 111°49 ’52.80” W; Identification: identifiedBy: Jesús Angel de León-González; Event: samplingProtocol: By hand; eventDate: July 20, 1985; year: 1985; month: 7; day: 20; habitat: Dead coral; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09 Type status: Other material. Occurrence: catalogNumber: UANL 8134 ; recordedBy: José María Aguilar-Camacho and Irving Daniel Ramírez-Santana; individualCount: 5; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Sinaloa; municipality: Ahome; locality: Topolobampo ; maximumDepthInMeters: 1 m; verbatimLatitude: 25°33.345'N; verbatimLongitude: 109°05.74'W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 9, 2011; year: 2011; month: 8; day: 9; habitat: Buoy fouling; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09 Type status: Other material. Occurrence: catalogNumber: UANL 8135 ; recordedBy: José María Aguilar-Camacho and Irving Daniel Ramírez-Santana; individualCount: 8; sex: 4 females, 4 males; lifeStage: Adult; reproductiveCondition: Ripe; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Sinaloa; municipality: Ahome; locality: Topolobampo ; maximumDepthInMeters: 1 m; verbatimLatitude: 25°34.097'N; verbatimLongitude: 109°04.361'W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 9, 2011; year: 2011; month: 8; day: 9; habitat: Buoy fouling; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09 Type status: Other material. Occurrence: catalogNumber: UANL 8136 ; recordedBy: María Ana Tovar-Hernández; individualCount: 1; lifeStage: juvenile; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Sinaloa; municipality: Mazatlán; locality: Mazatlan , Canal de Navegacion ; maximumDepthInMeters: 1 m; verbatimLatitude: 23°12 ’13’’ N; verbatimLongitude: 106°24 ’30.1’’ W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: March 24, 2009; year: 2009; month: 3; day: 24; habitat: Buoy fouling; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09 Type status: Other material. Occurrence: catalogNumber: UANL 8137 ; recordedBy: Tulio Fabio Villalobos-Guerrero TF and José María Aguilar-Camacho; individualCount: 1; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; specificEpithet: californica (Treadwell, 1906); Location: higherGeographyID: Pacific Ocean; higherGeography: Tropical Eastern Pacific; continent: America; waterBody: Gulf of California; country: México; countryCode: MX; stateProvince: Baja California Sur; municipality: La Paz; locality: La Paz, Club de Yates Palmira ; maximumDepthInMeters: 1 m; verbatimLatitude: 24°10.992'N; verbatimLongitude: 110°18.185'W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 15, 2011; year: 2011; month: 8; day: 15; habitat: Dock fouling; Record Level: language: Spanish; institutionID: Universidad Autónoma de Nuevo León; collectionID: Colección Poliquetológica, Universidad Autónoma de Nuevo León; institutionCode: UANL; collectionCode: UANL, NL INV 0002-05-09

Description

Figures 6-9 and 10A-F

Body shape and size. Specimens flattened dorso-ventrally along the body, ripe specimens with pyriform abdomen in transverse section. Body length 5.2-11.4 mm (X = 8.15 mm, n = 7), width 0.5-1.8 mm (X = 1.02 mm, n = 7).

Radiolar crown. Length 1.7-4.8 mm (X = 2.92, n = 7 mm) with 6-11 pairs of radioles (X = 9 pairs of radioles, n = 7). Base of radiolar crown (basal lamina or radiolar lobes) short, as long as the length of first three segments in lateral view (Fig. 8 I-J). Dorsal flanges prominent (triangular in side view), ventral flanges reduced (Fig. 8 B-C, I-J). Radioles not inrolled mid-ventrally. Radioles united by a palmate membrane as long as 1/2 the length of radiolar crown (Fig. 9 B) (or as long as two times the length of base of crown or basal lamina). Radioles not sectioned to count the number of skeletal cells (see remarks to genus level). Longest pinnules located at three quarters of the radiolar crown length. Radiolar tips with broad flanges and short, digitiform tips (Fig. 8 G), occupying the space of five pinnules. Radioles with 5-6 black ocelli in single rows on both outer sides of the radioles in smallest specimens (Fig. 8 C, E); 14-16 ocelli in largest specimen (Fig. 8 D), all bands of ocelli located at 3/4 of the radiolar crown length, each band as long as space of 4-6 pinnules. Dorsal lips as long as a half of the radiolar crown length, orange, erect, with mid-rib (Fig. 8 G). Ventral lips short, ear-shaped. Dorsal and ventral pinnular appendages absent.

Peristomium. Anterior peristomial ring not exposed beyond collar (not visible), high, rounded, slightly longer ventrally. Posterior peristomial ring collar: dorsal collar margins fused to faecal groove (Fig. 8 I). Lateral collar margins entire (Fig. 8 J). Ventral collar margins incised, forming two shallow, rounded lappets (Fig. 9 A). Ventral sacs and parallel lamellae absent.

Thorax. Chaetiger 1: with straight oblique rows of spine-like notochaetae (Fig. 8 I-J). Spine-like chaetae from collar with variable shape: stouter, straight with blades terminated by an elongated conical hood inflated at the base, usually bent or wavy in the slender distal half; and slender, sharply curved at the end, provided on the convex side with a short obliquely-striated wing (Fig. 9 C and Fig. 10 A). Ventral shield rectangular, whitish (Fig. 7 B and D). Chaetigers 2-8: notopodia with superior groups of short, spine-like chaetae (Fig. 9 E and Fig. 10 C) and inferior groups of paleate chaetae without mucros (Fig. 9 D and Fig. 10 B). Neurochaetae avicular uncini, handles as long as two times the length of crest, developed breast and several rows of minute, similarly-sized teeth occupying half of crest (Fig. 9 D and Fig. 10 D); neuropodial companion chaetae with rounded denticulate head and long, gently tapering asymmetrical membrane. Ventral shields broad, rectangular, laterally indented by neuropodial tori (Fig. 7 B-C and Fig. 9 A).Fig. 10

Abdomen. Segments: 64-76 chaetigers in complete specimens (n = 2). Abdominal ventral shields dark brown, rectangular, divided longitudinally by faecal groove. Anterior abdominal segments with paleate notochaetae, nearly rounded with mucros as long as palea width (Fig. 9 F and Fig. 10 F) and elongate, narrowly-hooded chaetae. Avicular uncini similar to thoracic ones, but handles short (handle as long as the length of crest) and dentition covering 3/4 of the crest (Fig. 9 G and Fig. 10 E). Posterior abdominal segments with oval paleae with a mucro longer than three times the palea width and modified, elongate narrowly-hooded chaetae. Pygidium rounded with two black, large, reniform eyespots (Fig. 8 H).

Variation. The number of radiolar ocelli may change according to ontogeny. Smallest specimens from the same lot have only 5-6 ocelli per row, whereas largest specimens which are ripe have 14-16 ocelli per row.

Colour in live specimens. Body yellow-greenish with ventral shields cream-coloured (Fig. 6 A-B). Radiolar crown with basal half translucent-whitish (Fig. 7 B and D); distal half orange with rows of black ocelli located at three quarters of radiolar crown length (Fig. 6 B, Fig. 7 A-B and D).

Colour in preserved specimens. Body yellow (Fig. 8 I-J) or pale (Fig. 8 H), peristomium purple (Fig. 9 A) and collar and some areas of thoracic segments purple (Fig. 8 I-J). Basal half of radiolar crown purple or whitish (Fig. 7 B-C).

Tubes: Organic tubes, covered with fine sand anteriorly near mouth, translucent posteriorly (Fig. 6 A).

Diagnosis

Ventral margin of collar incised, forming rounded lappets. Short bands of radiolar ocelli (each band as long as the space of 4-6 pinnules), ocelli distributed in single rows of five ocelli (in smallest, juvenile specimen) to 16 ocelli ocelli (largest, ripe specimens), bands located at three quarters of the radiolar crown length (Table 2 ).

Biology

Females with oocytes in anterior abdomen and males with spermatozoa along abdomen (UANL 8135) with spherical nucleus and rounded cap-like acrosome.

Taxon discussion

Potamilla californica was described from Monterey Bay, California by Treadwell (1906). It was included in Hartman (1942) as Hypsicomus sp. and later as Hypsicomus californica ( Hartman 1969). The original description is very brief and does not includes figures ( Treadwell 1906). In the description provided by Hartman (1969), it is mentioned that the collar is incised mid-ventrally and she emphasises the similarity of Hypsicomus lyra Moore in Moore and Bush (originally described from Suruga Bay, Japan) and Hypsicomus californica , except for the absence of radiolar ocelli in the second species. Later, Perkins (1984) had the opportunity to review and re-describe the holotype housed at the National Museum of Natural History (USNM 5222), transferring it to the genus Notaulax and confirming the presence of radiolar ocelli.

Perkins (1984) also suggest that N. californica is not the same as N. lyra as suggested by Hartman (1969), mainly based of chaetal differences. However, to judge the original description of N. lyra , there is another difference between both species: N. lyra may present 5-20 ocelli exhibiting much irregularity in arrangement and seldom occupy the entire distance in individual radioles, ocelli may be widely separated, much crowded or even coalesced. Notaulax californica has a low and consistent number of ocelli per radiole (5-6), these are always located at 3/4 of the radiolar length and close together one to another.

Fouling specimens here reviewed from the Gulf of California match with descriptions of the holotype provided by Hartman (1969) and Perkins (1984) to N. californica , as well as those records by Yáñez-Rivera et al. (2020) to Bahía de Chamela Islands Sanctuary, Jalisco (Mexico). Some specimens are embedded within their tubes and are hard to remove from these in order to count abdominal segments.

Data available for species of Notaulax suggest the presence of gonochorism, with gametes distributed in the abdominal segments, sperm developing in tetrads and having spherical heads: Notaulax midoculi (Hoagland, 1919), N. nudicollis , N. occidentalis (Baird, 1865) ( Rouse and Fitzhugh 1994), N. lyra (Moore and Bush, 1904) ( Sanders 2014) and N. tilosaula (Schmarda, 1861) ( Tovar-Hernández et al. 2017). This is confirmed here to Notaulax californica , as well as to the two new species described below.

Figures

Figure 6. Alive specimens of Notaulax californica (Treadwell, 1906). A) Dorsal view, tube partially removed, B) ventral view, tube removed. Scale bars: A-B) 2.6 mm. A-B) UANL 8134.

Figure 7. Alive specimens of Notaulax californica (Treadwell, 1906). A) Crown partially inside tube, B, D) crown and collar, ventral views, C) thoracic tori, ventral view. Scale bars: A) 1.5 mm, B, D) 1.3 mm, C) 0.3 mm. A-D) UANL 8134.

Figure 8. Selected features of Notaulax californica (Treadwell, 1906). A) Peristomium, frontal view, crown removed, B) base of radiolar crown, lateral view, C) radiolar crown with arrows pointed to eye bands, D) radiolar ocelli of largest specimen, E) radiolar ocelli of smallest specimen, F) radiolar tips, G) dorsal lip, H) pygidial eyes, I) thorax and base of radiolar crown, dorsal view, J) same, lateral view. Scale bars: A) 0.5 mm, B-C) 1 mm, D-E) 0.15 mm, F, H) 0.2 mm, G) 0.75 mm, I-J) 0.5 mm. Abbreviations: d) dorsal, df) dorsal flange, fl) flanges, m) mouth, v) ventral. A-J) UANL 8135.

Figure 9. Selected features of Notaulax californica (Treadwell, 1906). A) Collar and first thoracic segments, ventral view, B) palmate membrane, C) chaetae from collar, D) paleae from thorax, E) broadly-hooded thoracic chaetae, F) paleae from abdomen, G) abdominal uncini. Scale bars: A) 1.3 mm, B) 2.5 mm, C, E) 40 μm, D, F) 30 μm, G) 24 μm, Abbreviation: pm) palmate membrane. A-I) UANL 8134.

Figure 10. Chaetae and uncini of species in Notaulax. A-F) Notaulax californica (Treadwell, 1906), UANL 8134; G-N) N. nigroincrustata sp. n., UANL 8138 holotype; O-S) N. punctulata sp. n., UANL 8143 paratype. A, G, O) chaetae from collar, B, H, P) paleae from thorax, C) broadly-hooded thoracic chaeta, D, I, Q) thoracic uncini, E, J, R) abdominal uncini, F, L, S) paleae from anterior abdomen, K) abdominal, elongate, narrowly-hooded chaeta, M) palea from posterior abdomen, N) posterior abdominal chaetiger. Scale bars: 30 μm.