Notaulax nigroincrustata Tovar-Hernandez , Garcia-Garza & de Leon-Gonzalez, 2020

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 nigroincrustata Tovar-Hernandez, Garcia-Garza & de Leon-Gonzalez sp. n.

Materials

Type status: Holotype. Occurrence: catalogNumber: UANL 8138 ; recordedBy: José María Aguilar-Camacho and Irving Daniel Ramírez-Santana; sex: male; lifeStage: adult; reproductiveCondition: ripe; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; 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, Marina La Paz ; maximumDepthInMeters: 1 m; verbatimLatitude: 24° 09.296' N; verbatimLongitude: 110° 19.566' W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 14, 2011; year: 2011; month: 8; day: 14; 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 Type status: Paratype. Occurrence: catalogNumber: UANL 8139 ; recordedBy: José María Aguilar-Camacho and Irving Daniel Ramírez-Santana; individualCount: 2; sex: male; lifeStage: adult; reproductiveCondition: ripe; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; 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, Marina La Paz ; maximumDepthInMeters: 1 m; verbatimLatitude: 24° 09.296' N; verbatimLongitude: 110° 19.566' W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 14, 2011; year: 2011; month: 8; day: 14; 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 Type status: Paratype. Occurrence: catalogNumber: UANL 8140 ; recordedBy: Tulio Fabio Villalobos-Guerrero and José María Aguilar-Camacho; individualCount: 3; sex: 1 female, 2 males; lifeStage: adult; reproductiveCondition: ripe; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; 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, Marina La Paz ; maximumDepthInMeters: 1 m; verbatimLatitude: 24° 09.318' N; verbatimLongitude: 110° 19.630' W; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 14, 2011; year: 2011; month: 8; day: 14; 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 Type status: Other material. Occurrence: catalogNumber: UANL 8141 ; recordedBy: José María Aguilar-Camacho and Irving Daniel Ramírez-Santana; individualCount: 14; Taxon: phylum: Annelida; class: Polychaeta; order: Sabellida Levinsen, 1883; family: Sabellidae Latreille, 1825; genus: Notaulax Tauber, 1879; 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, Canal de Navegacion ; maximumDepthInMeters: 1 m; verbatimLatitude: 24° 16.447' N; verbatimLongitude: 110° 19.852' W,; Identification: identifiedBy: María Ana Tovar-Hernández; Event: samplingProtocol: By hand; eventDate: August 14, 2011; year: 2011; month: 8; day: 1; habitat: Hull 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 10G-N and 11-15

Body shape and trunk size. Specimens flattened dorso-ventrally along the body, ripe specimens with pyriform abdomen in transverse section. Body length 17.4 mm (6.3-34 mm, X = 14.3 mm, n = 4 paratypes incomplete, lacking some areas of abdomen), 2 mm width (1.2-2.7 mm, X = 2.7 mm, n = 5).

Radiolar crown. Length 9.4 mm (8-18 mm, X = 12.66, n = 5 mm) with 12 pairs of radioles (8-15 pairs of radioles, X = 12.8 radioles, n = 5). Radiolar lobes fused dorsally, whose union form a bridge internally (H-shaped). Base of radiolar crown (basal lamina or radiolar lobes) short, as long as the length of five segments in lateral view (Fig. 12 ) (3,3,3,3,5 segments each paratype). Dorsal flanges prominent (Fig. 11 A and Fig. 12 A), ventral flanges reduced (Fig. 11 B and Fig. 13 C). Radioles not inrolled mid-ventrally. Radioles fused by a palmate membrane, about 1/2 length of radiole length. There are thirteen pinnules above the palmate membrane and a group of 26-30 black ocelli in an oval group on both outer sides of the radioles, each group as long as the space of eight pinnules (Fig. 14 C-D). Radiolar flanges broad (Fig. 14 A-B). 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 flattened, erect, with broad flanges and digitiform tips, black or purple coloured (Fig. 14 B), occupying the space of 10 pinnules. Dorsal lips as long as 1/4 of the radiolar crown length, orange, erect, with mid-rib. Ventral lips short, ear-shaped. Dorsal and ventral pinnular appendages absent.

Peristomium. Anterior peristomial ring not exposed beyond collar (not visible), high, dome-shaped (Fig. 13 B). Posterior peristomial ring collar: dorsal collar margins fused to faecal groove (Fig. 12 D). Lateral collar margins entire (Fig. 12 A, C and Fig. 13 G) or with an asymmetrical lateral notch V-shaped (Fig. 12 A). Ventral collar margins entire, slightly higher than lateral collar margin (Fig. 11 B and Fig. 13 A-E). Ventral lappets, ventral sacs and parallel lamellae absent.

Thorax. Chaetiger 1: with straight oblique rows of spine-like notochaetae (Fig. 10 G and Fig. 12 A-B) or slightly curved basally (Fig. 11 A and Fig. 12 D). 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 (Fig. 10G and Fig. 15 A). Ventral shield rectangular, with an anterior whitish transverse band (Fig. 11 B and Fig. 13 A), sometimes covered partially by anterior margin of ventral collar (Fig. 13 B). Chaetigers 2-8: notopodia with superior group of short, spine-like chaetae and inferior groups of paleate chaetae without mucros. Neurochaetae avicular uncini, with handles as long as two times the length of crest, developed breast (Fig. 10 I and Fig. 15 C) and several rows of minute, similarly-sized teeth occupying half of crest length (Fig. 15 C); neuropodial companion chaetae with rounded denticulate head and long, gently tapering asymmetrical membrane. Ventral shields broad, trapezoidal, laterally indented by neuropodial tori (Fig. 13 A-E).

Abdomen. A total of 142 abdominal chaetigers (9-64 chaetigers, X = 24.8; n = 5, all incomplete specimens). Abdominal ventral shields dark brown, rectangular, divided longitudinally by faecal groove (Fig. 13 H). Anterior abdominal segments with paleate notochaetae, nearly rounded with mucros as long as palea width (Fig. 10L, Fig. 15 I, andJ-K) and elongate, narrowly-hooded chaetae (Fig. 10 K, Fig. 15 E, F and H). Avicular abdominal uncini similar to thoracic ones but handles shorter (as long as the length of crest) and dentition covering 3/4 of the crest length (Fig. 10 J and Fig. 15 D). Posterior abdominal segments with oval paleae with a mucro longer than three times the palea width (Fig. 10 M, Fig. 15 G and I) and modified, elongate narrowly-hooded chaetae (Fig. 10 N and Fig. 15 G). Pygidium unknown (incomplete specimens)Fig. 10

Variation: Amongst the set of specimens here reviewed, two were found with the lateral collar margin V-shaped, apparently natural, not damaged. Perkins (1984) (p. 338) attributed this variation to ontogeny in specimens grouped within " N. nudicollis ": collar of single lobe on adults or 2- or 4-lobed on juveniles. However, both specimens from La Paz with the lateral collar margin V-shaped were one of the largest (+34 mm) and one of medium-size (+13 mm). Thus, this anomaly cannot be attributed to ontogeny. In addition, the length of the basal lamina is also variable amongst specimens: it is as long as the length of three to five segments in lateral view (3,3,3,3,4,5,5, n = 7).

Colour in live specimens. Black thorax dorsally (Fig. 11 A). Collar ventral black with a white broad, transverse band near the base of the collar (Fig. 11 B). Anterior three thoracic segments black on ventral side (Fig. 11 B). Segments 4-8 with black colour surrounding each torus and ventral shields orange-cream coloured (Fig. 11 B). Base of radiolar crown purple-whitish (Fig. 11 A-B). Radiolar rachis and radiolar tips purple. Pinnules orange alternating with white and purple bands (Fig. 11 B). Ventral abdominal shields brown or black, with lateral sides of body yellow or cream-coloured.

Colour in preserved specimens (eight years post-fixation): black colour in life turns to deep purple-black in post-fixed specimens (Fig. 12 A-D). Other areas are cream-coloured (Fig. 12 C-D). Ventral abdominal shields purple (Fig. 13 H). Radiolar rachis and radiolar tips purple (Fig. 14 A-B).

Tubes: Organic, horny tube, translucent, some bioclaustrated in a carbonated matrix (sclerozoan) (Fig. 13 F).

Diagnosis

Ventral margin of collar entire. Short bands of radiolar ocelli (each band as long as the space of seven pinnules), ocelli distributed in oval groups of 26-30 ocelli each, groups located at the middle of the radiolar crown length.

Etymology

The name is a compound adjective, where the first term refers to the black colour of the worm (Latin nigrum, meaning black) and the second adjective (Latin incrustatĭo, meaning encrustation) makes reference to the encrusting nature of the species on the docks and hulls of the ships where the species was found.

Ecology

In La Paz (Gulf of California), Notaulax nigroincrustata sp. n, was found in hull fouling in densities of 16-40 ind.m-2. Additionally, it is remarkable that the black body pigmentation remains up tonine years after sampling.

Biology

Holotype (UANL 8138) and paratypes (UANL 8139, 8140) males with a huge mass of sperm in abdomen. Ripe worms have abdomen pyriform in cross section with a dorsal hump full of sperm. Males from additional samples (UANL 8141) have sperm between the body wall and internal layer of tubes. Spermatozoa has a spherical nucleus and rounded cap-like acrosome. Paratype female (UANL 8140) has full-developed oocytes in abdomen.

Taxon discussion

In his revision of Notaulax , Perkins (1984) includes specimens from Florida, Virgin Islands, North Carolina, Puerto Rico, Brazil, West Africa and western Mexico under the name of Notaulax nudicollis Krøyer, arguing minor differences amongst them attributed to geographic separation. However, as stated below, specimens from western Mexico, analysed in this study, have peculiar features that allow us the recognition of a separated species.

Specimens here reviewed from the Gulf of California, match only with the specimens reported from Zihuatanejo (western Mexico) by Perkins (1984) as " N. nudicollis ". Specimens from Zihuatanejo and adult specimens from La Paz have groups of up to about 30 ocelli, separated from palmate membrane by shorter length; and a collar entire ventrally. The holotype of Notaulax nudicollis has tight, elongate-oval groups of 30-70 ocelli, diminishing in number distally to form a row and separated from palmate membrane by about the length of the membrane; minute mucros in thoracic palea and mucro from the anterior abdominal palea are shorter than the palea width Perkins (1984). In specimens from western Mexico (Gulf of California, La Paz), ocelli are distributed in oval groups of 26-30 ocelli, not forming a row distally and that group is located 13 pinnules above the palmate membrane (less than a half of palmate membrane length); minute or reminiscent mucros in thoracic palea are not present; and mucros of paleate chaetae from the anterior abdomen are as long as the palea width. As these are features were constant in examined specimens, we decided to establish a new name for that species.

Amongst other species of Notaulax distributed in Western Mexico, N. nigroincrustata sp. n., differs from N. californica by the presence of an entire ventral collar margin (incised forming ventral lappets in N. californica ); and radiolar ocelli distributed in groups (single rows in N. californica ) (Table 2 ). Notaulax nigroincrustata sp. n. differs from N. punctulata sp. n. by the absence of interramal eyespots (present in N. punctulata sp. n.), amongst other features (Table 2 ).

Figures

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.

Figure 11. Alive holotype of Notaulax nigroincrustata sp. n. A) Base of crown and thorax, dorsal view, B) same, ventral view. Scale bars: A) 1 mm, B) 2 mm. A-B) UANL 8138 holotype.

Figure 12. Selected features of Notaulax nigroincrustata sp. n. A) Base of crown and thorax, lateral view, B) same, dorso-lateral view, C) same, ventro-lateral view, D) thorax, dorsal view. Scale bars: A-B) 1 mm, C) 1.5 mm, D) 2 mm, Abbreviations: cc) collar chaetiger, df) dorsal flange, vf) ventral flange. A-F) UANL 8139 paratypes.

Figure 13. Selected features of Notaulax nigroincrustata sp. n. A-E) Collar, different shapes, ventral view, F) sclerezoan worm inside a calcium carbonate matrix, G) base of radiolar crown and peristomium, lateral view, H) abdominal shields. Scale bars: A) 0.65 mm, B) 1 mm, C-D) 1 mm, E) 1.3 mm, F) 2.2 mm, G) 0.75 mm, H) 1 mm. Abbreviations: apr) anterior peristomial ring, df) dorsal flange. A) UANL 8138 holotype, B-H) UANL 8140 paratypes.

Figure 14. Selected features of Notaulax nigroincrustata sp. n. A-B) radiolar tips showing radiolar flanges, C) radiolar ocelli, D) same, detail. Scale bars: A-B) 0.3 mm, C) 0.25 mm, D) 0.1 mm. Abbreviations: fl) flange. A-D) UANL 8138 holotype.

Figure 15. Chaetae of Notaulax nigroincrustata sp. n. A) collar chaetae, B) paleae from thorax, C) thoracic uncinus, D) abdominal uncini, E) abdominal paleae and elongate, narrowly-hooded chaeta from anterior abdomen, F) chaetiger from anterior abdomen, G) chaetiger from posterior abdomen, H) palea from anterior abdomen, I) palea from posterior abdomen. Scale bars: A) 40 μm, B, E-I) 30 μm, C-D) 24 μm. A-I) UANL 8138 holotype.