Research Article |
Corresponding author: Víctor M. Conde-Vela ( victorconde2323@gmail.com ) Academic editor: Christopher Glasby
© 2015 Víctor M. Conde-Vela, Sergio I. Salazar-Vallejo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Conde-Vela VM, Salazar-Vallejo SI (2015) Redescriptions of Nereis oligohalina (Rioja, 1946) and N. garwoodi González-Escalante & Salazar-Vallejo, 2003 and description of N. confusa sp. n. (Annelida, Nereididae). ZooKeys 518: 15-49. https://doi.org/10.3897/zookeys.518.9564
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Type material of several polychaete species described by Enrique Rioja from Mexican coasts are lost, and the current status of some species is doubtful. Nereis oligohalina (Rioja, 1946) was described from the Gulf of Mexico, but it has been considered a junior synonym of N. occidentalis Hartman, 1945, or regarded as a distinct species with an amphiamerican distribution. On the other hand, N. garwoodi González-Escalante & Salazar-Vallejo, 2003, described from Chetumal Bay, Caribbean coasts, could be confused with N. oligohalina. In order to clarify these uncertainties, N. oligohalina is redescribed based on specimens from the Mexican Gulf of Mexico, including a proposed neotype; further, N. garwoodi is redescribed including the selection of lectotype and paralectotypes, and N. confusa sp. n. is described with material from the Gulf of California. A key for the identification of similar species and some comments about speciation in nereidid polychaetes are also included.
Amphiamerican, taxonomy, estuarine nereidids, cryptic species, Polychaeta
Among the non-marine polychaetes, the family Nereididae de Blainville, 1818 has the largest number of brackish and freshwater species (61), and 31 out of these species occur in estuaries and coastal lagoons (
Enrique Rioja documented extensively the Mexican polychaetes from Pacific or Atlantic coasts in a series of papers; unfortunately, his material is lost, and most species require designation of neotypes (
However, Nereis oligohalina and N. garwoodi have morphological differences that separate them. In this contribution our objectives were first, to redescribe N. oligohalina based upon material collected from Veracruz, including topotypes, and to propose a neotype. Second, to redescribe N. garwoodi to clarify some doubtful features in the original description, and to select lectotype and paralectotypes specimens from the syntype series. Third, to recognize what has been regarded as N. oligohalina from the Mexican Pacific as a distinct species and describe it as N. confusa sp. n., based upon material from the Gulf of California. Further, a key to identify similar Nereis species and comments about species delimitation are also included.
Specimens studied are deposited in the Reference Collection of El Colegio de la Frontera Sur, Chetumal (ECOSUR) including ethanol-fixed specimens (ECOSUR-OH), and in the Polychaetological Collection of the Universidad Autónoma de Nuevo León (UANL).
Topotypes of N. oligohalina from Estero Casitas, Nautla were examined, including additional specimens that were recently collected along the coast of Veracruz, Mexico, Gulf of Mexico. Specimens of N. confusa sp. n. from Bahía La Paz and Bahía de Los Ángeles, Gulf of California were found in unidentified material in ECOSUR, now formally deposited. To assess variation in paragnath morphology, specimens of N. pelagica Linnaeus from England (ECOSUR P2840), and Pseudonereis sp. from the Caribbean Sea (ECOSUR P1170), were also examined.
The best preserved specimens were used for designation of type material. Some specimens were fixed and preserved in 96% ethanol directly, otherwise the specimens were fixed with formalin and later preserved in 70% ethanol.
For analysis of variation, type materials and a number of non-type specimens for each species were measured. Total body length (TL), length up to chaetiger 3 (L3) or 10 (L10), width at same chaetigers (W3 and W10), number of chaetigers (nC), and length of longest tentacular cirrus (rTC) were measured with a millimeter rule under the stereomicroscope. TL was measured from palp tips to the end of the pygidium, W3 and W10 were measured excluding parapodia. Also, paragnath numbers in all areas were counted; if the pharynx was not everted, a ventral dissection was made, and areas VII-VIII were described as if the pharynx was exposed. With these results simple descriptive statistics (mean, range and standard deviation) were performed. As different fixation methods were used, a Mann-Whitney U-test was used for evaluating if there were significant differences in body measures as a result of fixation method.
For the microscopical observation of parapodial features and chaetae, right-side parapodia along body were removed and mounted in semi-permanent slides; the photographs were made with a digital camera, and distal-view drawings of parapodia were included to depict spatial disposition of ligules and chaetae. Descriptions of pigmentation patterns were included, because they are consistent and useful for recognizing the three species; other authors have noted their utility for identifying cryptic nereidid species (
Also, the reach of the dorsal cirrus and its relative length in respect of the dorsal or notopodial ligules were considered as separate attributes. For determining the reach of the dorsal cirrus, the tips of the both dorsal cirrus and dorsal or notopodial ligules were taken into account; if the dorsal cirrus tip extended beyond the tip of the dorsal or notopodial ligule, then we report “dorsal cirrus extended beyond dorsal/notopodial ligule” rather than “dorsal cirrus longer than dorsal/notopodial ligule”. On the contrary, if dorsal cirrus is shorter or not exceeding the dorsal or notopodial ligules, then we report “dorsal cirrus not extended beyond dorsal/notopodial ligule”. In the species herein treated, length of dorsal cirri and length of dorsal or notopodial ligules were generally subequal, and the dorsal cirri change their attachment along the body, but not their length necessarily.
For the designation of lectotype and neotype, the International Code of Zoological Nomenclature (
Nereis pelagica Linnaeus, 1758, by subsequent designation (
Although the species described here have more attributes than those included in the current diagnosis of the genus (
In his revision of Pseudonereis Kinberg, 1865,
The neuropodial postchaetal lobe has been considered absent for Nereis species in recent phylogenetic analyses (
Reproductive nereidids or epitokes can have two or three different regions; parapodial cirri and the pygidium are transformed is especially relevant for chemorreception, parapodial lobes or ligules are expanded and chaetae replaced for swimming (
Neanthes oligohalina
Nereis oligohalina
Veracruz, Mexico. Neotype ECOSUR 0172 and paraneotypes ECOSUR 0173 (5), mouth of Actopan River (19°25'2.95"N, 96°19'32.28"W), Chachalacas Sandbar, Gulf of Mexico, 1 m depth, on Crassostrea virginica reef, fine sediment, April 10 2012, Coll. V.M. Conde-Vela, A.E. Te-Gómez.
Veracruz, Mexico. ECOSUR P2826 (15), and ECOSUR P2827 (37), Mouth of Actopan River (19°25'2.95"N, 96°19'32.28"W), Chachalacas Sandbar, Gulf of Mexico, 1 m depth, on Crassostrea virginica reef, fine sediment, April 10 2012, Coll. C. Licona-Rosado, V.M. Conde-Vela, A.E. Te-Gómez. ECOSUR P2828 (1) Laguna de Alvarado, St. 8 (18°45'20.34"N, 95°46'29.04"W), December 6 2012, in rocks, Coll. J.M. Aguilar-Camacho. ECOSUR-OH-P0760 (23), Las Barrillas, St. 4 (18°11'20.15"N, 94°35'56.97"W), December 5 2012, on C. virginica culture, Coll. J. Cruz-Terrón. ECOSUR-OH-P0761 (15), Laguna Grande, Mandinga, St. 28 (19°1'54.96"N, 96°4'8.10"W), December 10 2012, on oyster, 11.41‰, 27.24 °C, Coll. T.F. Villalobos-Guerrero, MA. Tovar-Hernández, J.M. Aguilar-Camacho. ECOSUR-OH-P0762 (3), Laguna Grande, Mandinga, St. 26 (19°2'20.64’’ N 96°4'24.24"W), December 10 2012, on mangrove and oyster, 10.74 ‰, 26.63 °C, Coll. J.M. Aguilar-Camacho, T.F.Villalobos-Guerrero. UANL-3918 (6), Estero Casitas, Nautla, March 25 1990, Coll. A. Contreras-Arquieta.
Mouth of Actopan River (19°25'2.95"N, 96°19'32.28"W), Gulf of Mexico, in C. virginica (Gmelin) reef, in muddy sediment, 1 m depth.
Neotype complete (ECOSUR 0172), atokous female. Body tapering, 38 mm long, 2.1 mm wide, 74 chaetigers, filled with oocytes. Body yellowish, reddish brown pigmentation present dorsally on first quarter of body, discoloring towards midbody chaetigers; lateral oblique pale lines along chaetigers 1–9 (Fig.
Nereis oligohalina. Neotype female A–G (ECOSUR 0172); paraneotypes H–O (ECOSUR 0173). A Anterior end, dorsal view B Posterior end, dorsal view C Parapodium 2, anterior view D Parapodium 10, anterior view E Parapodium 46, anterior view F Parapodium 64, anterior view G Left jaw, dorsal view H Supra-acicular homogomph spiniger, parapodium 40 I Sub-acicular heterogomph spiniger, from same J Supra-acicular homogomph spiniger, from same K Notopodial homogomph falciger, from same L Supra-acicular heterogomph falciger, parapodium 28 M Sub-acicular heterogomph falciger, from same N Notopodial homogomph spiniger, parapodium 40 O Sub-acicular heterogomph spiniger, from same. Scale bars: 1 mm (A); 0.3 mm (B, G); 50 μm (C); 0.1 mm (D–F); 10 μm (H–M); 30 μm (N, O).
Prostomium 1.5 times longer than wide; antennae cirriform, slightly passing palps; eyes subequal, black, in trapezoidal arrangement (Fig.
Pharynx dissected; jaws light brown with 11 rounded teeth, extending to base (Fig.
Parapodial cirri pattern: Dorsal cirri longer than upper dorsal ligules throughout body; basally inserted on anterior region, displaced medially on midbody and posterior chaetigers. Ventral cirri shorter than neuropodial ligules throughout body, longer in few anterior chaetigers, basally inserted on anterior region, progressively distant throughout body.
First two chaetigers uniramous, remaining ones biramous. Uniramous parapodia (Fig.
In anterior parapodia (Fig.
In middle and posterior parapodia (Fig.
In anterior and midbody parapodia notochaetae homogomph spinigers; neurochaetae homogomph spinigers and heterogomph falcigers in supra-acicular fascicles, heterogomph spinigers and falcigers in sub-acicular fascicles. In posterior parapodia, notochaetae homogomph spinigers and falcigers; neurochaetae as in anterior parapodia. Chaetae decreasing rapidly in number toward posterior end.
Notopodial homogomph spinigers pectinate (i.e. blade narrow with parallel teeth), teeth decreasing distally (Fig.
Pygidium not modified; anal cirri cirriform, as long as last 5–6 chaetigers (Fig.
Male fully transformed (ECOSUR-OH-P0761) complete; body tapering, 9 mm long, 0.9 mm wide, 57 chaetigers. Partially transformed male (ECOSUR P2827) complete; body tapering, 38 mm long, 2.1 mm wide, 57 chaetigers. Partially transformed female (ECOSUR P2827) complete; body tapering, 20 mm long, 1.6 mm width, 55 chaetigers. All with body yellowish with brown pigmentation present dorsally on first quarter of body, discoloring towards midbody chaetigers; faint lateral lines (Fig.
Nereis oligohalina. Non-type male A–C, F–J, N (ECOSUR–OH–P0761); non-type partially transformed male D, L–M (ECOSUR P2827); non-type partially transformed female E, K (ECOSUR P2827). A Whole specimen, dorsal view B Anterior end, dorsal view C Posterior end, dorsal view D, E Anterior ends, dorsal view F Parapodium 1, anterior view G Parapodium 6, anterior view H Parapodium 10, frontal view I Parapodium 18, anterior view J Parapodium 31, anterior view K Parapodium 10, anterior view L Parapodium 24, anterior view M Parapodium 52, anterior view N Sesquigomph natatory chaetae, parapodium 31. Scale bars: 1 mm (A, D–E); 0.5 mm (B); 0.2 mm (C); 0.1 mm (F–M); 10 μm (N).
Prostomium longer than wide; antennae cirriform, slightly wider than those present in atokous female, as long as palps; eyes subequal, two (Fig.
Peristomium twice as long as first chaetiger, slightly pigmented; tentacular cirri present; dorsal tentacular cirri longer than ventral ones, posterodorsal ones reaching to chaetiger 10 (Fig.
Fully transformed male with pharynx everted, jaws amber with 10 teeth, inner edge toothed throughout. Maxillary ring: I = 8 cones in triangle, II = 30–32 cones in arc, III: 40 cones in rectangle, IV: 28–28 pointed cones in arc. Oral ring: V = 1 cone, VI: 4–4 pyramids in diamond, VII-VIII: 46 in two irregular rows, pyramids alternating with small cones in most-anterior row, pyramids with similar size alternating in most-posterior row.
Male body divided into two regions (Fig.
Parapodial cirri pattern: Anterior parapodia with dorsal cirri modified in chaetigers 1–7 in males, 1–5 in females; ventral cirri modified in chaetigers 1–5 in males, 1–4 in females; modification attenuated in females. Dorsal cirri subequal to upper dorsal ligules in anterior chaetigers, slightly longer throughout body; basally inserted in anterior-most region, displaced medially toward end of body. Ventral cirri shorter than neuropodial ventral ligules in unmodified chaetigers, subequal in modified region; basally inserted in anterior region, barely displacing ventrally throughout body.
Chaetigers 1–2 uniramous (Fig.
Chaetigers 3–7 in males (Fig.
Chaetigers 8–16 in males and 6–23 in females with parapodial proportions as in atokes, but with more acute ligules (Fig.
Remaining parapodia biramous, modified (Fig.
Prenatatory region with noto- and neurochaeta as in atokes, homogomph falcigers not observed, blades missing in most chaetae, number progressively reduced. In natatory region, noto- and neurochaetae sesquigomph chaetae with finely serrated, paddle-like blades (Fig.
In fully transformed males pygidium with anus surrounded by rosette of papillae (Fig.
Transformation in females discrete; small lamellae on base of dorsal cirri, neuroacicular ligules and ventral cirri (Fig.
The results of the analysis of body variation and paragnath numbers are summarized in Tables
Ranges, means and standard deviation of some body measures in three Nereis species (TL: total length, L3 and L10: length at chaetiger 3 and 10, respectively; W3 and W10: width at chaetiger 3 and 10, respectively; nC: number of chaetigers; rTC: reach of largest tentacular cirrus; SD: standard deviation).
Nereis oligohalina | TL | L3 | L10 | W3 | W10 | nC | rTC | |
---|---|---|---|---|---|---|---|---|
Formalin specimens (n = 23) |
Minimum | 10.0 | 1.1 | 1.6 | 0.6 | 0.6 | 55.0 | 4 |
Maximum | 40.0 | 3.0 | 6.3 | 2.2 | 1.9 | 80.0 | 7 | |
Mean | 24.8 | 2.3 | 3.8 | 1.6 | 1.4 | 66.9 | 5.7 | |
SD | 9.3 | 0.5 | 1.3 | 0.5 | 0.4 | 7.8 | 0.9 | |
Ethanol specimens (n = 19) |
Minimum | 9.0 | 0.6 | 1.5 | 0.7 | 0.6 | 57.0 | 5 |
Maximum | 36.0 | 3.0 | 6.5 | 2.2 | 2.0 | 78.0 | 14 | |
Mean | 20.4 | 1.5 | 3.5 | 1.4 | 1.3 | 67.0 | 9.3 | |
SD | 9.3 | 0.6 | 1.3 | 0.5 | 0.5 | 8.8 | 2.5 | |
N. garwoodi | TL | L3 | L10 | W3 | W10 | nC | rTC | |
Formalin specimens (n=33) |
Minimum | 9.0 | 0.7 | 2.4 | 1.0 | 0.8 | 46.0 | 6 |
Maximum | 36.0 | 3.0 | 6.9 | 2.0 | 2.1 | 96.0 | 13 | |
Mean | 22.1 | 1.8 | 3.8 | 1.5 | 1.5 | 72.7 | 9.3 | |
SD | 9.4 | 0.5 | 1.0 | 0.3 | 0.3 | 11.8 | 1.5 | |
N. confusa sp. n. | TL | L3 | L10 | W3 | W10 | nC | rTC | |
Formalin specimens (n=20) | Minimum | 13.0 | 1.4 | 2.6 | 1.0 | 0.9 | 66.0 | 4 |
Maximum | 35.0 | 2.5 | 5.8 | 1.9 | 1.7 | 89 | 7 | |
Mean | 24.5 | 1.8 | 4.0 | 1.5 | 1.3 | 78.6 | 5.3 | |
SD | 6.6 | 0.4 | 0.9 | 0.3 | 0.3 | 6.0 | 0.9 |
Ranges, means and standard deviations (SD) in number of paragnaths in three Nereis species (r: right, l: left).
Pharynx areas | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|
N. oligohalina (n=27) | I | II-r | II-l | III | IV-r | IV-l | V | VI-r | VI-l | VII–VIII |
Minimum | 8 | 25 | 25 | 32 | 22 | 22 | 1 | 4 | 3 | 40 |
Maximum | 15 | 38 | 37 | 71 | 45 | 39 | 2 | 4 | 4 | 47 |
Mean | 11.7 | 31.0 | 30.4 | 49.1 | 31.5 | 31.4 | 1.04 | 4.0 | 3.9 | 43.9 |
SD | 2.3 | 2.9 | 3.1 | 9.2 | 4.9 | 3.6 | 0.2 | 0.00 | 0.2 | 2.0 |
N. garwoodi (n=29) | I | II-r | II-l | III | IV-r | IV-l | V | VI-r | VI-l | VII–VIII |
Minimum | 4 | 19 | 20 | 28 | 22 | 19 | 1 | 4 | 3 | 42 |
Maximum | 19 | 42 | 40 | 59 | 43 | 38 | 1 | 4 | 4 | 46 |
Mean | 10.7 | 30.3 | 30.5 | 44.6 | 31.0 | 30.1 | 1.0 | 4.0 | 3.9 | 44.0 |
SD | 3.4 | 5.3 | 5.7 | 6.4 | 5.3 | 4.5 | 0.00 | 0.00 | 0.3 | 1.3 |
N. confusa sp. n. (n=30) | I | II-r | II-l | III | IV-r | IV-l | V | VI-r | VI-l | VII–VIII |
Minimum | 4 | 30 | 28 | 39 | 34 | 35 | 1 | 3 | 3 | 42 |
Maximum | 11 | 36 | 42 | 62 | 58 | 54 | 1 | 5 | 6 | 45 |
Mean | 7.3 | 32.3 | 34.2 | 51.2 | 45.6 | 45.1 | 1 | 3.6 | 4.0 | 42.9 |
SD | 2.1 | 2.3 | 3.9 | 8.1 | 8.9 | 6.6 | 0 | 0.7 | 0.9 | 1.1 |
In the case of N. oligohalina, fixation with 96% ethanol clearly affected the anterior portion of specimens with strong contraction of first segments, hence the tentacular cirri can reach more posterior chaetigers but without modifying their lengths, reaching up to chaetiger 14 (Table
In the maxillary ring, area I showed the least variation (Fig.
Nereis oligohalina (Rioja, 1946) is considered as a widespread species and even amphiamerican, but this stems from taxonomic confusion and the lack of type material. Designation of a neotype for N. oligohalina was considered necessary because there are no type specimens and being a problematic species, there must be an objective definition for it (
Neanthes oligohalina Rioja, 1946 was correctly transferred to Nereis by
The first synonymy involving these species was made by
Nereis oligohalina differs from N. occidentalis in some diagnostic features. In N. oligohalina there are 8–15 paragnaths on area I and 1–2 on area V, whereas in N. occidentalis there are 2–3 paragnaths on area I and no paragnaths on area V. Further, in N. oligohalina neuropodial ventral ligules are 2–3 times longer than neuroacicular ligules, but in N. occidentalis neuropodial ventral ligules from posterior chaetigers are shorter than neuroacicular ones. Regarding chaetae, and as an additional difference, in N. oligohalina the notopodial homogomph falciger has the distal tooth less developed than in N. occidentalis.
Another synonymy was made by
On the other hand, N. oligohalina has been recorded along American Atlantic coasts from northeastern Brazil, chiefly in ecological (community assemblages on Spartina alterniflora and mangroves), or population studies (secondary production and population dynamics), as well as part of taxonomic or genetic studies (
Records of N. oligohalina from the Eastern Tropical Pacific (
The species is associated with red mangrove Rhizophora mangle and with oysters. It has been found in Gulf of Mexico estuaries, including Tecolutla, Casitas-Nautla and Actopan (these estuaries have sand bars in their respective mouth rivers), and from coastal lagoons such as Mandinga and La Mancha. These systems have direct connection with the sea, some with seasonal closure of their mouths, with polyhaline to mesohaline waters (
The neotype and associated specimens were found in Crassostrea virginica (Gmelin) reef, in the Actopan river mouth. The specimens studied by
Restricted to the southwestern Gulf of Mexico.
Nereis garwoodi
Quintana Roo, Mexico. Lectotype ECOSUR 0065 and paralectotypes ECOSUR 0066 (7), Chetumal (18°29'38.88"N, 88°17'22.89"W), Chetumal Bay, 1 m depth, in calcareous sedimentary rocks, mixed bottom, September 24 1999, Coll. L.E. González-Escalante, S.I. Salazar-Vallejo.
Chetumal Bay, Quintana Roo, Mexico. ECOSUR P2829 (3), Alacranes (18°34'28.51"N, 88°14'24.21"W), May 1 1999, Coll. LEGE, SISV. ECOSUR P2830 (14), Chetumal, May 3 1999, Coll. LEGE, SISV. ECOSUR P2831 (29), Chetumal, 21 May 1999, Coll. LEGE, SISV. ECOSUR P2832 (17), Alacranes (18°34'28.51"N, 88°14'24.21"W), May 21 1999, Coll. LEGE, SISV. ECOSUR P2833 (2), Luis Echeverría (18°39'04’’N 88°12'07’’W), May 21 1999, Coll. LEGE, SISV. ECOSUR P2834 (28), Chetumal (18°29'38.88"N, 88°17'22.89"W), June 30 1999, Coll. LEGE, SISV. ECOSUR P2835 (9), Chetumal (18°29'38.88"N, 88°17'22.89"W), August 27 1999, Coll. LEGE, SISV.
Chetumal Bay, Mexico, Caribbean Sea, on rocks in mixed bottoms, 1 m depth.
Lectotype complete (ECOSUR 0065), atokous female, damaged with incisions at level of chaetigers 1, 15, and 30. Body tapering, 36 mm long, 1.7 mm wide, 95 chaetigers. Body pale, pigmentation faint, brown rectangle present dorsally on middle of anterior chaetigers, striated, discoloring toward end of body, lateral pale lines in anterior chaetigers only, oocytes present. Prostomium with brown pigment along inner margins of palps, two lines extending from antennae toward anterior pair of eyes separated by a longitudinal pale area, and two oval lateral patches; peristomium slightly pigmented, pale lines present (Fig.
Nereis garwoodi. Lectotype female A–B, D–P (ECOSUR 0065); paralectotype C (ECOSUR 0066). A Anterior end, dorsal view B Posterior end, dorsal view C Notopodial homogomph falciger, parapodium 75 D Parapodium 2, anterior view E Parapodium 9, anterior view F Parapodium 28 G Parapodium 56, anterior view H Parapodium 80, anterior view I Left jaw, dorsal view J Supra-acicular homogomph spiniger, parapodium 80 K Sub-acicular heterogomph spiniger, from same L Supra-acicular heterogomph falciger, parapodium 56 M Supra-acicular heterogomph falciger, parapodium 80 N Sub-acicular heterogomph falciger, from same O Supra-acicular heterogomph spiniger, from same P Sub-acicular heterogomph spiniger, from same. Scale bars: 1 mm (A–B, I); 50 μm (C, J–N); 0.1 mm (D–H); 0.3 mm (O–P).
Prostomium 1.5 times longer than wide; antenna cirriform, slightly passing palps; eyes subequal, black, in a rectangle (Fig.
Pharynx everted, jaws pale brown with 13 rounded teeth, extending to base (Fig.
Parapodial cirri pattern: Dorsal cirri longer than upper dorsal ligules throughout body; basally inserted on anterior region, displaced medially in midbody region, becomes subdistal in posterior chaetigers. Ventral cirri longer than neuropodial ligules in a few anterior chaetigers, progressively reduced throughout body; basally inserted on anterior region, barely migrating ventrally throughout body.
First two chaetigers uniramous, remaining ones biramous. In uniramous parapodia (Fig.
In anterior parapodia (Fig.
In middle parapodia (Fig.
In posterior parapodia (Figs
In anterior and midbody parapodia, notochaetae homogomph spinigers; neurochaetae homogomph spinigers and heterogomph falcigers in supra-acicular fascicles, heterogomph spinigers and falcigers in sub-acicular fascicles. In posterior parapodia, notochaetae homogomph spinigers and falcigers; neurochaetae as in anterior parapodia. Number of chaetae decreasing toward posterior end.
Notopodial homogomph spinigers pectinate, teeth decreasing in size distally (Fig.
Pygidium without modification; anal cirri cirriform, as long as last 4–5 segments (Fig.
Paralectotype fully transformed male (ECOSUR 0066) complete, body tapering, 9 mm long, 1 mm wide, 46 chaetigers; paralectotype partially transformed female (ECOSUR 0066) incomplete, body tapering, 16 mm long, 2 mm wide, 63 chaetigers; fully transformed female (ECOSUR P0066) complete, body tapering, 12 mm long, 1.6 mm wide, 63 chaetigers. All with body yellowish with brown pigmentation present dorsally on first quarter of body, discoloring toward midbody chaetigers (Fig.
Nereis garwoodi. Paralectotype male A, C, E, G–K (ECOSUR 0066); paralectotype female B, D, F, L–P (ECOSUR 0066). A, B Anterior ends, dorsal view. C, D Posterior ends, dorsal view E, F Whole specimens, dorsal view G Parapodium 2, anterior view H Parapodium 6, anterior view I Parapodium 10, anterior view J Parapodium 18, anterior view K Parapodium 36, anterior view L Parapodium 2, anterior view M Parapodium 5, anterior view N Parapodium 10, anterior view O Parapodium 26, anterior view P Parapodium 36, anterior view. Scale bars: A–D = 0.5 mm; E–F = 1 mm; G–P = 0.1 mm.
Prostomium longer than wide; antennae cirriform, as long as palps; eyes black, subequal, in a rectangle, three times larger than antennal basal width (Fig.
Male with pharynx everted, jaws amber with 9 teeth. Maxillary ring: I = 6 pointed cones in triangle, II = 19–20 pointed cones in arc, III = 28 pointed cones in rectangle, IV = 22–19 pointed cones in arc. Oral ring: V = 1 pointed cone, VI = 4–3 pyramids in diamond, VII-VIII = 42 in two irregular rows, P-bars alternating with small pyramids in most-anterior row, pyramids alternating with cones with similar size in most-posterior.
Male body divided into two regions (Fig.
Parapodial cirri pattern: Anterior parapodia with dorsal cirri modified in chaetigers 1–7 in males, 1–5 in females; ventral cirri modified in chaetigers 1–5 in males, 1–4 in females. Dorsal cirri subequal to upper dorsal ligules in anterior chaetigers, become subequal throughout body; basally inserted in most-anterior region, displaced medially throughout body. Ventral cirri shorter than neuropodial ligules in unmodified chaetigers, subequal in modified region; basally inserted in anterior region, barely migrating ventrally throughout body.
Chaetigers 1–2 with uniramous (Fig.
Chaetigers 3–7 in males (Fig.
Parapodial proportions as in atokous from chaetigers 8–16 in male and 6–26 in female (Fig.
Remaining parapodia modified (Fig.
Notopodial dorsal ligules subconical, longer than ventral ones in male, subequal in female; notopodial ventral ligules subconical, developing a large ventral lamella in males only, with a round projection. Neuroacicular ligules subconical, shorter than notopodial ventral ones; postchaetal lobes developing into flabellate lamellae with a round projection in dorsal edge in males, small lamellae in females, progressively increasing in size and decreasing in far posterior segments; neuropodial ventral ligules digitate, basally attached to neuroacicular ones. Ventral cirri subulate, slightly longer than neuroacicular ligules, with two basal lamellae of different sizes; dorsal cirri wider than ventral ones.
Prenatatory region with noto- and neurochaetae as in atokes, homogomph falcigers not observed. In natatory region, notochaetae and neurochaetae sesquigomph chaetae with finely serrated, paddle-like blades; atokous chaetae not completely replaced in specimens of either sex, homogomph falcigers observed in male (Fig.
Pygidium modified, anus surrounded by rosette of papillae in male, unmodified in female (Fig.
The results of the analysis of body variation and analysis of paragnath numbers are summarized in Tables
Regarding pigmentation, the striated rectangle seen in lectotype is more conspicuous in some specimens (Fig.
In an attempt to redefine the species, a lectotype has been selected (
Nereis garwoodi is closely allied with N. oligohalina, but they differ in some features in both atokous and epitokous forms, and in their habitats. In atokes, N. garwoodi never shows the dark brown coloration nor the fingerprint-like pattern found in N. oligohalina. The ranges of paragnath numbers of both species overlap and therefore are not useful to separate them, and the relative length of tentacular cirri would be useful if fixation method is the same (Table
In N. garwoodi, both dorsal and neuropodial ventral ligules are twice as long as neuroacicular ligules in uniramous chaetigers, whereas in N. oligohalina these are subequal and slightly longer, respectively. Also, in N. garwoodi the neuropodial postchaetal lobes are visible in the anterior and midbody only, whereas in N. oligohalina they are visible throughout body.
Further, N. garwoodi has notopodial ventral ligules twice as long as neuropodial ventral ones in posterior chaetigers, whereas in N. oligohalina these ligules are subequal to each other; further, in N. oligohalina neuropodial ventral ligules are medially attached in posterior chaetigers, whereas in N. garwoodi they are basally attached throughout body. Moreover, in N. garwoodi notopodial homogomph falcigers have more teeth and they are narrower than in N. oligohalina; also, in N. garwoodi the blades of supra-acicular heterogomph falcigers become broader and shorter in posterior chaetigers, but this modification is not present in N. oligohalina.
In epitokes, N. garwoodi has modified, cattail-like dorsal cirri present in biramous chaetigers with the basal sections as long as distal ones, whereas in N. oligohalina basal sections are longer. Also, in general N. garwoodi have better developed lamellae in natatory chaetigers than N. oligohalina, especially the basal lamellae of the dorsal cirri, the lamellae of both notopodial ventral and neuroacicular ligules. Moreover, epitokal transformation is more pronounced in females of N. garwoodi than in females of N. oligohalina. On the other hand, N. garwoodi is associated with calcareous rocks, while N. oligohalina is associated with reef-building bivalves and the mangrove R. mangle.
Chetumal Bay is a semi-closed, dynamic system linked to the Caribbean Sea by several freshwater tributaries, having a salinity gradient ranging 7–18 practical salinity units (psu) (
Apparently restricted to Chetumal Bay.
Nereis (Neanthes) oligohalina
Nereis oligohalina
Gulf of California, Baja California Sur. Holotype ECOSUR 0174 and paratypes ECOSUR 0175 (5), Bahía de La Paz (24°08'38.68"N, 110°20'44.40"W), March 1 2004, 70 m from shore, on wrinkled pen shell Pinna rugosa, sponges, PVC tube, and filamentous green algae, Coll. M.A. Tovar-Hernández, P. Salazar-Silva.
Gulf of California, Baja California. ECOSUR P2836 (16), Bahía de Los Ángeles (28°58'6.72"N, 113°32'43.24"W), Gulf of California, March 17 1985, on Atrina maura, Coll. E. Aguirre, C. Garza. Gulf of California, Sinaloa. ECOSUR P2837 (10), Estero el Yugo (23°18'8.30"N, 106°29'0.53"W), Mazatlán, February 24 2004, 50 cm depth, fine sediment, on filamentous green algae on mangrove roots, Coll. S. Rendón-Rodríguez, Nuri M., M.A. Tovar-Hernández, P. Salazar-Silva. Baja California Sur. ECOSUR P2838 (32), same data as holotype. ECOSUR P2839 (2), Bahía La Paz (24°12'6.51"N, 110°17'59.26"W), Gulf of California, March 2 2004, 1 m depth, on basalt rocks, sponges and algae, Coll. M.A. Tovar-Hernández, P. Salazar-Silva.
The specific name (L. confusa: confused, perplexed, troubled) indicates an earlier problematic delineation of the species; it is a noun in apposition.
Holotype complete (ECOSUR 0174), atokous female. Body tapering, 34 mm long, 1.7 mm wide, 81 chaetigers, inmature. Body yellowish, reddish brown pigmentation present dorsally on first quarter of body as three spots pattern, two lateral ones, and the other less pigmented, middorsal, forming discontinuous transverse bands up to chaetiger 10, remaining segments pale; lateral pale lines in anterior chaetigers only (Fig.
Nereis confusa sp. n. Holotype A–P (ECOSUR 0174). A Anterior end, dorsal view B Posterior end, dorsal view C Notopodial homogomph falciger, parapodium 72 D Parapodium 2, anterior view E Parapodium 11, anterior view F Parapodium 28, anterior view G Parapodium 51, anterior view H Parapodium 72, anterior view I Left jaw, dorsal view J Supra-acicular homogomph spiniger, parapodium 51 K Sub-acicular heterogomph spiniger from same L Sub-acicular heterogomph falciger from same M Supra-acicular heterogomph falciger, parapodium 72 N Supra-acicular heterogomph falciger from same O Supra-acicular homogomph spiniger from parapodium 51 P Sub-acicular heterogomph spiniger from same. Scale bars: 1 mm (A–B, I); 50 μm (C, J–N); 0.1 mm (D–H); 0.3 mm (O–P).
Prostomium longer than wide; antennae cirriform, extending beyond palps; eyes subequal, black, in a rectangle (Fig.
Pharynx dissected, jaws with 8 teeth, restricted to anteromedial edge, light brown (Fig.
Parapodial cirri pattern: Dorsal cirri longer than upper dorsal ligules throughout body; basally inserted on anterior region, displaced medially on midbody and posterior regions. Ventral cirri as long as neuropodial ligules in a few anterior chaetigers, progressively reduced throughout body; basally inserted in anterior region, migrating ventrally throughout body.
First two chaetigers uniramous, remaining biramous. In uniramous parapodia (Fig.
In anterior parapodia (Fig.
In midbody and posterior parapodia (Figs
In anterior and midbody parapodia, notochaetae homogomph spinigers; neurochaetae homogomph spinigers and heterogomph falcigers in supra-acicular fascicles, heterogomph spinigers and falcigers in sub-acicular fascicles. In posterior parapodia, notochaetae homogomph spinigers and falcigers; neurochaetae as in anterior parapodia.
Notopodial homogomph spinigers pectinate, teeth decreasing in size distally. Notopodial homogomph falcigers pectinate, 9 teeth, distal tooth stout, incurved, fused to blade (Fig.
Pygidium with broad margin, anus crenulated; anal cirri cirriform, as long as last 3–4 segments (Fig.
The results of the analysis of body variation and paragnath numbers are summarized in Table
All specimens examined show the same dorsal spotted pigmentation, but in some, especially the largest specimen, the middorsal spot disappears and only two discontinuous lines are visible along chaetigers 1–10; fingerprint-like or striated patterns were not observed. In mature specimens, the transformation starts in chaetiger 18 in males, 22 in females as previously noticed by
Variation of Nereis species studied. N. oligohalina A, D, J–L, O, Q from ECOSUR–OH–P0760. N. garwoodi B, E from paralectotype ECOSUR 0066; I, M, R from ECOSUR P2834. N. confusa sp. n. C, F–H, N, P from ECOSUR P2838. A–C Pharynges everted, dorsal view D–F Pharynges everted, ventral view G Pharynx everted, anterior view, showing merged paragnaths (arrows) H Close-up of merged paragnaths on area IV I–J Variations on area I K Variations on areas V and VI L Fingerprint-like pattern, dorsal view M–N Pigmentation patterns on anterior ends, dorsal view O Parapodial furrow in posterior end, lateral view (arrow indicates start) P Specimen on tube, dorsal view Q Parapodium from posterior chaetiger, anterior view R Ventral cirrus duplicated, chaetiger 5, anterior view. Scale bars: 0.5 mm (A–F, L); 1 mm (M–P); 0.1 mm (Q–R).
Nereis confusa sp. n. has been recorded as N. oligohalina; however, there are several differences between these two species. In N. confusa sp. n. the spotted pigmentation pattern extends up to chaetigers 10–14, and the jaws have 8 teeth restricted to the distal part of its inner edge, whereas in N. oligohalina the pale areas are replaced by fingerprint-like patterns from chaetiger 11, and its jaws have 11 teeth along its inner edge. Further, in N. confusa sp. n., both notopodial ligules and neuroacicular ligules are subequal to, or slightly longer than, neuropodial ventral ligules in midbody and posterior parapodia, whereas in N. oligohalina, they are twice as long as the neuroacicular ligules in midbody and posterior parapodia. On the other hand, N. confusa sp. n. has falcigers with broad blades, whereas in N. oligohalina they are narrower. Furthermore, in N. confusa sp. n. distal tooth of notopodial homogomph falciger is short and well developed, whereas in N. oligohalina it is longer and weakly developed.
The first records for N. confusa sp. n. (as N. oligohalina) from the Mexican Pacific were made by Berkeley and Berkeley; first, they reported Nereis (Neanthes) oligohalina males from Hipolito Bay (
Holotype found on wrinkled penshell Pinna rugosa Sowerby, 1835, sponges and filamentous green algae; other specimens were found in sponges and green algae near the type locality. Specimens from Bahía de Los Ángeles were associated with another penshell, Atrina maura (Sowerby, 1835), and specimens from Estero El Yugo were found on filamentous green algae on mangrove roots.
Gulf of California, Eastern Pacific coasts of Mexico. Probably extends to Costa Rica, in shallow water.
Some authors have emphasized the utility of reproductive patterns in taxonomy for species discrimination in closely related taxa (
Early studies considered that N. garwoodi presented a similar reproductive mode as A. virens or W. glauca, because the paralectotype female of N. garwoodi has a slight transformation; however, after further revision of additional material, a fully transformed female was found. We have no further details about its capacity for building mucous tubes or if females do not emerge to the water column or if gametes are retained in tubes.
In the material available of N. oligohalina, no completely transformed females were encountered, but perhaps with further sampling efforts they may appear. Another important consideration is the reduced size of males compared to females, even in the same sample; this has been noted for H. diversicolor (
Reproduction modes in estuarine species play a crucial role in their dispersal because the formation of planktonic larvae can determine their distribution range.
Based on the above, we hypothesize that N. oligohalina has an export strategy; it could disperse thanks to surface currents running parallel to the continental margin, and this would explain its presence along Gulf of Mexico estuaries. Similarly, distribution of N. confusa sp. n., with mainly marine habitats, could be due to current patterns along the Gulf of California.
However, sometimes the distribution patterns cannot be explained by currents and tidal dynamics. For example,
The present study demonstrates the need to encourage redescriptions of closely related and widely distributed species and, should it be necessary, the establishment of new species if there are conspicuous morphological differences.
Further, clarifying species delineation and distribution are urgent because they are essential for biogeography and phylogenetics.
(Modified from
1 | Area V without paragnaths | 2 |
– | Area V with paragnaths | 9 |
2(1) | Parapodial ligules long, slender; ceratophores distinct | N. caymanensis Fauchald, 1977 |
– | Parapodial ligules not enlarged; ceratophores indistinct | 3 |
3(2) | Notopodia with prechaetal lobes | N. goajirana Augener, 1933 |
– | Notopodia without prechaetal lobes | 4 |
4(3) | Dorsal cirri shorter than notopodial ligules; notopodial homogomph falcigers with oval blades | N. grayi Pettibone, 1956 |
– | Dorsal cirri subequal or longer than notopodial ligules; notopodial homogomph falcigers with falcate blades | 5 |
5(4) | Paragnaths on area I absent; notopodial homogomph falcigers with bifid blade | N. panamensis Fauchald, 1977 |
– | Paragnaths on area I present, notopodial homogomph falcigers with entire blade | 6 |
6(5) | Area VII-VIII with few paragnaths, usually 5–7 | 7 |
– | Area VII-VIII with numerous paragnaths, more than 40 | 8 |
7(6) | Longest tentacular cirri reaching chaetiger 7; area VI usually with 9 paragnaths | N. riisei (Grube & Ørsted in Grube, 1858) |
– | Longest tentacular cirri reaching chaetiger 4; area VI usually with 3 paragnaths | N. allenae Pettibone, 1956 |
8(6) | Notopodial homogomph falciger with numerous teeth, distal tooth recurved, one quarter of blade embedded in shaft | N. occidentalis Hartman, 1945 |
– | Notopodial homogomph falciger with few teeth, without distal tooth, one-half of blade embedded in shaft | N. pelagica Linnaeus, 1758 |
9(1) | Notopodial homogomph falcigers with cutting edge smooth | N. largoensis Treadwell, 1931 |
– | Notopodial homogomph falcigers with cutting edge denticulate | 10 |
10(9) | Dorsal pigmentation as a striated pattern, usually along first 10 chaetigers; tentacular cirri reaching beyond chaetiger 7 | 11 |
– | Dorsal pigmentation different; tentacular cirri reaching up to chaetiger 7 | 12 |
11(10) | Area I with paragnaths in an oval; longest tentacular cirri reaching up to chaetiger 9 | N. rigida Grube & Ørsted in Grube, 1858 |
– | Area I with paragnaths in a triangle; longest tentacular cirri reaching up to chaetiger 12 | N. garwoodi González-Escalante & Salazar-Vallejo, 2003 |
12(10) | Dorsal pigmentation with fingerprint-like pattern from chaetigers 10–11; notopodial ligules enlarged in posterior chaetigers | N. oligohalina (Rioja, 1946) |
– | Dorsal pigmentation with spotted pattern; notopodial ligules with similar proportions along body | N. confusa sp. n. |
Additional comments. Several species recorded from the Grand Caribbean are questionable, such as N. falcaria (Willey, 1905), N. jacksoni Kinberg, 1866, N. victoriana Augener, 1918, N. falsa de Quatrefages, 1865, N. callaona (Grube, 1857) and N. lamellosa Ehlers, 1868; consequently, they were not included in the key because their type localities are distant and different form the tropical American conditions. Only N. pelagica was included in order to contrast it with N. occidentalis, but its records from the Grand Caribbean Sea might belong to a different species.
We thank María A. Tovar-Hernández, Jesús. Á. de León-González (UANL), and Tulio F. Villalobos-Guerrero, for making available some of the material of N. oligohalina than made this study possible. We also thank Astrid E. Te-Gómez and Concepción Licona-Rosado help during fieldwork, and Luis F. Carrera-Parra by offering helpful suggestions about type-materials. This publication is part of the requirements for obtaining a M.S. degree at El Colegio de la Frontera Sur, Unidad Chetumal (ECOSUR), and supplied by a fellowship from CONACyT.
This species has not been found recently (
It has a doubtful amphiamerican status and there is no recent redescription. Here, descriptions by
After
In the original description of N. largoensis as N. brevicirrata,
Distributed in the Mexican Pacific, often confused with N. oligohalina.