Lumbrineris nonatoi Gallardoneris iberica Taxonomic review of Gallardoneris nonatoi (Ramos, 1976) comb. nov. (Annelida, Lumbrineridae), and description of a new species of Lumbrineris from the Gulf of Mexico Martin, Daniel Estefa, Jordi Gil, Joao ZooKeys 2022 2022-07-25 1114 35 57 9FYBZ EC8A3ED3-6ED2-5D06-92A3-B3941328F0CA (Ramos, 1976) Martin & Estefa & Gil 2022 Ramos 1976 Polychaeta Lumbrineridae Gallardoneris CoL Animalia Gallardoneris nonatoi Eunicida 0 35 Annelida species nonatoi comb. nov.   Figs 1 , 2 , 3 , 4 , 5   Lumbrineris nonatoiRamos, 1976: 124-127, figs 19-21 - Campoy (1982): 605-606; Papadopoulou et al. (1994): 263; Gil (2011): 398.  Gallardoneris ibericaMartins, Carrera-Parra, Quintino & Rodrigues, 2012: 6-10, fig. 2 - Bertasi et al. (2014): 3-5, figs 2-4;  D'Alessandroet al. (2016): 236-237, fig. 4;  Garcia Gomezet al. (2016): 1430-1432, fig. 3A; Katsiaras et al. (2018): 1611-1614, figs 2-3. [Syn. nov].  Material examined.   Western Mediterranean Sea.CEAB AP 986A, 3 specimens, Gulf of Fos, France, approx. 43.41°N, 04.93°E, 2008, coll. Creocean; CEAB AP 986B, 2 specimens, Gulf of Fos, France, approx. 43.41°N, 04.93°E, 2009, coll. Creocean; CEAB AP 986C, 30 specimens, Quai de Sete, France, approx. 43.40°N, 3.72°E, 2009, coll. Creocean; CEAB AP 986D, 5 specimens, Port la Nouvelle, France, approx. 43.01°N, 03.08°E, 2010, coll. Creocean. Allspecimens fixed with a 10% formalin/sea water solution, stained with Rose Bengal and preserved in 70% ethanol.  Additional material.   Gallardoneris iberica:  1 paratype(DBUA0001315.01), preserved in 96% ethanol, NW continental shelf of Portugal, Atlantic Ocean, 39°48.584'N, 09°13.773'W,  100.5 m, fine sand, MeshAtlantic St.3B, coll. R. Martins, June 2010; 3 non-type specimens (DBUA0001457.01), preserved in 70% ethanol, off Barcelona, Catalonia, Spain, western Mediterranean Sea, 41°23'27.14"N, 02°12'56.58"E,  21 m, muddy sand, coll. S. Garcia Gomez, October 2013.  Description. Prostomium conical, variable, short, almost equally longer than wide, lacking antennae and eyes; peristomium shorter and clearly wider than prostomium, with two rings of similar size; junction of prostomium with peristomium normally forming an open angle (Figs 1A, 2A, B, D, E). Maxillary apparatus with four pairs of maxillae; maxillary carriers as long as maxillae I (MI), almost triangular, joined along base of MI; MI forceps-like with wide recurved base, without attachment lamella; MII stout, as long as MI, with ligament, with three teeth, without attachment lamella; MIII edentate; MIV edentate plate, with lighter central area (Fig. 1B). Mandible completely fused, Y-shaped. Prechaetal lobe inconspicuous in most anterior chaetigers, becoming ovoid and longer along body until being clearly digitiform and longer than postchaetal lobe in most posterior chaetigers; postchaetal lobe auricular in anterior segments, becoming digitiform from ca. chaetiger 20 (Fig. 1C). Bilimbate capillary chaetae from chaetiger 1 to 7-12 dorsally and from 1 to 24-30 ventrally, with limb typically forming a clearly asymmetrical basal swelling, tapering swiftly to a slender and long tip curving smoothly but clearly on same side of basal swelling. Composite multidentate hooded hooks up to chaetiger 9 (ranging from 4-12, Fig. 4A), with very short blades, 6-8 teeth either all similar or with proximal tooth bigger (due to partial fusion of several smaller teeth) and a long hood, and a characteristic second swelling at shaft end, at shaft/blade joint level (Figs 1D, 3A, B, E, F); replaced from chaetiger 10 by simple multidentate hooded hooks, with seven or eight teeth similar in size, proximal one larger due to partial fusion of several smaller teeth (Figs 1E, 3C, D). Up to four aciculae, yellow, aristate, reducing to one or two in posterior parapodia. Pygidium conical, lacking anal cirri; anus dorsal (Figs 1P, 2C, F).   Figure 1.  Gallardoneris nonatoi(Ramos, 1976) comb. nov. Aanterior end B, F, Kmaxillary apparatus C, G, Mparapodia from midbody, chaetiger 88 and chaetiger 65, respectively; pre: prechaetal lobe; post: postchaetal lobe D, H, I, Ncomposite hooded hooks E, J, Osimple hooded hooks Lmandibles Pposterior end A-E, Predrawn from Ramos (1976) F-Jredrawn from Martins et al. (2012) K-Oredrawn from Bertasi et al. (2014). Scale bars: 10 µm( D, E, N, O); 12 µm( H-J); 25 µm( G); 50 µm( A, M); 100 µm( B, C, P); 200 µm( F, K, L).   Figure 2.  Gallardoneris nonatoi(Ramos, 1976) comb. nov. Stereomicroscope photos of entire specimens. A, Dwhole body, dorsal view B, Eanterior end, dorsal view C, Fposterior end, dorsal view ( Cnormal; Fregenerating last segments). Scale bars: 1.6 mm ( A, D), 0.6 mm ( B, C, E, F).   Figure 3.  Gallardoneris nonatoi(Ramos, 1976) comb. nov. Binocular microscope photos A, B, E, Fcomposite hooded hooks C, Dsimple hooded hooks A-Dfrom the Gulf of Fos (South France, Mediterranean Sea) Efrom the Portuguese continental shelf (Atlantic Ocean) Ffrom Barcelona (Catalunya, Mediterranean Sea). Scale bar: 20 µm.  Type locality. Gulf of Roses (= Bay of Roses), Catalan Sea (northwestern Mediterranean, 42°15'6"N, 3°9'6"E), 10 m depth, sandy mud ( Ramos 1976).  Known distribution. Western and eastern Mediterranean Sea, Adriatic Sea, Aegean Sea, Levantine Sea; Atlantic coasts of the Iberian Peninsula; on sandy and muddy bottoms; 4-180 m depth.  Morphometry.   Gallardoneris nonatoicomb. nov. is a relatively small species, and the studied population ranged from slightly <5 mm to ~ 17 mm long. This character showed the largest size-related variability (130%), followed by the number of chaetigers with composite hooded hooks (100%) (Table 1, Fig. 4A). The lowest variability occurred in prostomium and peristomium width (slightly <40%), followed by the widths at chaetigers 10 and 15 (between 50% and 60%) (Table 1, Fig. 5B, C, F). However, some characters proved to be significantly size-related (Table 1, Figs 4A, B, 5A-F). The best correlation occurred between prostomium and peristomium width, followed by the number of chaetigers vs. total body length and number of chaetigers with composite hooded hooks vs. total number of chaetigers (Table 1, Figs 4A, 5A, F). The weakest correlation occurred between number of chaetigers with bilimbate chaetae and total number of chaetigers, followed by body width at chaetiger 10 and total body length (Table 1, Figs 4B, 5B). Moreover, this relationship was much weaker than that between body width at chaetiger 15 and total body length (Table 1, Fig. 5B, C), indicating that the latter better represented the total size in case on finding fragmented worms.   Table 1.Measurements and relationships (linear regressions) for the main characters of  Gallardoneris nonatoicomb. nov. Abbreviations: N= number of examined specimens; p = significance level (probability); R2 = Square Pearson coefficient.    Morphometric characters Average Minimum Maximum  N  Total body length (mm) 9.50 4.92 17.31 40  Length at chaetiger 10 (mm) 1.48 1.20 1.94 40  Length at chaetiger 15 (mm) 2.18 1.64 2.90 40  Width at chaetiger 10 (mm) 0.36 0.24 0.53 40  Width at chaetiger 15 (mm) 0.36 0.19 0.52 40  Total number of chaetigers 78.44 50 121 39  Number of chaetigers with composite hooks 7.94 4 12 16  Number of chaetigers with bilimbate chaetae 16.75 11 24 20  Prostomium length (mm) 0.24 0.15 0.35 40  Prostomium width (mm) 0.25 0.19 0.31 40  Peristomium length (mm) 0.15 0.11 0.24 40  Peristomium width (mm) 0.37 0.27 0.45 40  Linear regressions R2 p  Number of chaetigers = 35.782+4.532*Body length 0.749 <0.0001  Total body length = -2.399+32.663*Width at 10 0.187 0.005  Total body length = 0.843+23.868*Width at 15 0.383 <0.0001  Prostomium length = -0.055+1.213*Prostomium width 0.576 <0.0001  Prostomium length = 0.052+1.295*Peristomium length 0.457 <0.0001  Prostomium width = 0.028+0.595*Peristomium width 0.814 <0.0001  Chaetigers with bilimbate chaetae = 9.092+0.101*Total chaetigers 0.291 0.014  Chaetigers with composite hooks = -0.392+0.106*Total chaetigers 0.740 <0.0001   Figure 4.  Gallardoneris nonatoi(Ramos, 1976) comb. nov. Morphometric relationships showing the regression model and the intervals of confidence (95%) over the average (avg.) and observed (obs.) values. Number of chaetigers with composite hooded hooks ( A) and bilimbate chaetae ( B) vs. total number of chaetigers.   Figure 5.  Gallardoneris nonatoi(Ramos, 1976) comb. nov. Morphometric relationships showing the regression model and the intervals of confidence (95%) over the average (avg.) and observed (obs.) values. Anumber of chaetigers vs. total body length (mm) Bbody length (mm) vs. body width at chaetiger 10 (mm) Cbody length (mm) vs. body width at chaetiger 15 (mm) Dprostomium length (mm) vs. width (mm) Eprostomium length (mm) vs. peristomium length (mm) Fprostomium width (mm) vs. peristomium width (mm).  Remarks. All Mediterranean specimens studied fully agree with the original description of the species by Ramos (1976), with minor morphometric and morphological variation, both in composite and simple hooks and parapodial characteristics. Some of them may result from the size-related morphometric relationships described above, but also from differences in observation procedures and drawing interpretations (Tables 1, 2, Figs 1B-O, 3A-F, 4A, B, 5A-F).   Table 2.Main morphological characteristics of  Gallardoneris nonatoicomb. nov. compared to  Lumbrineris jansp. nov. Abbreviations: CMHH = composite hooded hooks; L = length; SMHH = simple hooded hooks; W = width.    -   Gallardoneri nonatoi(Ramos, 1976) comb. nov.   Lumbrineris jansp. nov.  ( Ramos 1976)  Martins et al. (2012)  Bertasi et al. (2014)   Garcia Gomezet al. (2016)   D'Alessandroet al. (2016)  Katsiaras et al. (2018) This paper  Carrera-Parra (2001a, 2006b); this paper  Distribution Catalan Sea Portugal Adriatic Sea Catalan Sea Tyrrhenian Sea Eastern Mediterranean French NW Mediterranean Gulf of Mexico  Prostomium conical, variable conical, variable subconical to ovoid conical, variable conical conical, variable conical, variable globular, as long as wide  L10 (mm) 0.6-1.32 1.2-1.3 0.32-0.57 - - 0.27 - 2.05 1.20-1.93 1.3 - 2.7  W10 (mm) 0.4 0.2-0.5 1.45-2.02 0.4 - 0.16 - 0.54 0.24--0.53 0.3 - 0.9  Segments (max. number) 87 101 76 78 - 82 121 -  Mandibles fused fused fused fused fused fused fused slightly divided  Number of maxillae pairs 4 4 4 4 4 4 4 5  MII (left right) 3+3 3(4)+3 3(4)+3 3+3 3+3 3+3 3+3 3+3  MIII edentate edentate edentate edentate edentate edentate edentate unidentate  MIV edentate, white central area edentate, white central area edentate, white central area edentate, white central area edentate, white central area edentate, white central area edentate, white central area unidentate  MV absent absent absent absent absent absent absent present  CMHH - - - - - - - -  Blade size* short short short short short short short short  Teeth number 7-8 7 7 7 7 up to 7 6-8 up to 5  Teeth size similar proximal bigger proximal bigger, partially fused with next one proximal bigger proximal bigger proximal bigger proximal bigger all similar  Hood long, with second swelling long, with second swelling long, with second swelling long, with second swelling long, with second swelling - long, with second swelling long, with second swelling  SMHH  First chaetiger 10 7-10 7-10 8 - 7-10 5-13 7-20  Teeth number 9 7 7 7 - up to 7 7-8 up to 6  Teeth size proximal bigger proximal bigger proximal bigger proximal bigger proximal bigger proximal bigger proximal bigger proximal bigger  SMHH hood long long long long long - long short  Preacicular size twice equal twice - - twice twice equal  Bilimbate chaetae - - - - - - - -  Last ventral (chaetiger) - 7-13 8-12 - - 7-9 7-12 19  Last dorsal (chaetiger) - 25-35 18-19 - - 17-22 24-30 -  Prechaetal lobe  Anterior (shape) ovoid inconspicuous inconspicuous - inconspicuous ovoid inconspicuous elongated  Posterior (size) long small/similar/long small/similar/long small/similar/long small/long longer small/similar/long longer  Posterior (shape) - ovoid then digitiform ovoid then digitiform - digitiform ovoid then digitiform ovoid then digitiform digitiform  Postchaetal lobe  Anterior (shape) cylindrical auricular auricular auricular - auricular auricular digitiform  Posterior (shape) digitiform digitiform digitiform digitiform digitiform digitiform digitiform digitiform  Aciculae - - - - - - - -  Dorsal curved ? curved - - ? curved several, curved  Ventral (colour) yellow yellow yellow - - yellow yellow yellow  Ventral (maximum number) 1? 2 4 - 3 4 2 up to 3  Anal cirri absent absent absent absent absent absent absent two   Gallardoneris nonatoicomb. nov. characteristically lacks anal cirri, a feature seldom observed among lumbrinerids. They are also absent in members of  Lumbrineridesand  Lumbrineriopsis, two genera that also coincide in lacking MV. However, these two genera form a cohesive group sharing other characters that are absent in  Gallardoneris, including having bidentate simple hooded hooks, lacking composite hooks, or having completely pigmented MIV ( Carrera-Parra 2006a). The lack of anal cirri in  G. nonatoicomb. nov. in relation to  Lumbrinerides/  Lumbrineriopsiswould thus be due to homoplasy. However, as the posterior region of the other species in the genus has not been fully described, it is not possible to state definitely if it is a valid generic diagnostic character. According to Ramos (1976), the holotype of  L. nonatoiwas deposited in the collections of the MuseumNational d'HistoireNaturelle of Paris (MNHN). However, it is not referred in the MNHN catalogue of polychaete types by  Solis-Weisset al. (2004), neither was it possible to locate it in this museum or in any other collection according to Carrera-Parra (2006b). However, we do not consider it necessary to designate a neotype, as the holotype might still exist in the MNHN collections, being just misplaced or mislabelled. Moreover, this taxon is clearly defined and there is no obvious exceptional need for a neotype designation, in accordance to article 75.1 of the ICZN ( International Commission on Zoological Nomenclature 1999). 2022-01-01 2022-12-31 2022-01-01 Creocean France 43.41 Western Mediterranean Sea. 686 4.93 Gulf of Fos 3 2022-01-01 2022-12-31 2022-01-01 Creocean France 43.41 Gulf of Fos 686 4.93 2 2022-01-01 2022-12-31 2022-01-01 Creocean France 43.4 Quai de Sete 686 3.72 30 2022-01-01 2022-12-31 2022-01-01 Creocean. All France 43.01 Port la Nouvelle 688 3.08 5 2010-06 R. Martins Portugal 100 39.809734 Atlantic Ocean 1 -9.22955 MeshAtlantic St. 1 paratype 2013-10 S. Garcia Gomez Spain 21 41.390873 Barcelona 1 2.2157166 western Mediterranean Sea 1 Catalonia paratype