Taxonomic review of Gallardoneris nonatoi (Ramos, 1976) comb. nov. (Annelida, Lumbrineridae), and description of a new species of Lumbrineris from the Gulf of Mexico Author Martin, Daniel https://orcid.org/0000-0001-6350-7384 Centre d'Estudis Avancats de Blanes (CEAB-CSIC), carrer d'acces a la Cala Sant Francesc 14, ES 17300 Blanes, Catalunya, Spain dani@ceab.csic.es Author Estefa, Jordi Department of Organismal Biology, Evolutionary Biology Centre, Uppsala University, P. O. Box 256, SE- 751 05 Upsala, Sweden Author Gil, Joao Centre of Marine Sciences (CCMAR), University of Algarve, Campus de Gambelas, 8005 - 139 Faro, Portugal text ZooKeys 2022 2022-07-25 1114 35 57 http://dx.doi.org/10.3897/zookeys.1114.79492 journal article http://dx.doi.org/10.3897/zookeys.1114.79492 1313-2970-1114-35 8DCE8B645C0846BC8133C051F56FCD0D AB4463919789537BA3F4CC6631F27C63 Gallardoneris nonatoi (Ramos, 1976) comb. nov. Figs 1 , 2 , 3 , 4 , 5 Lumbrineris nonatoi Ramos, 1976: 124-127, figs 19-21 - Campoy (1982) : 605-606; Papadopoulou et al. (1994) : 263; Gil (2011) : 398. Gallardoneris iberica Martins, Carrera-Parra, Quintino & Rodrigues, 2012: 6-10, fig. 2 - Bertasi et al. (2014) : 3-5, figs 2-4; D'Alessandro et al. (2016) : 236-237, fig. 4; Garcia Gomez et 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. All specimens 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. A anterior end B, F, K maxillary apparatus C, G, M parapodia from midbody, chaetiger 88 and chaetiger 65, respectively; pre: prechaetal lobe; post: postchaetal lobe D, H, I, N composite hooded hooks E, J, O simple hooded hooks L mandibles P posterior end A-E, P redrawn from Ramos (1976) F-J redrawn from Martins et al. (2012) K-O redrawn 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, D whole body, dorsal view B, E anterior end, dorsal view C, F posterior end, dorsal view ( C normal; F regenerating 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, F composite hooded hooks C, D simple hooded hooks A-D from the Gulf of Fos (South France, Mediterranean Sea) E from the Portuguese continental shelf (Atlantic Ocean) F from 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 nonatoi comb. 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 nonatoi comb. 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. A number of chaetigers vs. total body length (mm) B body length (mm) vs. body width at chaetiger 10 (mm) C body length (mm) vs. body width at chaetiger 15 (mm) D prostomium length (mm) vs. width (mm) E prostomium length (mm) vs. peristomium length (mm) F prostomium 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 nonatoi comb. nov. compared to Lumbrineris jan sp. nov. Abbreviations: CMHH = composite hooded hooks; L = length; SMHH = simple hooded hooks; W = width.

-	Gallardoneri nonatoi (Ramos, 1976) comb. nov.							Lumbrineris jan sp. nov.
	( Ramos 1976 )	Martins et al. (2012)	Bertasi et al. (2014)	Garcia Gomez et al. (2016)	D'Alessandro et 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 nonatoi comb. nov. characteristically lacks anal cirri, a feature seldom observed among lumbrinerids. They are also absent in members of Lumbrinerides and 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. nonatoi comb. nov. in relation to Lumbrinerides / Lumbrineriopsis would 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. nonatoi was deposited in the collections of the Museum National d'Histoire Naturelle of Paris (MNHN). However, it is not referred in the MNHN catalogue of polychaete types by Solis-Weiss et 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 ).