FILOMENAE, Kienberger & Carmona & Pola & Padula & Gosliner & Cervera, 2016

Kienberger, Karen, Carmona, Leila, Pola, Marta, Padula, Vinicius, Gosliner, Terrence M. & Cervera, Juan Lucas, 2016, Aeolidia papillosa (Linnaeus, 1761) (Mollusca: Heterobranchia: Nudibranchia), single species or a cryptic species complex? A morphological and molecular study, Zoological Journal of the Linnean Society 177 (3), pp. 481-506 : 498-499

A EOLIDIA FILOMENAE SP. NOV.

LSID URN:LSID:ZOOBANK.ORG:ACT: F99139A8-04B3- 4F70-AA72-9F037C8FF05A

( FIGS 7C , 9C – F , 11A – C )

Aeolidia sp. A Carmona et al., 2013: 6.

Material examined. Holotype: MNCN (15.05/74475), one specimen, dissected, 25 mm in length alive, dissected, Spain, Galicia, A Coru na ~, Galicia Ribeira, iii.11, collected by Jacinto Perez Dieste. Paratype: MNCN (15.05/74474), one specimen, dissected, 45 mm in length alive, the Netherlands, Eastern Scheld , iv.12, collected by Peter H. van Bragt. Other material: MNCN (15.05/74472), one specimen, dissected, 25 mm in length alive, France, Cap Ferret , v.09, collected by Marina Poddubetskaia ; MNCN (15.05/74470), one specimen, dissected, 6 mm in length alive, France, Cap Ferret , v.09, collected by Marina Poddubetskaia ; MNCN (15.05/74471), one specimen, dissected, 12 mm in length alive, France, Cap Ferret , v.09, collected by Marina Poddubetskaia ; CASIZ 187742 , one specimen, dissected, 25 mm in length alive, Spain, Galicia, A. Coru na ~, Galicia Ribeira, iii.11, collected by Jacinto Perez Dieste ; MNCN (15.05/74473), one specimen, dissected, 17 mm preserved, Portugal, Cosata da Arrabida, Cabo Alfonso , v.12, collected by Goncalo Calado.

Type locality and habitat. Spain, Galicia, A. Coru na ~. Found under Laminaria spp. in 6 m of water .

Geographical distribution. From Scotland ( Picton & Morrow, 1994; as A. papillosa ; present study) to southern Lisbon ( Portugal; present study), including the Netherlands (present study).

Etymology. This species is dedicated to Matilde Filomena Lopez Gonzalez, born in Galicia ( Spain) and grandmother of the third author of this paper.

External morphology ( Fig. 11A–C ). The body is broad and relatively low, with a pointed posterior end of the foot. The foot corners are tentaculiform. The background colour is variable, ranging from white, light beige, salmon colour to greenish. White or lightbrown flecks are scattered all over the body. A white Y – shaped mark that runs from the oral tentacles to the pericardial area, passing between the rhinophores, may be present. This Y – shaped mark can be very evident and intense opaque white, or less noticeably beige or light brown. White or beige flecks may partly cover the Y – shaped pattern. The rhinophores are conical, blunt, and smooth. They are translucent with white or light-brown spots. The eyes are visible at the base of the rhinophores in lighter specimens ( Fig. 11B ). The oral tentacles are elongate, translucent, with opaque white or beige – brown flecks, depending on the general colour. Some specimens have white or beige tips, whereas the remaining species have translucent tips.

The cerata are flattened, broader at their base, and curved inwards. Those at the anterior region and near the posterior end of the foot are smaller than those in the middle. A bare zone from behind the rhinophores to the pericardium is present. The cerata have a lighter coloration than the rest of the body. Their apexes are white. The white – beige digestive gland is visible throughout the body wall. The cerata are arranged in up to 16 oblique rows, with each row including up to eight cerata. The anus is cleioproctic, situated between the ninth and tenth row of the right side. The gonopore is located between the fourth and fifth anterior rows of cerata.

Internal anatomy ( Figs 7C , 9C–F ). The jaws are more delicate than in A. papillosa , with a smooth masticatory edge ( Fig. 9C, E ). The radular formulae are 15 X 0.1.0 ( MNCN 15.05/74470, 6 mm) and 16 X 0.1.0 ( MNCN 15.05/74475, 25 mm). The teeth are progressively smaller towards the posterior region of the radula and pectinate with 25 – 31 denticles ( Fig. 9D, F ), which are relatively broad. Salivary glands were not found. Oral glands are absent.

The reproductive system is diaulic ( Fig. 7C ). The preampullary duct widens into the narrow and elongate ampulla. The ampulla bifurcates into the oviduct and the vas deferens. The vas deferens is extremely long and enters the wider proximal portion of the penial sac. The penial papilla is devoid of any armature. The pear-shaped receptaculum seminis joins the oviduct and enters the female gland. The vagina opens ventrally to the penis.

Remarks. Our study reveals that the specimens from several localities from the Atlantic coast of Europe, previously attributed to A. papillosa ( Carmona et al., 2013) , belong to a different species. The accepted variability in the colour pattern of the former species masked the existence of a second and pseudocryptic European species of this genus. Anatomically, both species A. papillosa and A. filomenae sp. nov. are very similar, but there are several consistent differences. The jaws of A. filomenae sp. nov. are more delicate than in A. papillosa . In addition, the number of radular teeth is significantly higher in A. papillosa : whereas A. papillosa can have 31 teeth, A. filomenae sp. nov. has a maxiumum of 16 teeth. Regarding the reproductive system, the vas deferens in A. filomenae sp. nov. is much longer than the vas deferens of A. papillosa . Also, some external differences have been observed. The cerata of A. filomenae sp. nov. are more flattened, slightly hook shaped, and usually show a paler coloration than the rest of the body, whereas the cerata in A. papillosa are usually darker and more slender.

After a careful examination of all the available literature concerning A. papillosa , it was not possible to identify our species amongst any of the specific European names considered as synonyms of A. papillosa ; however, considering the geographical range of A. filomenae sp. nov., there are several names that could match our species although their descriptions are very poor, vague, or almost inexistent. These names are:

? Doris spinis mollibus hirsute Baster, 1762: 81, plate X.

? Eolis cuvierii Lamarck, 1819: 302 .

Eolidia zetlantica Forbes & Goodsir, 1839: 647 .

Aeolis lesliana MacGillivray, 1843: 70 .

Aeolidia papillosa L., not A. papillosa (L., 1761): Walton, 1908: 227.

Eolis papillosa var. albina Dautzenberg & Durouchoux, 1913: 8 .

Aeolidia papillosa L., not A. papillosa (L., 1761): Ortea, 1980: 73.

Aeolidia papillosa L., not A. papillosa (L., 1761): Picton & Morrow, 1994: 130.

Aeolidia filomenae sp. nov. is sympatric to A. papillosa in the Netherlands and the British Isles.

Figures

Figure 7. Reproductive system: A, Aeolidia papillosa, the Netherlands (MNCN 15.05/65211); B, Aeolidia campbellii (ZSM 20041026, 20 mm), Chile; C, Aeolidia filomenae sp. nov., France; D, Aeolidia loui sp. nov. (CASIZ 102425), California, USA. Scale bars: 2.0 mm. Abbreviations: am, ampulla; fgm, female gland mass; ps, penial sac; rs, receptaculum seminis; va, vagina; vd, vas deferens.

Figure 9. Scanning electron micrographs. A, B, Aeolidia campbellii (ZSM 20041026, 20 mm); A, detailed view of the masticatory border, scale bar 1000 µm; B, radular teeth, scale bar 500 µm. C, D, Aeolidia filomenae sp. nov. (MNCN 15.05/74470); C, detailed view of the masticatory border, scale bar 300 µm; D, radular teeth, scale bar 200 µm; E, F, Aeolidia filomenae sp. nov. (MNCN 15.05/74475); E, detailed view of the masticatory border, scale bar 500 µm; F, radular teeth, scale bar 400 µm; G, D, Aeolidia loui sp. nov. (CASIZ 182214); G, detailed view of the masticatory bor- der, scale bar 10 µm; H, radular teeth, scale bar 30 µm.

Figure 11. Photographs of live Aeolidia filomenae sp. nov. specimens: A, specimen from the Netherlands, Eastern Scheld, photo by Peter H. van Bragt (MNCN 15.05/74474); B, specimen from France, Arcachon Bay, Cap Ferret, photo by Marina Poddubetskaia (MNCN 15.05/74471, 12 mm); C, specimen from Spain, Galicia, Ribeira, photo by Jacinto Perez Dieste (MNCN 15.05/ 74475, 12 mm).