Bull Mar Sci. 90(4):991–997. 2014 new taxa paper http://dx.doi.org/10.5343/bms.2014.1019

Description of a new species of Piseinotecus (Castropoda, , Piseinotecidae) from the northeastern Atlantic Ocean

1 Laboratoire des “Systèmes Naoufal Tamsouri 1 * Aquatiques: Milieu marin et 2 continental”, Département de Leila Carmona Biologie, Faculté des Sciences, Abdellatif Moukrim 1 BP8106, Cité Dakhla. 80000 Juan Lucas Cervera 2 Agadir, Morocco. 2 Departamento de Biología, Facultad de Ciencias del Mar y Ambientales, Campus de ABSTRACT.—A new species of aeolid of Excelencia Internacional del Mar the Piseinotecus Er. Marcus, 1955 is described based (CEI·MAR) Universidad de Cádiz. on several specimens from the Atlantic coast of Morocco Polígono Río San Pedro, s/n, Ap. and the southwestern Iberian Peninsula. This new species 40. 11510 Puerto Real (Cádiz), is characterized by a translucent violet ground color (very Spain. similar to several Flabellina species) with minute opaque * Corresponding author email: white spots over the cerata, latter emerge from stalks, . smooth rhinophores, uniseriate radula, and jaw with smooth masticatory border. Date Submitted: 5 March, 2014. Date Accepted: 24 April, 2014. Available Online: 10 July, 2014.

The genus Piseinotecus was introduced by Ernest Marcus (1955) and is currently constituted by only six accepted species (Cervera et al. 1987). Members of this taxon are characterized by having an uniseriate radula with strong median cusp, acleioproctic anus, penis unarmed, without penial gland, cerata in tufts arranged in rows or arches, and smooth or slightly annulate rhinophores (Er. Marcus 1955, Edmunds 1970, Schmekel and Portman 1982, Cervera et al. 1987, González-Duarte et al. 2008). So far, the genus Piseinotecus is distributed in the Atlantic Ocean, Mediterranean Sea, and western Indian Ocean (Er. Marcus 1955, Edmunds 1970, Cervera et al. 1987, Ortea et al. 1993, González-Duarte et al. 2008, Mavrič and Lipej 2012). Piseinotecus sphaeriferus (Schmekel, 1965), Piseinotecus gabinieri (Vicente, 1975), and Piseinotecus gaditanus Cervera, García-Gómez and García, 1987 are the three representatives of this genus from the eastern Atlantic Ocean and Mediterranean Sea. However, recent phylogenetic studies suggested that the genus Piseinotecus is not monophyletic (Gosliner et al. 2007, Carmona et al. 2011, 2013). Different species of this genus appear nested in the same clade of the type species of Flabellina Voight, 1834. Members of the latter genus have a triseriate radula, cerata in rows or clusters, or on short peduncles, great variability in the morphology of the rhinophores, and usually display a bright external coloration (Miller 1971, Gosliner and Willan 1991, Gosliner et al. 2008). Among all the Atlantic (including Mediterranean and Caribbean) aeolid species considered as Flabellina, only nine have a violet or pinkish ground color, which is similar to ground coloration of our new species. Indeed, the new species presented here externally resembles Flabellina.

Bulletin of Marine Science 991 © 2014 Rosenstiel School of Marine & Atmospheric Science of the University of Miami 992 Bulletin of Marine Science. Vol 90, No 4. 2014

Several field efforts along the Atlantic coast of Morocco and the southwestern Iberian Peninsula resulted in collection of specimens of an unknown aeolid, with an external appearance that resembled several Atlantic Flabellina species. However, the study of the internal anatomy was consistent with Piseinotecus. In the pres- ent study, we describe this new species, comparing it with the remaining Atlantic/ Mediterranean species of Piseinotecus, as well as with other violet/pinkish Atlantic flabellinids.

Materials and Methods

Most of samples were collected by scuba diving, except those from southern Spain that were collected intertidally. Specimens were dissected by dorsal incision. Their internal features were examined and drawn using a dissecting microscope and cam- era lucida. Special attention was paid to the morphology of the reproductive sys- tem. The buccal mass was removed and dissolved in 10% sodium hydroxide until the radula was isolated from the surrounding tissue. The radula was then rinsed in water, dried, and mounted for examination by scanning electron microscopy (SEM). The specimens examined were deposited in the Museo Nacional de Ciencias Naturales, MNCN (Madrid, Spain).

Species Description

Nudibrachia Blainville, 1814 (Odhner, 1939) Family Piseinotecidae Edmunds, 1970 Genus Piseinotecus Er. Marcus, 1955 Piseinotecus soussi new species urn:lsid:zoobank.org:pub:3A252709-D62E-4B94-BA4A-F75F1B13229A (Figs. 1–3)

Material Examined.—MNCN 15.05/67240, one specimen, dissected, 13 mm in length preserved, Morocco, Agadir, Cap Ghir (30°38´48˝N, 9°53´21˝W), 22.vi.09, col- lected by N Tamsouri; more material: MNCN 15.05/68722, one specimen, 8 mm in length preserved, Morocco, Mohammedia, David Beach (33°47´N, 7°13´W), 31.iii.10, collected by N Tamsouri; MNCN 15.05/68706, one specimen, 6 mm in length pre- served, Morocco, Agadir, Cap Ghir (30°38´48˝N, 9°53´21˝W), 27.iv.10, collected by N Tamsouri; MNCN 15.05/68711, one specimen, 11 mm in length preserved, Morocco, Agadir, Aghroud (30°36´N, 9°47´W), 13.v.10, collected by N Tamsouri; MNCN 15.05/68709, one specimen, 10 mm in length preserved, Morocco, Agadir, Aghroud (30°36´N, 9°47´W), 28.iii.09, collected by N Tamsouri; MNCN 15.05/68725, one specimen, 2 mm in length preserved, Spain, Cádiz, Santa María del Mar Beach (36°31´N, 6°17´W), 10.vii.13, collected by D Ortigosa; MNCN 15.05/68726, one speci- men, 2 mm in length preserved, Spain, Cádiz, Santa María del Mar Beach (36°31´N, 6°17´W), 10.vii.13, collected by D Ortigosa. Tamsouri et al.: New species of Piseinotecus 993

Figure 1. Photographs of the living of the genus Piseinotecus. (A) Piseinotecus sphaeriferus, specimen from Slovenia (Adriatic Sea), photo by B Mavrič; (B) Piseinotecus gabinieri, specimen from Es Caials, Cadaqués, Gerona, Spain, photo by A López-Arenas i Cama; (C) Piseinotecus gaditanus, specimen from Sesimbra, Portugal, photo by J Pedro Silva; (D–F) Piseinotecus soussi sp. nov. (D) specimen from Agadir, Cap Ghir, photo by N Tamsouri, 13.v.10, MNCN 15.05/68711; (E) specimen from Cádiz, Santa María del Mar Beach, photo by D Ortigosa, 10.vii.13, MNCN 15.05/68725; (F) specimen from Lecce, Italy, photo by F Vitale, 18.iii.14. 994 Bulletin of Marine Science. Vol 90, No 4. 2014

Figure 2. Piseinotecus soussi sp. nov. scanning electron micrographs. (A) frontal view of radular teeth, scale bar = 25 μm (MNCN 15.05/67240); (B) lateral view of radular teeth, scale bar = 100 μm (MNCN 15.05/67240); (C) complete view of the jaw, scale bar = 250 μm (MNCN 15.05/67239); (D) detailed view of the masticatory border, scale bar = 25 μm (MNCN 15.05/67239).

External Morphology (Fig. 1A–B).—The body is elongate and slender, with a long tail. The anterior margin of the foot constitutes into two short and hooked foot cor- ners. The body color, including the smooth rhinophores and oral tentacles is pinkish- purple, though the apical region of oral tentacles and rhinophores is translucent. The oral tentacles are elongate and longer than the rhinophores. The eyes are visible through the skin at the base of the rhinophores. The cerata are cylindrical, moder- ately long and translucent. The ramifications of the digestive gland are ochre or dark brown and end in a hyaline white cnidosac. The surface of some cerata has pinkish- purple pigmentation. Their subapical zone is dotted with small opaque white spots. The cerata emerge from stalks and are arranged in 7–9 groups on either side of the . Each group contains 1–13 cerata. The gonopore is located under the first group of cerata on the right, while the anus opens in an acleioproct position. Anatomy (Figs. 2A–C, 3).—The radular formulae are 27× 0.1.0 (MNCN 15.05/67240) and 25 × 0.1.0 (MNCN 15.05/67239). The radular teeth have a well-developed central cusp with 5–8 strong denticles on each side (Fig. 2A–B). The outermost denticles are smaller than the innermost. The jaws are translucent with smooth masticatory border (Fig. 2C). The reproductive system is diaulic (type II, see García-Gómez et al. 1990) (Fig. 3). The preampullary duct widens into the conspicuous, broad and bean-shaped ampul- la that narrows again before dividing into the oviduct and vas deferens. The slender Tamsouri et al.: New species of Piseinotecus 995

Figure 3. Piseinotecus soussi sp. nov. reproductive system (MNCN 15.05/67239), scale bar = 5 mm. Abbreviations: am, ampulla; bc, bursa copulatrix; fgm, female gland mass; ps, penial sac; rs, receptaculum seminis; vd, vas deferens vas deferens enters the wider proximal portion of the penial sac, which contains the unarmed penial papilla. The oviduct connects to a pyriform receptaculum seminis. There is a separate, large, and ovoid bursa copulatrix with a short and wide duct opening close to the male orifice. Etymology.—The specific name, soussi, refers to Souss Massa Drâa (province of Agadir), where most specimens of this new species were collected. Type Locality and Habitat.—Morocco, Agadir, Cap Ghir. Found in intertidal area, in tide pools and under the rocks. Geographical Distribution.—Piseinotecus soussi sp. nov. has been found along the Atlantic coast of Morocco, the southwestern Iberian Peninsula, and Italy.

Discussion

The body color of Piseinotecus soussi sp. nov. clearly separates this species from the remaining members of the genus. So far, the currently accepted species of Piseinotecus display a hyaline white or translucent body instead of pinkish-purple as in P. soussi sp. nov. Additionally, P. soussi sp. nov. is the only Piseinotecus species with smooth masticatory border (Er. Marcus 1955, Cervera et al. 1987). Nevertheless, recent studies have shown a strong intraspecific variability in the masticatory border 996 Bulletin of Marine Science. Vol 90, No 4. 2014 of other aeolids (Carmona et al. 2014a,b). Therefore, this difference between P. soussi sp. nov. and the remaining species of the genus could be due to this variability. On the other hand, nine species of Flabellina with a pinkish-purple body color are known from the Atlantic Ocean, including the Caribbean and Mediterranean seas [Flabellina affinis (Gmelin, 1791); Flabellina pedata (Montagu, 1815); Flabellina rubrolineata O’Donoghue, 1929; Flabellina llerae Ortea, 1989; Flabellina ischitana Hirano and Thompson, 1990; Flabellina marcusorum Gosliner and Kuzirian, 1990; Flabellina hamanni Gosliner, 1994; Flabellina arveloi Ortea and Espinosa, 1998; and Flabellina alternata Ortea and Espinosa, 1998] (Gosliner 1994, Ortea and Espinosa 1998, Calado et al. 2005). However, the only species with smooth rhinophores as Piseinotecus soussi sp. nov. is F. pedata. These species cannot be confused because of several external and internal features. Piseinotecus soussi sp. nov. has translucent cerata, with pinkish-purple pigmentation in some areas and conspicuous white dots, whereas the cerata of Flabellina pedata are covered by purple pigmentation, and have a subapical white ring. The cerata of the latter species arise from the notum directly and not from stalked clusters as in P. soussi sp. nov. Additionally, while F. pedata species has a triseriate radula, the radula of P. soussi is uniseriate. One of the main differences betweenPiseinotecus and Flabellina is the type of radula, uniserate, and triseriate, respectively. Nevertheless, recent phylogenetic results (Gosliner et al. 2007, Carmona et al. 2011, 2013) point to a close relationship between some species of Piseinotecus and the genus Flabellina. These phylogenies suggest that the presence of lateral teeth has less phylogenet- ic relevance than expected. Actually, Flabellina goddardi Gosliner, 2010 from the California coast was the first species of Flabellina described as having an uniseriate radula (Gosliner 2010). However, until the validity of the genus Piseinotecus could be tested properly (including the type species), we attribute this new species to the genus Piseinotecus.

Acknowledgments

Thanks to B Iazza from the University of Seville for helping us with the sampling, and J Pedro Silva, A López-Arenas i Cama (from GROC, Grupo de Recerca d’Opistobranquis de Catalunya), F Vitale and B Mavrič for providing us the photographs of other Piseinotecus species. We are deeply grateful to D Ortigosa that helped to collect and provided speci- mens and images to this study. Samplings were benefited by the following research grants: AECID-PCI A/8688/07, Spanish Agency of International Cooperation and Development; and CGL2010-17187, Spanish Ministry of Economy and Competitiveness, which includes the for- mer Ministry of Sciences and Innovation. This is CEI-MAR journal publication 63.

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