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The study of a well-preserved collection of discodorid nudibranchs collected in Koumac, New Caledonia, revealed the presence of seven species new to science belonging to the genera Atagema, Jorunna, Rostanga, and Sclerodoris, although some of the generic assignments are tentative as the phylogeny of Discodorididae remains unresolved. Moreover, a poorly known species of Atagema originally described from New Caledonia is re-described and the presence of Sclerodoris tuberculata in New Caledonia is confirmed with molecular data. All the species described herein are highly cryptic on their food source and in the context of the present study the term “cryptic” is used to denote such species. This paper highlights the importance of comprehensive collecting efforts to identify and document well-camouflaged taxa.

Species of Kaloplocamus Bergh, 1880 are enigmatic Nudibranchia sea slugs, and only two valid species are reported in the northwestern Pacific. Kaloplocamus japonicus (Bergh, 1880) was initially described based on alcohol-fixed specimens. In the latest revision of Kaloplocamus, it was synonymized with Kaloplocamus ramosus (Cantraine, 1835). Recently, several nudibranchs were collected from Tianheng, Shandong Province, China, and one of them is identified as an undescribed species described here as Kaloplocamus albopunctatus sp. nov. based on integrated approaches incorporating morphological observations, internal anatomy, and phylogenetic analyses of two mitochondrial (COI, 16S rRNA) genes. The other species is identified as K. japonicus Bergh, 1880 based on the anatomy of the reproductive system. The new species K. albopunctatus sp. nov. is similar to K. ramosus in having a bright orange-red color pattern but differs significantly in the structure of appendages and reproductive system. Kaloplocamus japonicus can be easily distinguished from other Kaloplocamus species by its translucent, white-pink coloration and unique features of the female reproductive organ. Both species are supported as distinct species in all molecular analyses. The phylogenetic analyses propose a new estimate of the relationship between Kaloplocamus and Plocamopherus, and the evolution of bioluminescence within Triophinae is discussed. Our results also suggest cryptic biodiversity within the K. ramosus species complex.

Conformity of trans-Arctic Coryphellidae genera with the most recent revised taxonomy for the group was tested. Both morphological and molecular data were used to compare more than 200 specimens from various localities in the North Pacific, North Atlantic and Arctic Oceans. The molecular study included four markers (COI, 16S, H3 and 28S) and implemented Bayesian and maximum likelihood phylogenetic approaches. Species delimitation was investigated with automatic barcode gap discovery (ABGD), Poisson tree processes (PTP) and generalized mixed Yule coalescent (GMYC) methods. The inter- and intraspecific divergence were studied using the TCS-based haplotype analysis and calculation of p-distances. Morphological analysis included standard anatomical techniques and scanning electron microscopy. Based on integrative analysis, we conclude that Coryphella verrucosa and C. pseudoverrucosa represent a single species (C. verrucosa), which is highly heterogeneous at morphological and molecular levels. For Microchlamylla gracilis and M. amabilis, their synonymization under the name M. gracilis is suggested, but increased sample size is needed for further clarification of their identities. Finally, the amphiboreal species Borealea nobilis and the North Pacific B. sanamyanae are distinct species displaying secondary sympatry in the north-west Pacific. Based on these results, we propose a new classification system for the family Coryphellidae uniting all currently considered genera in the single genus Coryphella, and suggest a new approach for further revision of Flabellinidae s.l. taxa.

Berghia rissodominguezi Muniain & Ortea, 1999 (Figures 6A, 6B, 7A) Material examined. Argentina, Río Negro, Caleta Los Hornitos, two specimens, intertidal, 23/02/2017 (ESCM-Ma-37). Description. Length up to 30 mm. Body elongated, translucent white color, with oblique bright orange lines at the base of the cerata and rhinophores. Translucent cerata with white tip and a small, terminal cnidosac. Oral tentacles long, distal portion yellow. Rhinophores yellow (Figure 6B). Radular formula 25 x 0.1.0 (ESCM-Ma-37). Rachidian teeth bicurved with up to 45 denticles on each side of the central triangular cusp (Figure 7A). Geographic distribution and depth range. Western Atlantic, from Florida to San Jose gulf, Argentina (Muniain & Ortea 1999, Padula et al. 2011). Found in soft and intertidal bottoms under rocks up to 15 m depth. Biology. The white egg mass is a cylindrical belt with up to four complete turns in a spiral. The egg mass is deposited on rocks (Figure 6B). Remarks. This species was described and illustrated by Muniain & Ortea (1999). It is a warm water species that extends its distribution to the Magallanes Province. This is the first record of the B. rissodominguezi in the San Matías gulf., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 466-467, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Muniain, C. & Ortea, J. (1999) First records of the genus Berghia Trinchese, 1877 (Opisthobranchia: Aeolidiidae) from Argentina, with description of a new species. Avicennia, 10 / 11, 143 - 150.","Padula, V., Bahia, J., Vargas, C. & Lindner, A. (2011) Mollusca, Nudibranchia: New records and southward range extensions in Santa Catarina, southern Brazil. Check List, 7 (6), 806 - 808. https: // doi. org / 10.15560 / 11037"]}

Aeolidia cf. campbellii (Cunningham, 1871) (Figures 4G, 4F, 5G) Material examined. Argentina, Río Negro: Las Grutas, two specimens, (MLP-Ma 14575); Bahía San Antonio, one specimen, intertidal, March 2014. Description. Length up to 40 mm, body broad and relatively low, tapering to the posterior end of the foot. Gray, reddish, or transparent violet with white and gray spots. 20 rows of flattened cerata wider at the base. Each row contains up to 25 cerata that decrease in size towards the posterior end of the foot. Rhinophores conical, blunt and smooth. Oral tentacles translucent, longer than rhinophores and with white tips (Figure 4G, 4F). Radular formula 19 x 0.1.0 (MLP-Ma 14575). The radular teeth are progressively smaller towards the posterior region of the radula and have up to 100 denticles each (Figure 5G). Geographic distribution and depth range. Until now recorded from South America, in the Malvinas (Faklands) Islands, and Chilean Patagonia to Valparaiso (Kienberger et al. 2016). Here extended to SMG. Found under rocks in the intertidal zone. Remarks. Based on molecular analysis Kienberger et al. (2016) considered specimens from the Chilean coast, previously attributed to Aeolidia papillosa, as a different species: Aeolidia campbellii (Cunningham 1871), originally described from Tierra del Fuego. Scarabino (1977) reported A. papillosa in the San Matías gulf, however, specimens from the western Atlantic coast were not included in the work of Kienberger et al. (2016), so morphoanatomical and molecular studies of are necessary to confirm their identification., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 463-466, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Kienberger, K., Carmona, L., Pola, M., Padula, V., Gosliner, T. M. & Cervera, J. L. (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), 481 - 506. https: // doi. org / 10.1111 / zoj. 12379","Scarabino, V. (1977) Moluscos del Golfo San Matias (provincia de Rio Negro, Republica Argentina). Inventario y claves para su identificacion. Comunicaciones de la Sociedad Malacologica del Uruguay, 4, 177 - 285."]}

Coryphella cf. verta Ev. Marcus, 1970 (Figures 6E, 7B) Material examined. Argentina, Río Negro: Plataforma, one specimen, 8 m, 21/02/2017 (ESCM-Ma-82); one specimen, 8 m, 01/02/2021 (ESCM-Ma-55); one specimen, 6 m, 17/08/2021 (ESCM-Ma-42). Description. Length up to 10 mm, body translucent white. A longitudinal dorsal thin white line runs from the oral tentacles to the posterior end of the foot. Conical rhinophores with fine wrinkles and white tip. Cerata cylindrical, bright orange-red with white tips. Cerata grouped in widely spaced clusters along the dorsum. The anterior group begins just behind the rhinophores and contains 8 to 11 cerata, arranged in rows with no more than four per row. There are four to six posterior groups on each side of the body with a decreasing number of cerata in each group, up to 11 in the former, as few as one in the posterior one. Oral tentacles thin, elongated and slightly flattened (Figure 6E). Radular formula is 21 x 1.1.1 (ESCM-Ma-82). Rachidian teeth with a narrow central cusp, slightly longer than the 5-8 denticles on each side. Lateral teeth with a broad base and 5-7 long denticles (Figure 7B). Geographic distribution and depth range. Recorded from Florida, Gulf of Mexico, Venezuela to southeastern Brazil (Millen & Hamann 2006). Found on rocky bottoms between four and 12 m depth. Here extended to San Matías gulf, Argentina. Biology. Found on hard bottoms associated with hydrozoans. Remarks. Coryphella verta is a tropical species distributed from Florida to Brazil (Millen & Hamman 2006) and is recorded here for the first time in the Argentine Sea. C. verta can be distinguished externally from Flabellina dushia because F. dushia has oral tentacles longer than rhinophores, shorter cerata grouped in arcs, a wider body and broad opaque white bands on the notum, sides, head, and distal parts of the rhinophores and oral tentacles. Internally the lateral teeth of F. dushia are more acute, the seminal receptacle is shorter and larger and has a conical penis with a pointed tip (Millen & Hamann 2006). Another tropical species is F. dana, which can be easily distinguished because it has annulate rhinophores (Millen & Hamann 2006). The identification here is provisional due to observed differences in the body color pattern., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 467, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Millen, S. & Hamann, J. C. (2006) A new nudibranch species, genus Flabellina (Opisthobranchia: Aeolidacea) from the Caribbean with redescriptions of F. verta (Marcus 1970), and F. dushia (Marcus & Marcus 1963). Proceedings of the California Academy of Sciences, 57 (30), 928 - 936."]}

Diaulula punctuoalata (d'Orbigny, 1837) (Figures 2C, 2D, 3D) Material examined. Argentina, Río Negro: Las Grutas, one specimen, 10m, 2007 (MLP-Ma 14655); Las Grutas, one specimen (MLP-Ma 13528); Caleta Los Hornitos, one specimen, intertidal, 02/2017; San Antonio Este, Punta Villarino, two specimens (MACN 13368); San Matías gulf, SAO campaign, station 117, one specimen (MLP-Ma 14177). Description. Length up to 100 mm, body color varies from whitish to yellowish, some specimens presenting two rows of dark spots that extend from the rhinophores to the gill. Dorsal mantle densely covered by thin caryophyllidia. Rhinophoral and branchial sheaths elevated, covered with caryophyllidia. Oral tentacles digitiform. 6-7 bipinnate gill leaves around the anal papilla (Figure 2C). Foot bilabiate; upper lip notched. The radular formula is 24 x 27.0.27. (MLP-Ma 14655). Lateral teeth simple and smooth (Figure 3D). Geographic distribution and depth range. Argentina, Buenos Aires (37°50´S; 56°11´W) (Carcelles 1950), Nuevo gulf, in the Magellanic Province (Atlantic Ocean), throughout the Pacific Magellanic Province, to Arica, northern Chile, in the Peruvian Province (Pacific Ocean) (Valdés & Muniain 2002). Common in the SMG on rocky bottoms between six and 25 m depth. Biology. The egg mass in a whitish wavy gelatinous ribbon about 70 mm long, with up to four complete spiral turns (Figure 2D). Remarks. Magellanic species occasionally found in the Peruvian Province. This species is clearly differentiated from Diaulula hispida by the absence of the crest note (Valdés & Muniain 2002)., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 460, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Carcelles, A. R. (1950) Catalogo de los moluscos marinos de la Patagonia. Anales del Museo Nahuel Huapi, 1950, 41 - 49.","Valdes, A. & Muniain, C. (2002) Revision and taxonomic reassessment of Magellanic species assigned to Anisodoris Bergh, 1898 (Nudibranchia: Doridoidea). Journal of Molluscan Studies, 68, 345 - 351. https: // doi. org / 10.1093 / mollus / 68.4.345"]}

Trapania rocheae Cetra & Roche, 2019 (Figures 4A, 5A, 5B) Material examined. Holotype: Argentina: Río Negro, Balneario Las Grutas, one specimen, 8m, 04/2017 (MLPMa 14834, 9mm). Additional material: Plataforma: one specimen, 12m, 03/2017; one specimen, 8 m, 01/02/2018. Description. Length up to 12 mm, body elongated and smooth, pale white with brownish spots extending over the entire mantle, including gills and rhinophores. A white notal crest runs along the posterior dorsal area, from the base of the gills to the end of the tail. It has two pairs of dorsal processes, one on the rhinophores and the other on the gills. Short and conical oral tentacles. Rhinophores foliated with 13 leaves. Tripinnate gills, arranged in a semicircle around the anus (Figure 4A). Radular formula 32 x 0.1.0.1.0. (MLP-Ma 14834). Lateral teeth are rectangular, with eight large denticles and between each of these there are three to six smaller denticles. External cusp of each lateral tooth more elongated and robust. Masticatory armature composed of plates. Masticatory edge slightly curved with denticles that decrease in size towards the interior of the plates (Figure 5A, 5B). Geographic distribution and depth range. Only known from Balneario Las Grutas (40°49.291'S; 065° 04.498'W), Río Negro, Argentina (Cetra & Roche 2019). Found on rocky bottoms between four and 12 m depth. Remarks. This species has been recently described (see Cetra & Roche 2019). The family Goniodorididae has two species in the Argentine Sea, Ancula fuegiensis Odhner, 1926 and Trapania rocheae. The first one lacks records in the Argentine Sea since its original description by Odhner (1926) from Ushuaia, being recorded in the Chilean Pacific by Fischer and Ortea (1996) and Schrödl (2003). Both species have a similar coloration pattern differing in the location of the spots that, in A. fuegiensis, are found only on the dorsum around the gills, while in T. rocheae the spots extend throughout the body including rhinophores, digitiform processes and gills. The radulae are also different, A. fuegiensis presents marginal teeth, which are absent in the genus Trapania., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 462, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Cetra, N. & Roche, A. (2019) Una nueva especie del genero Trapania (Nudibranchia: Goniodorididae) para el Mar Argentino. Acta Zoologica Lilloana, 62 (2), 1 - 10. https: // doi. org / 10.30550 / j. azl / 2019.63.2 / 1","Odhner, N. (1926) Die Opisthobranchien. In: Further Zoological Results of the Swedish Antarctic Expedition 1901 - 1903 under the direction of Dr. Otto Nordenskjold. Norstedt, Stockholm, pp. 1 - 100.","Fischer, M. A. & Ortea, J. (1996) New records of the genus Ancula Loven, 1846 (Nudibranchia: Goniodorididae) on the American Pacific coast. The Veliger, 39, 90 - 92.","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp."]}

Rostanga pulchra MacFarland, 1905 (Figures 2B, 3B, 3C) Material examined. Argentina, San Matías gulf, Caleta Los Hornitos, REDE 2009, one specimen, 110m (MLP-Ma 14654). Description. Length up to 25 mm, body oval, uniform reddish orange, densely covered with thin caryophyllidia. Rhinophores with approximately ten vertical lamellae and apical knob. Oral tentacles short and conical.Ten unipinnate gill leaves around the anal papilla (Figure 2B). Foot broad and bilabiate. Radular formula is 60 x 30.44.0.44.30. (MLP-Ma 14654). Innermost teeth small with up to eight pointed denticles on the inner edge. Lateral teeth increase in size, becoming longer and thinner. Marginal teeth very thin and elongated, divided in the form of fine threads in the upper third (Figures 3B, 3C). Geographic distribution and depth range. Recorded from Eastern Pacific and Southern South America. Alaska to Mexico (Schrödl 2003), central Chile (Schrödl 1996, 1997a, 1999) and Bahía Camarones, Argentina (Schrödl 2003). Found on rocky bottoms between four and 110 m depth. Remarks. Rostanga pulchra is clearly distinguishable from other nudibranchs in this study by the orange color of the mantle and the lamellate rhinophores. The material studied matches the original description of R. pulchra from Monterey Bay, California (MacFarland 1905). R. pulchra has a disjunct geographic distribution in America, with a great geographic gap between north and south. Marcus & Marcus (1969) extended its distribution from Chile to Argentina, with one specimen collected in Camarones bay (44°45’57´´S; 65°34’47´´W) at 102 m depth. Schrödl (1996, 1997a) confirmed the presence of R. pulchra in the southern hemisphere. R. pulchra is very similar to its congener Rostanga byga Marcus, 1958 recorded from northern Brazil to Patagonia. According to Muniain & Valdés (2000) and Schrödl & Grau (2006) the differences between R. pulchra and R. byga are the presence of white dots on the central notum in R. byga and the double number of lamellae in the rhinophores., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 458, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["MacFarland, F. M. (1905) A preliminary account of the Dorididae of Monterey Bay, California. Proceedings of the Biological Society of Washington, 18, 35 - 54.","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp.","Schrodl, M. (1996) Nudibranchia y Sacoglossa de Chile: Morfologia exterior ydistribution. Gayana Zoologica, 60, 17 - 62.","Schrodl, M. (1997 a) Range extensions of Magellanic nudibranchs (Opisthobranchia) intothe Peruvian faunal province. The Veliger, 40, 38 - 42.","Schrodl, M. (1999) Zoogeographic relationships of the Magellan Nudibranchia (Mollusca: Opisthobranchia) with particular reference to species from adjacent regions. Scientia Marina, 63 (1), 409 - 416. https: // doi. org / 10.3989 / scimar. 1999.63 s 1409","Marcus, Ev. & Marcus Er. (1969) Opistobranchian and lamellarian gastropods collected by the \" Vema \". American Museum Novitates, 2368, 1 - 31.","Muniain, C. & Valdes, A. (2000) Rostanga byga Er. Marcus, 1958 from Argentina: redescription and comparison to Rostanga pulchra MacFarland, 1905 (Mollusca: Nudibranchia: Doridina). Proceedings of the California Academy of Sciences, 52, 1 - 10.","Schrodl, M. & Grau, J. H. (2006) Nudibranchia from the remote southern Chilean Guamblin and Ipun islands (Chonos Archipelago, 44 - 45 S), with re-description of Rostanga pulchra MacFarland, 1905. Revista Chilena de Historia Natural, 79 (1), 3 - 12. https: // doi. org / 10.4067 / S 0716 - 078 X 2006000100001"]}

Gargamella immaculata Bergh, 1894 (Figures 2E, 2F) Material examined. Argentina, San Matías gulf, Puerto Lobos, two specimens, (MLP-Ma 14578); Río Negro: Isla Mejillón, one specimen, intertidal, 03/11/2017; Plataforma, one specimen, 9m, 10/01/2018; one specimen, 6 m, 01/20/18. Description. Length up to 30 mm. Body oval, white, yellowish to orange. Dorsum densely covered with caryophyllidia of different sizes, the tips of some white in color. Rhinophores foliated. 8 to 12 bipinnate or tripinnate gill leaves around the anal papilla (Figure 2E). The radular formula is 60 x 81.0.81, lateral teeth very dense near the middle of the radula and are smaller towards the external ones. Geographic distribution and depth range. North of Argentina (37°50´S, 56°11´W), Argentinean and Chilean Patagonia to Bahia Coliumo, central Chile (Schrödl 1997b, 2003). Found in soft and intertidal bottoms under rocks up to 15 m depth. Biology. Egg mass is a gelatinous ribbon yellowish about 20 mm long with up to four complete spiral turns, and deposited on rocks (Figure 2F). Remarks. The genus Gargamella Bergh, 1894 currently contains seven species. Only one of them is listed for South America, Gargamella inmaculata, since the Chilean species Gargamella latior Odhner, 1926 was included as a junior synonym of G. inmaculata (Schrödl 2003). This species differs from Gargamella gravastella Valdés & Gosliner, 1999 and Gargamella bovina Garovoy, Valdés & Gosliner, 1999, from South Africa, by the absence of dark grey mantle color and dense carophyllidia present in G. garvastella. It differs from G. bovina by the absence of the white notum colored with dark brown spots and short caryophyllidia., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 460-461, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Schrodl, M. (1997 b) On the Magellanic nudibranch Gargamella immaculata Bergh, 1894, and its synonymy to G. latior Odhner, 1926. Spixiana, 20, 81 - 92.","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp.","Odhner, N. (1926) Die Opisthobranchien. In: Further Zoological Results of the Swedish Antarctic Expedition 1901 - 1903 under the direction of Dr. Otto Nordenskjold. Norstedt, Stockholm, pp. 1 - 100."]}

The San Matías gulf is a marine biodiversity hotspot located in the Argentinean Patagonia. Although there are several studies on gastropod mollusks from Argentina, nudibranchs remain less studied. In this faunistic assessment 15 species of nudibranchs were found in San Matías gulf, including two new family records for Argentina: Coryphellidae and Dotidae. Although Magellanic nudibranchs are clearly predominant in this area, warm water species such as Doto uva and Coryphella cf. verta are recorded and the known geographic range of five species is extended. In addition, Rostanga pulchra and Diaulula hispida extend their range to the north of the Argentine Patagonia.

Tritonia odhneri Er. Marcus, 1959 (Figures 6F, 7E, 7F) Material examined. Argentina, San Matías gulf: Caleta Los Hornitos, two specimens, 65 m, BENTOS 2008; one specimen, REDE, 2004; ten specimens, REDE 2005, 133 m; nineteen specimens, REDE 2009; Plataforma, one specimen, 9 m, 03/11/2017 (ESCM-Ma-10); sixty specimens, San Matías gulf, (MACN 21311). Diagnosis. Length up to 90 mm, elongated body slightly angled laterally, pink or orange coloration. Dorsum covered with irregular spots. Oral tentacles bilobed with several finger-like processes on each lobe. Lamellar rhinophores with lobed folds; abundant dorsolateral gills of different sizes (Figure 6F). Radular formula is 58 x 64.1.1.1.1.64 (ESCM-Ma-10). Rachidian teeth broad and tricuspid, all cusps strong. First lateral teeth broad-based with a short cusp. Outer lateral teeth simple, without cusps (Figures 7E, 7F). Mandibles are large with denticles on the chewing edges. Geographic distribution and depth range. From northern Uruguay to Central Chile; San Jorge gulf, Argentina (García-Matucheski & Muniain 2011); Gulf of Ancud to Coliumo Bay (Schrödl 2003). Found on rocky bottoms between four and 25 m depth. Remarks. The morphological characteristics of the specimens examined in this work agree with the original description of Tritonia odhneri Marcus, 1959. The Chilean and Argentinean populations of warm-adapted T. odhneri may be geographically separated, but still appear morphologically identical (Schrödl 2009). Moles et al. (2021) consider that Tritonia odhneri is a single species distributed around Cape Horn, Chile or that it is undergoing a process of speciation. This species was previously confused with the similarly colored Marionia cucullata (Couthouy, 1852) found in northern Argentina and Brazil but molecular analysis confirmed that they are distinct (Moles et al. 2021). Although M. cucullata has been mentioned by Scarabino (1977) in the SMG, it has not been collected again and no material of the species belonging to the gulf has been found in the museum collections consulted. A comparative revision of South Atlantic Tritonia species is needed., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 469-470, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Marcus, Er. (1959) Lamellariacea und Opisthobranchia. Reports of the Lund University Chile Expedition 1948 - 49, No. 36. Lunds Universitets Arsskrift, Ny Foljd, 55, 1 - 133.","Garcia-Matucheski, S. M. & Muniain, C. (2011) Predation by the nudibranch Tritonia odhneri (Opisthobranchia: Tritoniidae) on octocorals from the South Atlantic Ocean. Marine Biodiversity, 41, 287 - 297. https: // doi. org / 10.1007 / s 12526 - 010 - 0063 - y","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp.","Schrodl, M. (2009) Opisthobranchia - Babosas de Mar. In: Haussermann, V. & Forsterra, G. (Eds). Fauna Marina Bentonica de la Patagonia Chilena. Nature in Focus, Santiago de Chile, pp. 505 - 542.","Moles, J., Berning, M. I., Hooker, Y., Padula, V., Wilson, N. G. & Schrodl, M. (2021) Due South: the evolutionary history of Sub- Antarctic and Antarctic Tritoniidae nudibranchs. Molecular Phylogenetics and Evolution, 162, 107209. https: // doi. org / 10.1016 / j. ympev. 2021.107209","Scarabino, V. (1977) Moluscos del Golfo San Matias (provincia de Rio Negro, Republica Argentina). Inventario y claves para su identificacion. Comunicaciones de la Sociedad Malacologica del Uruguay, 4, 177 - 285."]}

Geitodoris patagonica Odhner, 1926 (Figures 2G, 2H, 3E, 3F) Material examined. Argentina, Río Negro, Caleta Los Hornitos, four specimens, intertidal, December 2017 (one specimen deposited, MLP-Ma 14656). Description. Length up to 40 mm, body white, yellow to orange or dark gray, with dark spots on the dorsum. Dorsal mantle is densely covered with tubercles of two different sizes. Rhinophores lamellate, 12 to 16 lamellae. 6-8 bi or tripinnate gill leaves around the anal papilla (Figure 2G). Radular formula is 16 x 12.24.0.24.12 (MLP-Ma 14656). The lateral teeth with small curved and pointed cusp. Marginal teeth delicate, very close standing lamellae (Figure 3E, 3F). Geographic distribution and depth range. Argentina, San Matías gulf (Scarabino 1977), Northern Chubut, Patagonia to Malvinas (Falkland) Islands (Schrödl 2000, Muniain 2001). Found in hard and intertidal bottoms under rocks up to 25 m depth. Biology. The egg masses of this species are gelatinous yellowish color tape of about 20 mm in length, with up to three full spiral turns. The egg mass is deposited on rocks (Figure 2H). Remarks. Only one species of Geitodoris is recorded for the Magellanic Province: Geitodoris patagonica. Another species, Geitodoris pusae (Marcus 1955), known from warm waters from Florida along Brazil to the Argentine coast, is an amphiatlantic species recorded by Marcus and Marcus (1969) on the north coast of Argentina, off Bahía Blanca at 68 m depth. According to Muniain (2001), G. pusae differs from G. patagonica by the presence of a reticulum and spicules present in the external duct of the vestibular gland. Muniain (2001) discussed the doubtful record of G. patagonica made by Carcelles (1944) from the littoral zone of Puerto Quequén (38º34'S) as it did not fit the original description given by Odhner (1926), suggesting that it should be disregarded., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 461-462, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Odhner, N. (1926) Die Opisthobranchien. In: Further Zoological Results of the Swedish Antarctic Expedition 1901 - 1903 under the direction of Dr. Otto Nordenskjold. Norstedt, Stockholm, pp. 1 - 100.","Scarabino, V. (1977) Moluscos del Golfo San Matias (provincia de Rio Negro, Republica Argentina). Inventario y claves para su identificacion. Comunicaciones de la Sociedad Malacologica del Uruguay, 4, 177 - 285.","Muniain, C. (2001) Taxonomical and ecological aspects of the nudibranch Geitodoris patagonica Odhner, 1926 (Opisthobranchia, Doridina) from Argentina. Bollettino Malacologico, 37, 171 - 176.","Marcus, Er. (1955) Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo. Zoologia, 20, 89 - 200. https: // doi. org / 10.11606 / issn. 2526 - 3382. bffclzoologia. 1955.120213","Marcus, Ev. & Marcus Er. (1969) Opistobranchian and lamellarian gastropods collected by the \" Vema \". American Museum Novitates, 2368, 1 - 31.","Carcelles, A. (1944) Catalogo de los moluscos marinos de Puerto Quequen (Republica Argentina). Revista del Museo de La Plata, 3 (23), 233 - 309."]}

Berghia rissodominguezi Muniain & Ortea, 1999 (Figures 6A, 6B, 7A) Material examined. Argentina, Río Negro, Caleta Los Hornitos, two specimens, intertidal, 23/02/2017 (ESCM-Ma-37). Description. Length up to 30 mm. Body elongated, translucent white color, with oblique bright orange lines at the base of the cerata and rhinophores. Translucent cerata with white tip and a small, terminal cnidosac. Oral tentacles long, distal portion yellow. Rhinophores yellow (Figure 6B). Radular formula 25 x 0.1.0 (ESCM-Ma-37). Rachidian teeth bicurved with up to 45 denticles on each side of the central triangular cusp (Figure 7A). Geographic distribution and depth range. Western Atlantic, from Florida to San Jose gulf, Argentina (Muniain & Ortea 1999, Padula et al. 2011). Found in soft and intertidal bottoms under rocks up to 15 m depth. Biology. The white egg mass is a cylindrical belt with up to four complete turns in a spiral. The egg mass is deposited on rocks (Figure 6B). Remarks. This species was described and illustrated by Muniain & Ortea (1999). It is a warm water species that extends its distribution to the Magallanes Province. This is the first record of the B. rissodominguezi in the San Matías gulf.

Polycera marplatensis Franceschi, 1928 (Figures 4C, 4D, 5E, 5F) Material examined. Argentina, San Matías gulf: Plataforma, one specimen, 10 m, 21/01/2018; Isla, two specimens, 9 m, 30/03/2018. Description. Length up to 30 mm, body elongated, white with orange dorsal lines, laterally spotted with yellow. Nine unipinnate posterior dorsal gill leaves, surrounding the anal papilla. A pair of digitiform projections surrounding the gills. Rhinophores lamellate with 18-20 lamellae. 6-7 digitiform processes in the oral veil (Figure 4C). The radular formula is 14 x 4.2.0.2.4. (MLP-Ma 14661). Rachidian teeth absent. The lateral teeth are simple with a spur on the end, the first inner lateral teeth differ in shape from the second one, usually larger; marginal teeth reduced. Jaw with a winglike process (Figure 5E, 5F). Geographic distribution and depth range. Brazil, S„o Paulo (Rios 1975) and Argentina, Mar del Plata (Franceschi 1928), Puerto Quequén (Carcelles 1944) both in Buenos Aires Province; Punta Pardelas, Chubut, Argentina (as “Patagonia”) (42°36´S, 64° 16´W) (Muniain & Ortea 1998) and Camarones Bay (Schrödl 1996). Found on rocky bottoms between four and 25 m depth. Biology. The white eggs masses are a slightly wavy ribbon with a full spiral turn. The egg mass is deposited on algae (Figure 4D). Remarks. Franceschi (1928) described a Polycera marplatensis as a variation of the European species Polycera quadrilineata (M̧ller 1776). Muniain and Ortea (1998) reevaluated its taxonomic status considering Polycera marplatensis as a distinct species. These authors redescribed it with the reexamination of the type material from Mar del Plata and based on material collected in Nuevo gulf, Patagonia. In that redescription the authors argue that P. marplatensis differs from P. quadrilineata in the body coloration pattern, with the absence of black coloration on the mantle, the larger size of the mandibles and radula, and a greater number of digitiform projections on the mantle in P. marplatensis. Our observations of the specimens studied agree with the descriptions of the species P. marplatensis, recording it for the first time in the San Matías gulf., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 463, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Franceschi, G. J. (1928) Una nueva forma de nudibranquio de la Argentina (Polycera quadrilineata var. marplatensis, n. var.). Anales del Museo Nacional de Historia Natural de Buenos Aires, 34, 577 - 586.","Rios, E. C. (1975) Brazilian Marine Mollusks Iconography. Fundac \" o Universidade do Rio Grande. Centro de Ciencias do Mar, Museu Oceanografico, Rio Grande, 331 pp.","Carcelles, A. (1944) Catalogo de los moluscos marinos de Puerto Quequen (Republica Argentina). Revista del Museo de La Plata, 3 (23), 233 - 309.","Muniain, C. & Ortea, J. (1998) The taxonomic status and redescription of Polycera marplatensis Franceschi, 1928 (Nudibranchia: Polyceratidae) from Argentina. The Veliger, 41 (2), 142 - 147.","Schrodl, M. (1996) Nudibranchia y Sacoglossa de Chile: Morfologia exterior ydistribution. Gayana Zoologica, 60, 17 - 62."]}

Diaulula hispida (d'Orbigny, 1834) Material examined. Argentina, San Matías gulf, Caleta Los Hornitos, BENTOS 2008, two specimens, 65 m (ESCM-Ma-22). Description. Length up to 80 mm, body whitish with brown dots, or body completely brown to dark brown. Undulating dorsal ridge. Mantle covered by very thin caryophyllidia, forming a collar at the base of the rhinophores. 6-7 tripinnate gill leaves surround the anal papilla. Oral tentacles digitiform. Foot bilabiate, with upper lip notched. The radular formula is 24 x 32.0.32. The lateral teeth are simple and smooth. Geographic distribution and depth range. Argentina, from Comodoro Rivadavia, Chubut to central Chile in Valparaiso, Chile (Schrödl 1996, 1997a, 2000, 2003). Found at 65 m depth. Remarks. According to Valdés & Gosliner (2001) D. hispida is characterized by a notum ridge, common to the members of the genus Atagema Gray, J.E. 1850 (= Trippa see Marcus 1959). On the contrary, Marcus (1959) considered that this species fits better within the genus Diaulula, due to morphological differences in the mouth of Trippa species. Until more information is available, D. hispida should be maintained in Diaulula (Schrödl 2003)., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 460, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Schrodl, M. (1996) Nudibranchia y Sacoglossa de Chile: Morfologia exterior ydistribution. Gayana Zoologica, 60, 17 - 62.","Schrodl, M. (1997 a) Range extensions of Magellanic nudibranchs (Opisthobranchia) intothe Peruvian faunal province. The Veliger, 40, 38 - 42.","Valdes, A. & Gosliner, T. M. (2001) Systematics and phylogeny of the caryophyllidia-bearing dorids (Mollusca, Nudibranchia), with descriptions of a new genus and four new species from Indo-Pacific deep waters. Zoological Journal of the Linnean Society, 133 (2), 103 - 198. https: // doi. org / 10.1111 / j. 1096 - 3642.2001. tb 00689. x","Marcus, Er. (1959) Lamellariacea und Opisthobranchia. Reports of the Lund University Chile Expedition 1948 - 49, No. 36. Lunds Universitets Arsskrift, Ny Foljd, 55, 1 - 133.","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp."]}

Trapania rocheae Cetra & Roche, 2019 (Figures 4A, 5A, 5B) Material examined. Holotype: Argentina: Río Negro, Balneario Las Grutas, one specimen, 8m, 04/2017 (MLPMa 14834, 9mm). Additional material: Plataforma: one specimen, 12m, 03/2017; one specimen, 8 m, 01/02/2018. Description. Length up to 12 mm, body elongated and smooth, pale white with brownish spots extending over the entire mantle, including gills and rhinophores. A white notal crest runs along the posterior dorsal area, from the base of the gills to the end of the tail. It has two pairs of dorsal processes, one on the rhinophores and the other on the gills. Short and conical oral tentacles. Rhinophores foliated with 13 leaves. Tripinnate gills, arranged in a semicircle around the anus (Figure 4A). Radular formula 32 x 0.1.0.1.0. (MLP-Ma 14834). Lateral teeth are rectangular, with eight large denticles and between each of these there are three to six smaller denticles. External cusp of each lateral tooth more elongated and robust. Masticatory armature composed of plates. Masticatory edge slightly curved with denticles that decrease in size towards the interior of the plates (Figure 5A, 5B). Geographic distribution and depth range. Only known from Balneario Las Grutas (40°49.291'S; 065° 04.498'W), Río Negro, Argentina (Cetra & Roche 2019). Found on rocky bottoms between four and 12 m depth. Remarks. This species has been recently described (see Cetra & Roche 2019). The family Goniodorididae has two species in the Argentine Sea, Ancula fuegiensis Odhner, 1926 and Trapania rocheae. The first one lacks records in the Argentine Sea since its original description by Odhner (1926) from Ushuaia, being recorded in the Chilean Pacific by Fischer and Ortea (1996) and Schrödl (2003). Both species have a similar coloration pattern differing in the location of the spots that, in A. fuegiensis, are found only on the dorsum around the gills, while in T. rocheae the spots extend throughout the body including rhinophores, digitiform processes and gills. The radulae are also different, A. fuegiensis presents marginal teeth, which are absent in the genus Trapania.

Doto uva Er. Marcus, 1955 (Figures 6C, 6D, 7C, 7D) Material examined. Argentina, San Matías gulf: Plataforma, one specimen, 10 m, 21/02/2017 (ESCM-Ma-50); Plataforma, one specimen, 7 m, 03/2017; La Salvadora, three specimens, 11 m, 04/2017 (ESCM-Ma-46); Plataforma, three specimens (two specimens deposited MLP-Ma 14657) 10 m, 08/2017; Plataforma, four specimens, 5 m, 09/2017; Plataforma, three specimens, 8 m, 01/2018 (ESCM-Ma-18). Description. Length up to 8 mm, body elongate and smooth, translucent whitish with darker spots. Smooth rhinophores with blunt apices, surrounded by rhinophore sheaths. Up to six cerata are present on each side of the body, it has 3-5 rings of 3-6 semiglobular tubercles and a tubercle at the top. The tubercles decrease in size towards the peduncle (Figure 6C). Radular formula 75-90 x 0.1.0 (MLP-Ma 14657). The rachidial teeth are arched with up to five lateral denticles on each side of the central cusp (Figures 7C, 7D). Geographic distribution and depth range. Key Largo, Florida (Marcus & Marcus 1960); S„o Paulo, Brazil (Marcus 1957); Gulf of Ancud to Bay of Coliumo, Chile (Schrödl 2003), whereas Fischer & Cervera (2005) considerably extended its known geographical distribution to northern Chilean coast (see Fischer et al. 2006) and Peru (Uribe et al. 2013). Found on rocky bottoms between four and 12 m depth. Biology. Found on hard bottoms associated with hydrozoans. Difficult to find due to its small size. The lightcream colored egg masses are thin, undulating ribbon of about 5 mm length. It is filled with numerous tiny, oval, encapsulated eggs (Figure 6D). Remarks. The genus Doto has three species in the Argentinean Province, Doto caramella Marcus, Er. 1957, Doto pita Marcus, Er. 1955 and Doto uva Marcus, Er. 1955 (García & Bertsch 2009). This species was originally described for S„o Sebasti„o, Brazil (Marcus 1955), later Marcus (1959) recorded it in the Gulf of Ancud, extending its distribution to the southern coast of Chile. Schrödl et al. (2005) recorded it from the southern Chile in the Comau Fjord. Fischer & Cervera (2005), considerably extend the known distribution range to Tongoy and Tocopilla and the Chilean coast. Uribe et al. (2013) extend the distribution north to Ica, Bahia Independencia, Peru. In the present work, the know range of the species is expanded to the Argentine Sea. This wide southern South America distribution should be confirmed with further integrative studies., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 467-468, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Marcus, Er. (1955) Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo. Zoologia, 20, 89 - 200. https: // doi. org / 10.11606 / issn. 2526 - 3382. bffclzoologia. 1955.120213","Marcus, Ev. & Marcus, Er. (1960) Opistobranchs from American Atlantic Warm Waters. Bulletin of Marine Science, 10, 129 - 203.","Marcus, Er. (1957) On Opisthobranchia from Brazil (2). Zoological Journal of the Linnean Society, 43, 390 - 486. https: // doi. org / 10.1111 / j. 1096 - 3642.1957. tb 01559. x","Schrodl, M. (2003) Sea slugs of Southern South America. ConchBooks, Hackenheim, 165 pp.","Fischer, M. A. & Cervera, J. L. (2005) Checklist of the opisthobranchs (Mollusca: Gastropoda) from the Chilean coast deposited at the \" Coleccion de Flora y Fauna Profesor Patricio Sanchez Reyes, Pontificia Universidad Catolica de Chile \". Iberus, 23 (2), 165 - 181.","Fischer, M. A., van der Velde, G. & Roubos, E. W. (2006) Morphology, anatomy and histology of Doto uva Marcus, 1955 (Opisthobranchia: Nudibranchia) from the Chilean coast. Contributions to Zoology, 75, 145 - 159. https: // doi. org / 10.1163 / 18759866 - 0750304004","Uribe, R. A., Nakamura, K., Indacochea, A., Pacheco, A. S., Hooker, Y. & Schrodl, M. (2013) A review on the diversity and distribution of opisthobranch gastropods from Peru, with the addition of three new records. Spixiana, 36 (1), 43 - 60.","Garcia, F. J. & Bertsch, H. (2009) Diversity and distribution of the Gastropoda Opisthobranchia from the Atlantic Ocean: a global biogeographic approach. Scientia Marina, 73 (1), 153 - 160. https: // doi. org / 10.3989 / scimar. 2009.73 n 1153","Marcus, Er. (1959) Lamellariacea und Opisthobranchia. Reports of the Lund University Chile Expedition 1948 - 49, No. 36. Lunds Universitets Arsskrift, Ny Foljd, 55, 1 - 133.","Schrodl, M., Alarcon, M., Bedrinana, M. A., Bravo, L. R., Bustamante, C. M., Carvalho, R., Forsterra, R., Gallardo, G., Hausserman, C. V. & Salmen, A. (2005) Nudipleura (Gastropoda, Opisthobranchia) from the southern Chilean Comau Fjord, with redescription of Polycera priva Er. Marcus, 1959. Vita Malacologica, 3, 23 - 33."]}

Cadlina sparsa (Odhner, 1921) (Figures 4B, 5C, 5D) Material examined. Argentina, Río Negro: Las Grutas, four specimens (MLP-Ma 14576); El Fuerte, four specimens, intertidal, 03/2012 (one specimen deposited, MLP-Ma 14657); La Salvadora, one specimen, 11 m, 19/04/2017; One specimen, Plataforma, at 4 m, 10/2017; Isla Mejillón, two specimens, intertidal, 11/11/2017 (one specimen deposited, MLP-Ma 14658). Description. Length up to 30 mm, body white, with a row of orange spots on the edge of the mantle. Dorsum covered with scattered tubers of different sizes. 10 to 16 unipinnate or bipinnate gill leaves that surround the anal papilla. Rhinophores lamellate with 10-15 lamellae (Figure 4B). Radular formula 64 x 0.30.1.30.0. (MLP-Ma 14658). Rachidian teeth in form of claws with four cusps. The first lateral teeth have some denticles and a large median cusp. The outermost lateral teeth become increasingly longer with numerous denticles (Figure 5C, 5D). Geographic distribution and depth range. California, from San Diego (Behrens 1991) to Marin County (Jaeckle 1983), Chile, Juan Fernandez Islands (Odhner 1921) and Coliumo Bay (36° 32´S 72° 57´W) (Schrödl 1996), Argentina, Camarones Bay (44° 53´S 65° 39´´ W) (Schrödl 1996). Found in hard intertidal bottoms under rocks up to 18 m depth. Remarks. Cadlina laevigata (Odhner 1926) cited for Puerto Madryn and Cadlina berghi (Odhner 1926) for Delgado Cape, in northern Argentina, have been revised and considered as synonyms of C. sparsa (Schrödl 2000)., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on pages 462-463, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["Odhner, N. H. (1921) Mollusca from Juan Fernandez and Easter Island. In: Scottsberg, C. (Ed.), The Natural History of Juan Fernandez and Easter Island. Vol. III. Zoology. Almqvist & Wiksells Boktryckeri, Uppsala, pp. pp. 219 - 254.","Behrens, D. W. (1991) Pacific coast nudibranchs: a guide to the opisthobranchs, Alaska to Baja California. Sea Challengers, No. 594.36 B 44. [unknown pagination]","Jaeckle, W. B. (1983) Additions to the opisthobranch mollusk fauna of Marin County, California, with notes on the biology of certain species. The Veliger, 26 (2), 93 - 95.","Schrodl, M. (1996) Nudibranchia y Sacoglossa de Chile: Morfologia exterior ydistribution. Gayana Zoologica, 60, 17 - 62.","Odhner, N. (1926) Die Opisthobranchien. In: Further Zoological Results of the Swedish Antarctic Expedition 1901 - 1903 under the direction of Dr. Otto Nordenskjold. Norstedt, Stockholm, pp. 1 - 100."]}

Spurilla braziliana (MacFarland, 1909) (Figures 4E, 4F, 5H) Material examined. Argentina, Río Negro: Caleta Los Hornitos, 15 specimens, intertidal, 18/12/2016 (two specimens deposited, MLP-Ma 14659); four specimens, intertidal, 03/02/2017; San Antonio Este, one specimen, intertidal, 24/08/2016 (ESCM-Ma-79); Fuerte Argentino, two specimens, intertidal, 03/03/2011 (ESCM-Ma-81); El Buque, two specimens, intertidal, 15/06/2016 (ESCM-Ma-31); Bahía San Antonio, two specimens, intertidal, 28/05/2016 (ESCM-Ma-32); Plataforma; one specimen, 12 m, 03/2017 (ESCM-Ma-80); Chubut, Puerto Lobos, one specimen (MLP-Ma 13108). Description. Length up to 50 mm, body elongated, orange. Rhinophores lamellate, with a white apex. Cerata arranged in 6 to 9 arches, with the tips clearer and curved towards the dorsal area of the body. Digestive glands are visible in the cerata (Figure 4E). Radular formula 23 x 0.1.0. (MLP-Ma 14659). Teeth progressively smaller in the posterior region of the radula. Rachidian teeth bicurved, with between 55 to 70 denticles on each side of the central, small, and triangular cusp. (Figure 5H). Geographic distribution and depth range. Pacific and Atlantic Ocean. Hawaiian Islands, Costa Rica, Peru, Japan, China, and Australia. From Florida to Argentinean Patagonian (Carmona et al. 2014). San Matías gulf, Argentina (Ģller & Zelaya 2017). Found in soft and intertidal bottoms under rocks up to 10 m depth. Biology. The egg mass is a spiral white ribbon, about 30 mm in length. The egg masses are deposited on the rocks (Figure 4F). Remarks. This species has often been recorded under the name Spurilla neapolitana (Delle Chiaje 1841). Externally, the color and morphology are not sufficient to differentiate the two species. Based on genetic data, Carmona et al. (2014) identified the southwestern Atlantic species as S. braziliana., Published as part of Cetra, Nicolás & Roche, Andrea, 2023, Nudibranchia (Mollusca: Gastropoda) from San Matías Gulf, Northern Argentine Patagonia, pp. 455-473 in Zootaxa 5244 (5) on page 466, DOI: 10.11646/zootaxa.5244.5.3, http://zenodo.org/record/7663830, {"references":["MacFarland, F. M. (1909) The opisthobranchiate Mollusca of the Branner-Agassiz expedition to Brazil. Leland Stanford Junior University Publications, University Series, 2, 1 - 104. https: // doi. org / 10.5962 / bhl. title. 23300","Carmona, L., Lei, B. R., Pola, M., Gosliner, T. M., Valdes, A. & Cervera, J. L. (2014) Untangling the Spurilla neapolitana (Delle Chiaje, 1841) species complex: a review of the genus Spurilla Bergh, 1864 (Mollusca: Nudibranchia: Aeolidiidae). Zoological Journal of the Linnean Society, 170 (1), 132 - 154. https: // doi. org / 10.1111 / zoj 12098","G ¸ ller, M. & Zelaya, D. G. (2017) A hot-spot of biodiversity in Northern Patagonia, Argentina. Biodiversity and Conservation, 26 (14), 3329 - 3342. https: // doi. org / 10.1007 / s 10531 - 017 - 1408 - 7"]}

The nudibranch diversity of the western Indian Ocean is comparatively one of the least studied in the world. In this paper a sponge-like Discodoridae nudibranch Jorunna liviaesp. nov. is described. The description is based on integrative anatomy, including molecular analysis of two genes (the mitochondrial COI and the nuclear H3), dissections, electron microscopy (SEM) of buccal elements, micro tomography of the spicule’s arrangements and ecological observations. This study provides the first ever molecular data of Jorunna species from the western Indian Ocean, helping to fill the gap to further understand this apparent paraphyletic genus.

21. Cadlina rumia Er. Marcus, 1955 (Fig. 3F) Material examined: Praia de Pirambúzios (mangrove), Nísia Floresta, 13.III.2009, one specimen, 12 mm (body length), leg. V. Padula (MZSP 97064). Praia de Baia Formosa, Baia Formosa, 04.VII.2009, one specimen, 10 mm (body length), leg. F. Santos (GEEFAA 311). Description: Body flattened, elliptical, 13 mm in length. Coloration white with yellow glands at the border of the mantle. Rhinophores with white basis graduating to brown in the apical portion. Anteriorly with a projecting notum, which covers the foot; and a pair of retractile, lamellate rhinophores; eyes close to the proximal part of the rhinophores. Mid-dorsally with a ring of glands around the notum. Posteriorly, five flattened, retractile gill leaves around the anus. Geographic distribution: Eastern Atlantic: Ghana (Edmunds, 1981); Western Atlantic: Aruba, Bahamas, Belize, Bonaire, Cuba, CuraÇao, Dominican Republic, Grenada, Guadeloupe, Jamaica, Panama, Puerto Rico, St. Lucia, St. Martín / St. Maarten, Saint Vicent & the Grenadines, USA, Venezuela, Brazil (Alagoas,Bahia, Rio de Janeiro, Rio Grande do Norte – present study, São Paulo) (Marcus, 1955; Domínguez et al., 2006; Valdés et al., 2006; García et al., 2008; Ourives et al., 2011; Padula et al., 2012; Sales et al., 2013). Family Chromodorididae Bergh, 1891, Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 13, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Marcus, E. 1955. Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras, 207 (20): 89 - 261. https: // doi. org / 10.11606 / issn. 2526 - 3382. bffclzoologia. 1955.120213.","Edmunds, M. 1981. Opisthobranchiate Mollusca from Ghana: Chromodorididae. Zoological Journal of the Linnean Society, 72 (2): 175 - 201. https: // doi. org / 10.1111 / j. 1096 - 3642.1981. tb 01657. x.","Dominguez, M.; Garcia, F. J. & Troncoso, J. S. 2006. Some aspects of the family Chromodorididae (Opisthobranchia: Nudibranchia) from Brazil, with description of a new species. Scientia Marina, 70 (4): 621 - 634. https: // doi. org / 10.989 / scimar. 2006.70 n 4621.","Valdes, A.; Hamann, J.; Behrens, D. & DuPont, A. 2006. Caribbean Sea Slugs. A field guide to the Opisthobranch mollusks from the tropical Northwestern Atlantic. Gig Harbor, WA, Sea Challengers. 289 p.","Garcia, F. J.; Dominguez-Alvarez, M. & Troncoso, J. S. 2008. Opistobranquios de Brasil: descripcion y distribucion de Opistobranquios del litoral de Brasil y del Archipielago Fernando de Noronha, Vigo, Feito, 215 p.","Ourives, T. M.; Guerrazzi, M. C. & Simone, L. R. 2011. Gastropods from Camamu Bay, state of Bahia, Brazil. Check List, 7 (3): 328 - 336. https: // doi. org / 10.15560 / 7.3.328.","Padula, V.; Bahia, J.; Correia, M. D. & Sovierzoski, H. H. 2012. New records of opisthobranchs (Mollusca: Gastropoda) from Alagoas, Northeastern Brazil. Marine Biodiversity Records, 5: 1 - 11. https: // doi. org / 10.1017 / S 1755267212000346.","Sales, L.; Queiroz, V.; Padula, V.; Neves, E. & Johnsson, R. 2013. New records of nudibranchs (Mollusca: Gastropoda) from Bahia State, northeastern Brazil. Check List, 9 (3): 689 - 691. https: // doi. org / 10.15560 / 9.3.689."]}

Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana, Brandão, Simone Nunes (2022): Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil. Papéis Avulsos de Zoologia 62: 1-26, DOI: 10.11606/1807-0205/2022.62.063, URL: http://dx.doi.org/10.11606/1807-0205/2022.62.063

22. Felimida clenchi (Russel, 1935) (Fig. 3G) Material examined: Praia de Santa Rita, Extremoz, 09.III.2009, one specimen, 4 mm (body length), leg. V. Padula (MZSP 97081). Praia de Pirambúzios, Nísia Floresta, 06. V.2008, one specimen (lost), 10 mm (body length), phot. reg. N. Filho. Praia de Baia Formosa, Baia Formosa, 04.VII.2009, two specimens, 6-9 mm (body length), leg. G. Grimaldi (MZSP 97070), 05.VII.2009, six specimens, 4-15 mm (body length), leg. F. Santos (GEEFAA 263a, 329). 03.I.2011, one specimen, 10 mm (body length), leg. M. Delgado (GEEFAA 332). Description: Body oblong, up to 25 mm (body length) in length. Border of mantle reddish-orange-yellowish reticulated pattern. Anteriorly, elongated notum covering head and foot, dark-colored from midlength to posterior, where visceral mass is located. Rhinophores and gills white and purple. Retractile, lamellate rhinophores curved posteriorly. Anus surrounded by flattened, retractile, gill leaves. Posterior of body acute, orange to reddish dorsally, and white ventrally. Geographic distribution: Western Atlantic: Aruba, Bermuda, Bonaire, Costa Rica, Colombia, Cuba, CuraÇao, Guadeloupe, Jamaica, Panama, St. Vicent & the Grenadines, USA, Venezuela, Virgin Islands, Brazil (Ceará, Rio Grande do Norte, Alagoas, Bahia, Rio de Janeiro and São Paulo) (Marcus, 1957; Domínguez et al., 2006; García et al., 2008; Padula et al., 2012, Sales et al., 2013; GalvãoFilho et al., 2015; Padula et al., 2016)., Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 13, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Marcus, E. 1957. On opisthobranchia from Brazil (2). Zoological Journal of the Linnean Society, 43 (292): 390 - 486. https: // doi. org / 10.1111 / j. 1096 - 3642.1957. tb 01559. x.","Dominguez, M.; Garcia, F. J. & Troncoso, J. S. 2006. Some aspects of the family Chromodorididae (Opisthobranchia: Nudibranchia) from Brazil, with description of a new species. Scientia Marina, 70 (4): 621 - 634. https: // doi. org / 10.989 / scimar. 2006.70 n 4621.","Garcia, F. J.; Dominguez-Alvarez, M. & Troncoso, J. S. 2008. Opistobranquios de Brasil: descripcion y distribucion de Opistobranquios del litoral de Brasil y del Archipielago Fernando de Noronha, Vigo, Feito, 215 p.","Padula, V.; Bahia, J.; Correia, M. D. & Sovierzoski, H. H. 2012. New records of opisthobranchs (Mollusca: Gastropoda) from Alagoas, Northeastern Brazil. Marine Biodiversity Records, 5: 1 - 11. https: // doi. org / 10.1017 / S 1755267212000346.","Sales, L.; Queiroz, V.; Padula, V.; Neves, E. & Johnsson, R. 2013. New records of nudibranchs (Mollusca: Gastropoda) from Bahia State, northeastern Brazil. Check List, 9 (3): 689 - 691. https: // doi. org / 10.15560 / 9.3.689.","Padula, V.; Bahia, J.; Stoger, I.; Camacho-Garcia, Y.; Malaquias, M. A. E.; Cervera, J. L. & Schrodl, M. 2016. A test of color-based taxonomy in nudibranchs: molecular phylogeny and species delimitation of the Felimida clenchi (Mollusca: Chromodorididae) species complex. Molecular Phylogenetics and Evolution, 103: 215 - 229. https: // doi. org / 10.1016 / j. ympev. 2016.07.019."]}

30. Taringa telopia Er. Marcus, 1955 (Fig. 3O) Material examined: Praia de Areia Preta, Natal, 22. VI.2009, one specimen, 18 mm (body length), leg. F. Santos (MZSP 97071). Description: Body flat, elliptical, yellowish notum with dark brown, small spots from the medial portion of the body to the margins of the notum. Rhinophores with whitish bases and light brown lamellae.Gills light brown. Head with eyes covered by edge of the mantle, bilobated labium. Lamellate retractile rhinophores.Body with small tubercles medially. Six folliculated, retractile, tripinnated gill leaves around the anus. Foot narrower than notum. Geographic distribution: Western Atlantic: Colombia, Brazil (Alagoas, Pernambuco, Rio de Janeiro, Rio Grande do Norte – present study, São Paulo) (Bergh, 1894; Er. Marcus, 1955; Ev. Marcus & Er. Marcus, 1967a; Valdés & Gosliner, 2001; García et al., 2008; Padula et al., 2012; Alvim & Pimenta, 2013). Family Dorididae Rafinesque, 1815, Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on pages 15-16, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Marcus, E. 1955. Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras, 207 (20): 89 - 261. https: // doi. org / 10.11606 / issn. 2526 - 3382. bffclzoologia. 1955.120213.","Bergh, L. S. R. 1894. Die Opisthobranchien. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish Commission steamer \" Albatross \", during 1891, Lieut. Commander Z. L. Tanner, U. S. N., commanding. Bulletin of the Museum of Comparative Zoology, 25: 125 - 233. https: // www. biodiversitylibrary. org / item / 48546 # page / 5 / mode / 1 up.","Marcus, E. D. B. R. & Marcus, E. 1967 a. American opisthobranch mollusks Part I, Tropical American opisthobranchs. Studies in Tropical Oceanography, 6 (1 - 2): 1 - 256.","Valdes, A. & Gosliner, T. M. 2001. Systematics and Phylogeny of the caryophyllidia-bearing dorids (Mollusca, Nudibranchia), with descriptions of a new genus and four new species from Indo-Pacific deep waters. Zoological Journal of Linnean Society, 133 (2): 103 - 198. https: // doi. org / 10.1111 / j. 1096 - 3642.2001. tb 00689. x.","Garcia, F. J.; Dominguez-Alvarez, M. & Troncoso, J. S. 2008. Opistobranquios de Brasil: descripcion y distribucion de Opistobranquios del litoral de Brasil y del Archipielago Fernando de Noronha, Vigo, Feito, 215 p.","Padula, V.; Bahia, J.; Correia, M. D. & Sovierzoski, H. H. 2012. New records of opisthobranchs (Mollusca: Gastropoda) from Alagoas, Northeastern Brazil. Marine Biodiversity Records, 5: 1 - 11. https: // doi. org / 10.1017 / S 1755267212000346.","Alvim, J. & Pimenta, A. D. 2013. Taxonomic review of the family Discodorididae (Mollusca: Gastropoda: Nudibranchia) from Brazil, with descriptions of two new species. Zootaxa, 3745 (2): 152 - 198. https: // doi. org / 10.11646 / zootaxa. 3745.2.2."]}

29. Taringa iemanja Alvim & Pimenta, 2013 (Fig. 3N) Material examined: Praia de Baia Formosa, Baia Formosa, 05.VII.2009, two specimens, 24-25 mm (body length), leg. F. Santos (MZSP 97062; GEEFAA 337). Geographic distribution: Western Atlantic: Brazil (Rio de Janeiro, Rio Grande do Norte – present study) (Alvim & Pimenta, 2013). Description: Body flat,elliptical,dorsally dark brown with oval-shaped, dark brown spot. Ventrally, light brown notum and foot with oval-shaped,small spots. Rhinophores and gills light brown with small, irregular-shaped, dark brown spots. Bilobated labium. Lamellated, retractile rhinophores within a basal projection. Body with smallsized, rounded tubercles medially. Seven folliculated, retractile, tripinnated gill leaves around the anus. Foot smooth, narrower than and covered by the notum. Remarks: First record from northeastern Brazil., Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 15, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Alvim, J. & Pimenta, A. D. 2013. Taxonomic review of the family Discodorididae (Mollusca: Gastropoda: Nudibranchia) from Brazil, with descriptions of two new species. Zootaxa, 3745 (2): 152 - 198. https: // doi. org / 10.11646 / zootaxa. 3745.2.2."]}

29. Taringa iemanja Alvim & Pimenta, 2013 (Fig. 3N) Material examined: Praia de Baia Formosa, Baia Formosa, 05.VII.2009, two specimens, 24-25 mm (body length), leg. F. Santos (MZSP 97062; GEEFAA 337). Geographic distribution: Western Atlantic: Brazil (Rio de Janeiro, Rio Grande do Norte – present study) (Alvim & Pimenta, 2013). Description: Body flat,elliptical,dorsally dark brown with oval-shaped, dark brown spot. Ventrally, light brown notum and foot with oval-shaped,small spots. Rhinophores and gills light brown with small, irregular-shaped, dark brown spots. Bilobated labium. Lamellated, retractile rhinophores within a basal projection. Body with smallsized, rounded tubercles medially. Seven folliculated, retractile, tripinnated gill leaves around the anus. Foot smooth, narrower than and covered by the notum. Remarks: First record from northeastern Brazil.

34. Doto divae Marcus & Marcus, 1960 (Fig. 4D) Material examined: Parrachos de Rio do Fogo, Rio do Fogo, 25. VI.2010, one specimen, 2 mm (body length), leg. M. Delgado (GEEFAA 285). Description: Body cylindric and elongated, whitish with dark green spots irregularly distributed over the body. Rhinophores whitish with dark green spots. Conspicuous, well-developed head with small eyes close to the basis of the rhinophores, elongated, retractile rhinophores with robust basal projection. Anal and genital pores present mid-laterally and below the first cerata, and between second and third ceratae, respectively.Two longitudinal rows of ceratae parallel to each other, with narrow basis and with globular tubercles. Second cerata with well-developed and vertically elongated, small tubercle. Foliaceous gills between the tubercles, irregularly distributed. Posteriorly, smooth foot elongated forming a prominent, translucent tail. Geographic distribution: Western Atlantic: Bonaire, Mexico, Puerto Rico, St. Lucia, USA, Brazil (Bahia, Ceará, Rio Grande do Norte – present study) (Marcus & Marcus, 1960; Valdés et al., 2006; Rios, 2009). Superfamily Aeolidioidea Gray, 1827, Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 16, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Marcus, E. D. B. R. & Marcus, E. 1960. Opisthobranchs from American Atlantic warm waters. Bulletin of Marine Science of the Gulf and Caribbean, 10 (2): 129 - 203.","Valdes, A.; Hamann, J.; Behrens, D. & DuPont, A. 2006. Caribbean Sea Slugs. A field guide to the Opisthobranch mollusks from the tropical Northwestern Atlantic. Gig Harbor, WA, Sea Challengers. 289 p.","Rios, E. C. 2009. Compendium of Brazilian Sea Shells. Rio Grande, Evangraf. 668 p."]}

25. Diaulula greeleyi (MacFarland, 1909) (Fig. 3J) Material examined: Praia de Rio do Fogo, Rio do Fogo, 28.I.2010, one specimen, 7 mm (body length), leg. M. Delgado (GEEFAA 302). Praia de Santa Rita, Extremoz, 11.X.2008, two specimens, 5-9 mm (body length), leg. M. Delgado (MZSP 97046), 26.III.2009, two specimens, 7-10 mm (body length), leg. S. Delgado, 28.III.2009, two specimens, 6-13 mm (body length), leg. M. Delgado (GEEFAA lot 316), 26.IV.2009, five specimens, 3-20 mm (body length), leg. F. Santos (GEEFAA 259 a), 24.VI.2009, four specimens, 3-14 mm (body length), leg. M. Delgado (GEEFAA lot 316), 09.VII.2009, three specimens, 7-11 mm (body length),leg. M.Delgado (GEEFAA lot 316), 05.I.2011, one specimen, 11 mm (body length), leg. M. Delgado (GEEFAA 290). Praia de Pirambúzios, Nísia Floresta, 11.VII.2009, three specimens, 5-10 mm (body length), leg. F. Santos (GEEFAA 317), 08.XII.2013, one specimen, 6 mm (body length), leg. M. Delgado (GEEFAA 348), 15.XII.2013, three specimens, 5-10 mm (body length), leg. S. Delgado (GEEFAA 283), 18.I.2014, two specimens, 8-10 mm (body length), leg. M. Delgado (GEEFAA 341). Praia de Baia Formosa, Baia Formosa, 04.VII.2009, 15 specimens, 4-10 mm (body length), leg. M. Delgado (GEEFAA 288 – 13 expl.; MZSP 97045 – 2 expl.), 05.VII.2009, three specimens, 3-4 mm (body length), leg. F. Santos (GEEFAA 301). Praia de Pitangui, Extremoz, 11.I.2019, four specimens, 4-14 mm (body length), leg. M. Delgado (GEEFAA 1313). Description: Body flat and elliptical, yellow-orange coloration with small, subcircular, brown-black patches, and brown-black longitudinal,subparallel lines from the base of rhinophores to the gills. Rhinophores dark brown in the laminar portion and yellow-orange in the basis. Gills brown-black with translucent orange on the margins and center portion. Conical oral tentacles, lamellate retractile rhinophores. Medial portion of notum with abundant tubercles. Mantle margins with glands. Unipinnate and retractile gills. Muscular foot narrower than the mantle, anteriorly bifurcated and short in the posterior portion. Geographic distribution: Eastern Pacific Ocean: Mexico, Costa Rica (Camacho-García & Valdés, 2003; Valdés et al., 2006; Dayrat, 2010); Western Atlantic: Bahamas, Costa Rica, USA, Brazil (Alagoas, Ceará, Pernambuco, Rio de Janeiro, Rio Grande do Norte – present study, São Paulo) (MacFarland,1909:as P.greeleyi; Marcus, 1955; Ev.Marcus & Er. Marcus, 1967a; Rios, 1994; García et al., 2008; Padula et al., 2012; Alvim & Pimenta,2013; Camacho-García et al., 2014; Galvão-Filho et al., 2015)., Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 14, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Macfarland, F. M. 1909. The Opisthobranchiate Mollusca of the Branner- Agassiz Expedition to Brazil. California, Stanford University. 104 p. (Leland Stanford Junior University Publications, University Series, 2) https: // doi. org / 10.5962 / bhl. title. 23300.","Camacho-Garcia, Y. & Valdes, A. 2003. Caryophyllidia-bearing dorid nudibranchs (Mollusca, Nudibranchia, Doridacea) from Costa Rica. Proceedings of the California Academy of Sciences, 54: 65 - 79.","Valdes, A.; Hamann, J.; Behrens, D. & DuPont, A. 2006. Caribbean Sea Slugs. A field guide to the Opisthobranch mollusks from the tropical Northwestern Atlantic. Gig Harbor, WA, Sea Challengers. 289 p.","Dayrat, B. 2010. A monographic revision of basal Discodorid sea slugs (Mollusca: Gastropoda: Nudibranchia: Doridina). Proceedings of the California Academy of Sciences, 61 (1): 1 - 403.","Marcus, E. 1955. Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras, 207 (20): 89 - 261. https: // doi. org / 10.11606 / issn. 2526 - 3382. bffclzoologia. 1955.120213.","Marcus, E. D. B. R. & Marcus, E. 1967 a. American opisthobranch mollusks Part I, Tropical American opisthobranchs. Studies in Tropical Oceanography, 6 (1 - 2): 1 - 256.","Rios, E. C. 1994. Seashells of Brazil, Editora da Fundacao Universidade Federal do Rio Grande, Rio Grande, Brazil, 368 pp.","Garcia, F. J.; Dominguez-Alvarez, M. & Troncoso, J. S. 2008. Opistobranquios de Brasil: descripcion y distribucion de Opistobranquios del litoral de Brasil y del Archipielago Fernando de Noronha, Vigo, Feito, 215 p.","Padula, V.; Bahia, J.; Correia, M. D. & Sovierzoski, H. H. 2012. New records of opisthobranchs (Mollusca: Gastropoda) from Alagoas, Northeastern Brazil. Marine Biodiversity Records, 5: 1 - 11. https: // doi. org / 10.1017 / S 1755267212000346.","Alvim, J. & Pimenta, A. D. 2013. Taxonomic review of the family Discodorididae (Mollusca: Gastropoda: Nudibranchia) from Brazil, with descriptions of two new species. Zootaxa, 3745 (2): 152 - 198. https: // doi. org / 10.11646 / zootaxa. 3745.2.2.","Camacho-Garcia, Y. E.; Pola, M.; Carmona, L.; Padula, V.; Villani, G. & Cervera, J. L. 2014. Diversity and distribution of the heterobranch sea slug fauna on the Caribbean of Costa Rica. Cahiers de Biologie Marine, 55 (1): 109 - 127. https: // doi. org / 10.21411 / CBM. A. 8 B 3 AE 5 D 2.","Galvao-Filho, H.; Araujo, A.; Silva, F.; Azevedo, V.; Meirelles, C. A. O. & Matthews-Cascon, H. 2015. Sea slugs (Gastropoda: Heterobranchia) from a poorly known area in North-east Brazil. Filling gaps in Atlantic distributions. Marine Biodiversity Records, 8 (e 115). https: // doi. org / 10.1017 / S 1755267215000494."]}

40. Glaucus atlanticus Forster, 1777 (Fig. 4J) Material examined: Praia de Santa Rita, Extremoz, 15.VIII.2016, one specimen, 35 mm (body length), phot. reg. P. Rocha. Praia de Ponta Negra, Natal, 02.XI.2020, two specimens, 30-32 mm (body length), phot. reg. Í. Sérgio. Description: Elongate and flat body, up to 35 mm (body length) in length. Color silver, rhinophores, cerata, oral tentacles, back dorsum and tail predominantly dark blue and white on ventral side. Head small with a ventral, small oral tentacles and very short rhinophores on the dorsal side. There are three pairs of clusters of cerata arranged on the lateral side of the body. Clusters of cerata mobile, allowing for retraction and distension movements similar to swimming. Geographic distribution: Widespread in the tropical and subtropical oceans (Churchill et al., 2014). Mediterranean Sea:Indian Ocean: Andaman & Nicobar Islands, Australia; Cook Island, India, Lord Howe, Madagascar, Norfolk Islands, Seychelles, South Africa, Sri Lanka, Vietnam, Yemen (Srinivasulu et al., 2012); Eastern Pacific: Costa Rica, El Salvador, Galapagos, Hawaii, Mexico, USA (Srinivasulu et al., 2012); Western Pacific: Japan, New Zealand (Srinivasulu et al., 2012); Eastern Atlantic: Azores, Canaries, Madeira, South Africa (Srinivasulu et al., 2012); Western Atlantic: Aruba, Argentina, Barbados, Bermuda, Jamaica, Mexico, Uruguay, USA, and Brazil (Alagoas, Bahia, Rio Grande do Norte – present study, São Paulo and Rio Grande do Sul) (Valdés et al., 2006; Segovia & López, 2015; Pinotti et al., 2019). Remarks: First record from Rio Grande do Norte. The specimens from both localities sampled were found at the seaside stranded close to Velella velella (Linnaeus, 1758)., Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on pages 19-20, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Churchill, C. K. C.; Valdes, A. & Foighil, D. O. 2014. Molecular and morphological systematics of neustonic nudibranchs (Mollusca: Gastropoda: Glaucidae: Glaucus), with descriptions of three new cryptic species. Invertebrate Systematics, 28: 174 - 195. https: // doi. org / 10.1071 / IS 13038.","Srinivasulu, B.; Srinivasulu, C. & Kumar, G. C. 2012. First record of the blue sea slug (Glaucus atlanticus) from Andhra Pradesh - India. Taprobanica: The Journal of Asian Biodiversity, 4 (1): 52 - 53. https: // doi. org / 10.4038 / tapro. v 4 i 1.4386.","Valdes, A.; Hamann, J.; Behrens, D. & DuPont, A. 2006. Caribbean Sea Slugs. A field guide to the Opisthobranch mollusks from the tropical Northwestern Atlantic. Gig Harbor, WA, Sea Challengers. 289 p.","Segovia, J. & Lopez, G. 2015. Registro de Glaucus atlanticus en la costa de El Salvador, Pacifico de Centroamerica. Revista Mexicana de Biodiversidad, (86): 1089 - 1090. https: // doi. org / 10.1016 / j. rmb. 2015.05.015.","Pinotti, R. M.; Bom, F. C. & Muxagata, E. 2019. On the occurrence and ecology of Glaucus atlanticus Forster, 1777 (Mollusca: Nudibranchia) along the Southwestern Atlantic coast. Anais da Academia Brasileira de Ciencias, (91): 1 - 6. https: // doi. org / 10.1590 / 0001 - 3765201920180154."]}

33. Doto chica Marcus & Marcus, 1960 (Fig. 4C) Material examined: Praia de Baia Formosa, Baia Formosa, 03.I.2011, three specimens, 5-6 mm (body length), leg. M. Delgado (GEEFAA 315). Description: Body flat and elongated, whitish with light brown spots over the entire body and also white, spots. Ceratae creamish, with light pink tubercles and blue spots. Rhinophores dark brown with minute, whitish spots. Head well developed, with small eyes, elongated oral tentacles, one pair of retractile rhinophores with basal projection. Medially, with anal and genital pores elevated. Two rows of longitudinal ceratae with globose tubercles. Posteriorly, smooth foot elongated forming a prominent tail. Geographic distribution: Western Atlantic: Bahamas, Costa Rica, Cuba, CuraÇao, Mexico, Panama, Puerto Rico, USA, Venezuela, Brazil (Ceará, Rio Grande do Norte – present study, São Paulo) (Ev.Marcus & Er. Marcus,1960; Valdés et al., 2006; García et al., 2008; Galvão-Filho et al., 2015)., Published as part of Delgado, Marlon, Freire, Fúlvio Aurélio de Morais, Meirelles, Carlos Augusto Oliveira de, Padula, Vinicius, Bahia, Juliana & Brandão, Simone Nunes, 2022, Sea slugs (Gastropoda: Heterobranchia) from Rio Grande do Norte, Northeastern Brazil, pp. 1-26 in Papéis Avulsos de Zoologia 62 on page 16, DOI: 10.11606/1807-0205/2022.62.063, http://zenodo.org/record/7617672, {"references":["Marcus, E. D. B. R. & Marcus, E. 1960. Opisthobranchs from American Atlantic warm waters. Bulletin of Marine Science of the Gulf and Caribbean, 10 (2): 129 - 203.","Valdes, A.; Hamann, J.; Behrens, D. & DuPont, A. 2006. Caribbean Sea Slugs. A field guide to the Opisthobranch mollusks from the tropical Northwestern Atlantic. Gig Harbor, WA, Sea Challengers. 289 p.","Garcia, F. J.; Dominguez-Alvarez, M. & Troncoso, J. S. 2008. Opistobranquios de Brasil: descripcion y distribucion de Opistobranquios del litoral de Brasil y del Archipielago Fernando de Noronha, Vigo, Feito, 215 p.","Galvao-Filho, H.; Araujo, A.; Silva, F.; Azevedo, V.; Meirelles, C. A. O. & Matthews-Cascon, H. 2015. Sea slugs (Gastropoda: Heterobranchia) from a poorly known area in North-east Brazil. Filling gaps in Atlantic distributions. Marine Biodiversity Records, 8 (e 115). https: // doi. org / 10.1017 / S 1755267215000494."]}

In this study five new records and two probably undescribed species of heterobranch sea slugs placed in four genera, three families, and two orders are reported from Kuwait, northwestern Arabian / Persian Gulf with details and photographs. The present study increases the heterobranch diversity in Kuwaiti waters from 35 to 40 species. The range of habitats in Kuwait provides a vital opportunity for further investigation to understand the actual faunal diversity.

TRAPANIA TIGGER SP. NOV. (FIGS 2H, 4H, 11) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 3295F8AD-8D24-4204-A98E-8F6910B14367. Trapania sp. 6 Nakano, 2004: 95, top photograph. Trapania sp. 7 Gosliner et al., 2015: 143, bottom-left photograph. Trapania sp. 8 Gosliner et al., 2018: 61, middle-right photograph. Type material: Holotype: NMP 041328, originally CASIZ 180412, one specimen, dissected and sequenced, Mabini, Batangas Province, Luzon Island, Philippines, November 2008, collected by Peri Paleracio. Type locality: Mabini, Batangas Province, Luzon Island, the Philippines. Geographical distribution: Known only from Mabini, Luzon, the Philippines. Etymology: Trapania tigger is named for its resemblance to Tigger, a fictional character illustrated by E. H. Shepard from the Winnie-the-Pooh series of stories (Milne, 1928). It is a noun in apposition. External morphology: Living animal (Fig. 2H) is 12 mm. Entire body is mottled pale tan with darker brown patches and a few small opaque white spots over the entire surface of the body. Posterior portion of the body has alternating bands of light and dark pigment, while the middle and anterior of the body is characterized by discrete light and dark patches of pigment. Rhinophores are thick, club-shaped with about eight lamellae and a pointed tip. The small gill has a central, branching stalk flanked by two smaller branches. The extra-branchial and extra-rhinophoral appendages are thick, stocky and have a slight bulge near the base. The anterior margins of the foot extend laterally as elongate appendages. Oral tentacles are short. Posterior foot is bluntly rounded and less pointed than typical of Trapania. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of welldeveloped jaws. The jaws contain several rows of acutely pointed, slightly curved, cylindrical jaw elements, equally spaced with regularly sized gaps (Fig. 11A). The radular formula of the holotype is 50 × 1.0.1 (Fig. 11B). The older teeth are much smaller than the newer ones and the radula widens towards the more newly developed teeth. The teeth bear numerous denticles with the smallest ones being found on the inner edge of the tooth. There are approximately five to ten denticles on the inner side of the much larger narrow primary cusp and one denticle on the outer side of the cusp. The older teeth (Fig. 11C) have fewer denticles than the more recently developed ones (Fig. 11D). Some of the larger more recently formed teeth have small denticles between some of the adjacent larger ones. Reproductive system: The mature reproductive system is triaulic (Fig. 4H). The tubular pre-ampullary duct is narrow where it connects to the ovotestis. From there it abruptly widens into the saccate ampulla proper, which leads to a point where it divides into the short oviduct and vas deferens. The oviduct enters the female gland mass. The vas deferens gradually widens into a prostatic portion and gradually transitions into the narrow, muscular ejaculatory portion that is continuous with the narrow penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The narrow vagina maintains its width from its opening through a long region prior to its junction with the large, pyriform bursa copulatrix and short, narrow receptaculum duct. The receptaculum duct joins the spherical receptaculum seminis near the long division of the uterine duct, which enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: In our molecular phylogeny Trapania tigger is sister to a clade that contains T. reticulata and T. darvelli. Our ABGD analysis and bPTP analysis reveal a strong genetic divergence of 8.0% in the COI gene and 8.2% in the 16S gene between T. tigger and T. reticulata and 8.7%/7.7−8.2% between T. tigger and T. darvelli, further supporting T. tigger as a distinct species within Trapania (Tables 3, 4). These three species are uniquely characterized by having thick extra-rhinophoral and extra-branchial appendages, thick rhinophores and a relatively blunt posterior end of the foot. Both T. tigger and T. reticulata have mottled brownish pigment, but in T. tigger the pattern occurs as distinct dark and light patches, whereas T. reticulata has brownish pigment surrounding yellow spots. In contrast, T. darvelli has a translucent white body with brown on the rhinophores, oral tentacles, extrarhinophoral and extra-branchial appendages and gill. All three species have radular teeth with a single, large cusp situated near the outside of the tooth and a series of smaller denticles on the inside of the cusp. In T. tigger and T. darvelli there are few denticles on the inside of the cusp (five to ten) as compared to T. reticulata (11– 18 denticles in largest teeth). The reproductive systems of T. tigger, T. reticulata and T. darvelli have significant differences. In T.tigger and T.reticulata, the pre-ampulla enters the proximal end of the ampulla subterminally, whereas it enters the proximal ampulla terminally in T. darvelli. In T. tigger, the penial sac is narrow and is the same width as the ejaculatory portion of the vas deferens, whereas in T. reticulata and T. darvelli the penial sac is much wider than the short ejaculatory portion. The bulbous penial sac of T. darvelli is short in T. darvelli and elongate in T. reticulata. Both T. tigger and T. reticulata have a straight vaginal duct, while it is curved in T. darvelli. In T. tigger and T. reticulata, the bursa copulatrix and receptaculum seminis are about the same size, while the bursa of T. darvelli is much larger than the receptaculum.

MURPHYDORIS MARACABRANCHIA SP. NOV. (FIGS. 2E, 3G, H, 6A–C) ‘ Goniodoridella sp. 4 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 4 ’ – Gosliner et al. (2018: 50). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 79EC6092-71AA-4222-9E61-1728A47A142D. Material examined: Holotype. CASIZ 186120 (Figs 1, 2E) (Philippines, Batangas Province, Maracaban Island, Sepok, 13°41′16.98′′N, 120°49′37.884′′E; coll. P. Paleracio, 25 May, 2011), one preserved specimen, 3 mm long. Paratypes. CASIZ 222056, CASIZ 224657 (Philippines, Romblon, Romblon Province, Logbon Reef). CASIZ 222056. (coll. P. Eschweiler, 3 April 2017). CASIZ 224657 (Figs 3H, 6 A–C) (coll. T. M. Gosliner, 8 March 2018), dissected (SEM: Radula). CASIZ 222015 (Philippines, Romblon Province, Alad Island, 0.0– 0.5 m depth; coll. T. M. Gosliner, 7 April 2017). Distribution: Papua New Guinea (Anderson, 2015, Gosliner et al., 2015, 2018), Japan (Gosliner et al., 2015, 2018) and the Philippines: Romblon Island, Batangas (Kim, 2021, present study). Ecology: The species is found on the underside coral rubble (Gosliner etal., 2008, 2015, 2018), from intertidal (present study) to 14 m deep (Anderson, 2015). Etymology: Murphydoris maracabranchia is named after the shape of the gill (Latin branchia), having the tips wide and rounded, which resemble the musical percussion instrument called maracas (from Guaraní mbaracás). External morphology (Figs 2E, 3G): Preserved specimens between 3 and 5 mm length. Body limaciform and narrow. Foot tipped at end, rounded at anterior part, with small tubercles in dorsal part. Notal border well developed, reduced and serrated due to presence of spicules. Spicules form small tubercles on dorsal part of tail. Notal border ends in two lateral processes in back-posterior part of notum, one on each side of anal opening. Processes narrow at base, ending in widened, rounded tip. They are likely a modification of the branchial gill (Fig. 3G). Dorsal or lateral papillae absent. Rhinophores non-retractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, without oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense network of spicules. Colour pattern (Fig. 2E): Body uniformly opaque white with tips of rhinophores red or orange. A thin maroon line surrounds body behind rhinophores. Foregut anatomy (Fig. 6A–C): Buccal bulb muscular. Dorsal buccal pump elongates backwards. Radular sac short descending ventrally. Thin oesophagus originates at posterior end of buccal bulb, behind buccal pump. Nervous system surrounds this area. Oesophageal pump and salivary glands were not observed due to the small size of specimens. Labial cuticle surrounds lips and expands within buccal pump. Radula could not be opened due to the small size. Estimated radular formula 13 × 1.1.0.1.1. Inner lateral tooth large and robust, with masticatory margin bearing denticles (Fig. 6A, B). Outer lateral tooth with two thin and sharp cusps, the upper somewhat longer than lower (Fig. 6C). Base of outer tooth wide and rectangular. Reproductive system (Fig. 3H): Reproductive system located at anterior-third of body. Thin hermaphroditic duct begins at ovotestis, located inside digestivehermaphroditic gland. Hermaphroditic duct expands in wide, pear-shaped ampulla. Thin post-ampullary duct arises from ampulla and divides into prostatic portion of vas deferens and oviduct. Oviduct enters in female gland mass. Prostate not differentiated, same width as ejaculatory portion. Thin and elongate vas deferens terminates in wide penial sac in most distal part. Vagina wider than vas deferens, connecting with small, rounded bursa copulatrix. Short duct arises from base of bursa copulatrix and divides into two thin ducts; one connects with small, oval receptaculum seminis. Receptaculum seminis slightly smaller than bursa copulatrix. Second duct corresponds to thin and short uterine duct, which enters in female gland mass. Remarks: Murphydoris maracabranchia is the second white species of Murphydoris described. Murphydoris maracabranchia and M. cobbi have a white body with a brown line behind the rhinophores. However, M. cobbi has small brown spots along the body, whereas M.maracabranchia lacksthese.Inaddition, therhinophores of M. maracabranchia have red to orange tips, while the rhinophores of M. cobbi are entirely white. The gill of M.maracabranchia is narrow at base with a wide, rounded tip, whereas M. singaporensis and M. adusta have several branches arising from same stalk and M. cobbi has wingshaped gill. Murphydoris maracabranchia also differs from other species of Murphydoris in the internal anatomy. It is the only species of Murphydoris without a differentiated prostate. Moreover, the uterine duct emerges in the middle of a duct between the bursa copulatrix and seminal receptacle in M. maracabranchia, whereas in M. adusta and M. cobbi the uterine duct emerges at distal part of the vagina and at the base of the receptaculum seminis in M. singaporensis (Table 3). Molecular analyses support the difference among species (Fig. 1). Moreover, the p -distance for COI between M. cobbi and M. maracabranchia is 21.4–21.9%. The p -distance for 16S is 18.0–18.6% between M. maracabranchia and M. adusta and 17.7–18.6% between M. maracabranchia and M. cobbi (Table 2)., Published as part of Paz-Sedano, Sofía, Smirnoff, Dimitri, Candás, María, Gosliner, Terrence M. & Pola, Marta, 2022, Rediscovering the overlooked genus Murphydoris (Nudibranchia: Goniodorididae): the first phylogeny and addition of four new species, pp. 250-269 in Zoological Journal of the Linnean Society 196 on pages 260-262, DOI: 10.1093/zoolinnean/zlac002, http://zenodo.org/record/7037692, {"references":["Gosliner TM, Valdes A, Behrens D. 2015. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Gosliner TM, Valdes A, Behrens D. 2018. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Anderson J. 2015. Goniodoridella sp. 4. In: Papua New Guinea nudibranchs & sea slugs. Available at: http: // www. nudibranch. org / Papua % 20 New % 20 Guinea % 20 Sea % 20 Slugs / html / nudibranchs / Goniodoridella % 20 sp 4 % 2001. html","Kim L. 2021. Goniodoridella sp. 4 (NSSI), Romblon Island, Philippines. In: The Three P Beach Resort Romblon (Flickr photostream). Available at: https: // www. flickr. com / photos / the _ three _ p _ romblon / 50280994712"]}

TRAPANIA LEMANIOIDES SP. NOV. (FIGS 2D, E, 4D, 7) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 172793CC-C647-44F6-8926-8AB7593490B2. Trapania sp. 2 Gosliner et al., 2018: 59, top-right photograph. Type material: Holotype: NMP 041335, originally CASIZ 208433, one specimen, partially dissected and sequenced, Manila Channel (13.52230°N 120.94850°E), Puerto Galera, Mindoro Oriental Province, Mindoro, Philippines, 3−23 m, 13 April 2015, collected by P. J. Aristoreanas. Paratypes: CASIZ 221979, one specimen, sequenced, Romblon Island (12.53267°N 122.24998°E), Romblon Province, Philippines, 7–16 m depth, 05 April 2017, collected by Joseph Maestro. CASIZ 222066, one specimen, dissected and sequenced, Romblon Island (12.55660°N 122.24940°E), Romblon Province, Philippines, 4–21 m depth, 04 April 2017, collected by Joseph Maestro. Ty p e l o c a l i t y: M a n i l l a C h a n n e l (1 3.5 2 2 3 0°N 120.94850°E), Puerto Galera, Mindoro Oriental Province, Mindoro, the Philippines. Geographical distribution: Known only from the Philippines. Natural history: Trapania lemanioides is often associated with large colonies of the octocoral genus Lemnalia Gray, 1868. It is cryptically coloured on these soft coral colonies. It does not appear to feed on the soft coral but is frequently associated with it. Etymology: This species named for its resemblance to the octocoral genus Lemnalia, with which this species is often associated. External morphology: Living animals are 3−5 mm (Fig. 2D, E). Body opaque, slightly pink and covered unevenly in a frosting of white pigmentation. The rhinophores, evenly wide along their length, have four to five lamellae with a pointed tip and are particularly white at their apex. The bipinnate gill plume is small and white, consisting of three branches. Both extrarhinophoral and extra-branchial appendages are average sized and particularly white. The anterior margins of the foot extend laterally as elongate appendages are more translucent than the rest of the body and are slightly longer than the oral tentacles. The long, conical oral tentacles and oral veil are particularly white. The white pigmentation on the dorsal ridge of the most posterior portion of the foot appears as if it is clumpy. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. It was not possible to discern any details of the structure of the jaws. The radular formula in the specimen CASIZ 222066 is 16–17 × 1.0.1 (Fig. 7A) with one additional tooth being present on the right side of the radula. The left side of the radula appears to have one additional tooth, hence the variation in formula. The radula of the holotype was not removed. The older teeth (Fig. 7B) are much smaller than the newer ones and the radula widens towards the more newly developed teeth (Fig. 7C, D). The teeth bear numerous evenly graded, elongate denticles with the smallest ones being found on the inner edge of the tooth. There are approximately three to six denticles on the inner side of the much larger, narrow primary cusp and one denticle on the outer side of the cusp. The older teeth have fewer denticles than the more newly developed teeth. Reproduction system: In the holotype NMP 041335, the mature reproductive system is triaulic (Fig. 4D). The reproductive system of the paratype (CASIZ 222066) was immature. The narrow pre-ampullary duct enters the spherical ampulla at its base, adjacent to the vas deferens. The oviduct departs the ampulla on the other side of the vas deferens enters the female gland mass. The vas deferens is equally sized throughout its length until a sharp turn where it transitions into an equally sized muscular ejaculatory portion that is continuous with the wide penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The short vagina is uniformly narrow throughout its length and joins the small, pyriform bursa copulatrix and short narrow receptaculum duct. The receptaculum duct joins the equally small pyriform receptaculum seminis near the division of the long uterine duct, which enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: In our molecular phylogeny, Trapania lemanioides is sister to a clade composed of T. circinata and T. naeva. Our ABGD and bPTP analyses reveal a strong genetic divergence of 13.9–15.7% in the COI gene and 10.3−10.6% in the 16S gene between T. lemanioides and T. naeva and 12.2−14.0% in the COI gene between T. lemanioides and T. circinata, while intraspecific variation ranged between 0.9 and 1.5% in the three Philippine specimens of T. lemanioides studied here (Tables 3, 4). Morphologically, T. lemanioides is distinct from T.circinata and T. naeva. The latter two species have black markings on the rhinophores, oral tentacles, extrarhinophoral appendages extra-branchial appendages and gill that are not present in T. lemanioides, which has a uniformly, frosted white or pinkish white coloration. Additionally, T. naeva has large, black spots on the body. The only other species that has a largely uniform white coloration is Trapania armilla Gosliner & Fahey, 2008. However, this species has a distinctive dark ring on the oral tentacles and large tubercles on the body, which are both absent in T. lemanioides. The radular teeth of T. lemanioides are similar to those of T. circinata and T. naeva, with widely separated denticles and a single, large cusp near the outer margin of the tooth. Both T. circinata and T. naeva have more denticles (eight to nine and seven to nine, respectively) inside the large cusp, whereas there are only three to six in T. lemanioides. In all three species, the ampulla is rounded and the pre-ampullary duct enters the ampulla near its distal end. In T. circinata and T. naev, there is a narrowing of the ejaculatory duct prior to its entry into the wide portion of the penial sac. In T. lemanioides, the prostatic and ejaculatory portions of the male duct and penial sac are all of uniform diameter. In both T. naeva and T. lemanioides, the distal portion of the vagina is much wider than the proximal portion, while in T. circinata the vagina is uniformly narrow throughout., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 288-289, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Gosliner TM, Fahey SJ. 2008. Systematics of Trapania (Mollusca: Nudibranchia: Goniodorididae) with descriptions of 16 new species. Systematics and Biodiversity 6: 53 - 98."]}

MURPHYDORIS COBBI SP. NOV. (FIGS 2C, D, 3D–F, 4E–H, 5) ‘ Goniodoridella sp. 2 ’ – Debelius & Kuiter (2007: 29). ‘ Goniodoridella sp. 1 ’ – Cobb & Willian (2006: 79). ‘ Goniodoridella sp. 8 ’ – Gosliner et al. (2015: 134). ‘ Goniodoridella sp. 8 ’ – Gosliner et al. (2018: 51). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 399921BD-F4E3-4204-A76A-8DA92ED07636. Material examined: Holotype. QMMO 85978 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 29 June 2017), one preserved specimen, 2 mm long. Paratypes. QMMO 85979, QMMO 85981 - QMMO 85984, MNCN 15.05 / 200142 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41 ′ S, 153°08′E, 3–6 m depth; coll. G. Cobb). QMMO 85979 (Fig. 2D) (29 June 2017), one preserved specimen, 4 mm long, dissected (SEM: Radula). QMMO 85981 (29 September 2019), one preserved specimen 2 mm long, µCT. QMMO 85982 (Fig. 3F) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85983 (Figs 3E, 4F–H) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85984 (Fig. 4E) (29 September 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula, labial cuticle). MNCN 15.05 /200142 (29 September 2019), 18 preserved specimens. CASIZ 186114, CASIZ 186119, CASIZ 186123 (Philippines, Luzon, Batangas Province, Mabini, Balayan Bay, Matotonngil Point, 13°45′N, 120°54′E). CASIZ 186114 (coll. A. Hermosillo and P. Paleracio, 8 May 2011), one preserved specimen, 5 mm long, dissected (SEM: Radula). CASIZ 186119 (coll. T. M. Gosliner, 3 May 2011), one preserved specimen, 4 mm long, dissected (SEM: Radula). CASIZ 186123 (coll. T. M. Gosliner, 10 May 2011). CASIZ 186116, CASIZ 217417 (Philippines, Luzon, Batangas Province, Balayan Bay, Ligpo, 13°49′N, 120°54′E). CASIZ 186116 (coll. A. Hermosillo, 2 May 2011). CASIZ 217417 (coll. T. M. Gosliner, 17 April 2016). CASIZ 217284 (Philippines, Negros, Negros Oriental Province, Siaton, Siit; coll. T. M. Gosliner, 5 April 2016). CASIZ 206795 (Figs 1, 2C) (Philippines, Luzon, Batangas Province, Lobo, Malabrigo, 13°36′N, 121°15′E; coll. P.J. Aristorenas, 15 April 2015). Distribution: Indian and Western Pacific Oceans (Cobb & Willan, 2006; Gosliner et al., 2015, 2018), including New Caledonia: Noumea (Huaux, 2021); the Philippines: Mudjimba Island (Mullins, 2009) and Batangas, (De Marchini, 2021; present study); and Australia: Queensland (Cobb, 2005; Debelius & Kuiter, 2007; Mullins, 2009; present study). Ecology: Murphydoris cobbi is found in shallow sandy slopes among coral rubble (Gosliner et al., 2015). Etymology: Murphydoris cobbi is named after Gary Cobb, who collected, photographed and made numerous comments on the specimens studied, which greatly facilitated its description. External morphology (Figs 2C, D, 3D): Preserved specimens between 2 and 4 mm length. Body tall, limaciform and narrow. Foot tapered posteriorly, with slightly rounded edge at its front end. Foot narrow, does not protrude from sides of notum. Notal border well developed, reduced, ending in two lateral processes in posterior part of notum, one on each side of anal opening. Processes wide and elongated, wing-shaped. They are likely a modification of the branchial gill (Fig. 3D). No dorsal or lateral papillae. Rhinophores nonretractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, lacking oral tentacles. Reproductive opening located halfway up right side of body. Whole body, including lateral processes, covered by network of spicules. Colour pattern (Fig. 2C, D): Body hyaline white, with dark brown line surrounding the area behind rhinophores and widens towards sides. Second transverse band often present posterior to wingshaped processes. Some specimens with some small brown spots, dispersed randomly on body or in middle and behind lateral processes. Processes with same coloration of body. Rhinophores translucent white. Foregut anatomy (Figs 3E, 4E–G, 5A): Buccal bulb muscular. Elongated and small buccal pump located dorsally. Radular sac short, descending ventrally and expanding backwards. Oesophagus originates at buccal bulb, behind buccal pump. Nervous system surrounds this area. One small rounded salivary gland presents on each side at junction of oesophagus and buccal pump (Fig. 3E). Oesophagus continues and inserts into digestive-hermaphroditic gland (Fig. 5A). Oesophagus slightly wider at middle part, but oesophageal pump not differentiated (Fig. 5A). Stomach and oesophagus meet in diffuse chamber inside digestive-hermaphroditic gland. Elongated stomach, almost as wide as intestine at its beginning, left part of body. Intestine continues laterally towards right side of body and ends in dorsal, posterior anus (Fig. 5A). Labial cuticle surrounds lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 4E). Radular formula 14–19 × 1.1.0.1.1. Inner lateral tooth large and robust, with one upper cusp. Masticatory margin with small notches (Fig. 4F, G). Base of these teeth wide and straight (Fig. 4F, G). Outer lateral tooth much smaller, hook-shaped, with two sharp cusps. Upper cusp somewhat longer than lower (Fig. 4G). Base of outer tooth wide, robust and elongated. Reproductive system (Figs 3F, 4H, 5B): Reproductive system located in anterior-third of body. Thin and elongate hermaphroditic duct begins at ovotestis, located inside digestive-hermaphroditic gland. Hermaphroditic duct expands into kidney-shaped ampulla. Ampulla becomes thin proximally and continues as a short post-ampullary duct that divides into prostatic portion of vas deferens and oviduct. Oviduct enters female gland. Prostate wide and elongated, becomes narrow and continues as thin vas deferens, ending in penial sac at most distal part. Penial sac spherical. Penis armed. Penial spines small, thin, and pointed (Fig. 4H). Vagina short, curves and widens near origin of ducts connecting with bursa copulatrix, receptaculum seminis and uterine duct. Bursa copulatrix large and pyriform. From base of bursa copulatrix, short, narrow, curly duct merges into receptaculum seminis. Receptaculum seminis pyriform, smaller than bursa copulatrix. Thin and large uterine duct connects distal part of vagina with female gland mass. Remarks: Murphydoris cobbi differs from M. singaporensis and M. adusta in its colour pattern. Murphydoris cobbi has a general white body, whereas M. singaporensis is translucent white with many dark reddish-brown blotches and spots (Sigurdson, 1991; Swennen & Buatip, 2012) and M. adusta has a chocolate brown body with pale blue, iridescent white and yellow patches. Also, M. singaporensis and M. adusta have one gill branch on each end of the mantle edge (Paz-Sedano et al., 2021b), whereas these structures are modified in M. cobbi, having wing-shaped processes that likely have a respiratory function. Internally, the radula of M. adusta has only one row of teeth per side while M. cobbi has two. Moreover, the radula of M. singaporensis has inner teeth with a denticulate masticatory margin with small, thin denticles and outer teeth with one cusp, while the masticatory margin in M. cobbi has small, discrete notches and the outer teeth have two cusps (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Within the reproductive system, the size of the vagina is notably different in M. singaporensis and M. cobbi, being thin and elongated in the former and short and wider in M. cobbi (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Also, the uterine duct arises near the base of the receptaculum seminis in M. singaporensis, while in M. cobbi it arises from the vagina (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Murphydoris adusta has a rounded bursa copulatrix and elongated receptaculum seminis, whereas M. cobbi has a pyriform bursa copulatrix and receptaculum seminis (Table 3). The molecular analyses support the difference between M. adusta and M. cobbi (Fig. 1), with a p -distance of 14.5–15.1% for 16S (Table 2).

TRAPANIA TAMARAW SP. NOV. (FIGS 12A, 4I, 13) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: F4449CAF-D81F-41CF-8FA2-FD8BE7F3710C. Trapania sp. 13 Gosliner et al., 2018: 62, middle-right photograph. Type material: Holotype: NMP 041334, originally CASIZ 208391, one specimen, dissected and sequenced, Puerto Galera (13.51350°N 120.95833°E), Mindoro, Mindoro Oriental Province, Philippines, 15–20 m depth, 28 March 2015, collected by P. J. Aristorenas. Type locality: Puerto Galera (13.51350°N 120.95833°E), Mindoro, Mindoro Oriental Province, the Philippines. Geographical distribution: Known only from Puerto Galera, the Philippines. Etymology: A tamaraw is the native name for the dwarfed hoofed buffalo endemic to Mindoro, Bubalus mindorensis (Heude, 1888). The horns of this buffalo resemble the well-developed and large jaws of this species compared to other members of the genus Trapania. External morphology: Live animal 4 mm (Fig. 12A). Live animal coloured with alternating blotches of dark brown/ black and yellowish tan across the entire body.Rhinophores are cylindrical halfway up and conical to a blunt tip the rest of the way with five to six lamellae. Top two-thirds of the rhinophores have a purple-brown appearance, while the bottom third has light coloration, similar to other parts of the body. Gill plume consists of three branches that have a translucent yellow-tan appearance with the same coloration as the rhinophores along the gill plume tips. Extra-branchial and extra-rhinophoral appendages are elongated and curved, typical of Trapania. Each is coloured yellowish tan with a single spot of purple/brown coloration halfway along their length. Digitiform oral tentacles are dark brown/black at their base and yellowish tan at their distal halves. The anterior margins of the foot extend laterally as elongate appendages are dark brown/black at their base and translucent yellowish at their distal halves. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of extremely well-developed jaws.The jaws (Fig. 13A) contain a single row of massive, acutely pointed, slightly curved, triangular jaw elements, equally spaced with regularly sized gaps (Fig. 13B). The radular formula of the holotype is 20 × 1.0.1 (Fig. 13C). The older teeth are much smaller than the newer ones (Fig. 13D) and the radula widens towards the more newly developed teeth. The teeth bear numerous denticles with the smallest ones being found on the inner edge of the tooth. There are approximately six to eight denticles on the inner side of the much larger, narrow primary cusp, and either no denticle on the outer side of the cusp or a series of up to eight minute, almost vestigial, short denticles on a few teeth. The denticles on the inner side of the primary cusp are generally larger nearer the primary cusp, but are not evenly graded. Reproductive system: The mature reproductive system is triaulic (Fig. 4I). The saccate ampulla is narrow at the distal end where it connects to the ovotestis. From there, it abruptly widens before leading to a point where it divides into the short oviduct and vas deferens. The short, oviduct enters the female gland mass. The vas deferens gradually widens into a curved, thick prostatic portion and gradually transitions into the narrow, muscular ejaculatory portion that is continuous with the narrow, moderately short penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The narrow vagina maintains its width from its opening through a long region prior to its junction with the large, ovoid bursa copulatrix and short narrow receptaculum duct. The receptaculum duct joins the smaller spherical receptaculum seminis near the long division of the short uterine duct, which enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: This species is compared to the four other members of its species complex below in the remarks following Trapania kamagong., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 296-297, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc."]}

TRAPANIA RETICULATA RUDMAN, 1987 (FIGS 2G, 4F, 9) Trapania reticulata Rudman, 1987: 201−203, figs 7C, 10; Wägele et al., 2006: fig. 12R; Gosliner et al., 2008: 138, second photograph from the bottom; Hervé, 2010: 156, middle photograph; Humann & Loach, 2010: 307, bottom-left photograph; Gosliner et al., 2015: 140, top photograph; Gosliner et al., 2018: 57, bottom photograph. Material examined: CASIZ 191431, one specimen, dissected and sequenced, Tab Island, Madang Province, Papua New Guinea, 9–12 m depth, 26 November 2012, collected by Vanessa Knudson. Geographical distribution: Known from Australia (R u d m a n, 1 9 8 7; W ä g e l e e t a l., 2 0 0 6; G o s l i n e r et al., 2015, 2018), New Caledonia (Hervé, 2010), Thailand (Neal, 2009) and Papua New Guinea (present study). External morphology: The living animal has been well described previously Rudman (1987) and the specimen depicted in Figure 2G closely resembles that described by him. Buccal mass: The buccal mass is muscular with a moderately enlarged buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of well-developed jaws. The jaws contain several rows of acutely pointed jaw elements of various sizes that are tightly packed together with a few gaps between them (Fig. 9A). The radular formula is 34 × 1.0.1 (Fig. 9B). The older teeth (Fig. 9C) are somewhat smaller than the newer ones and the radula widens gradually towards the more newly developed teeth (Fig. 9D). All of the teeth are narrow at the inner base and broaden out at the outer edges. The teeth bear numerous denticles, with the smallest ones being found on the inner edge of the tooth. Despite the inner denticles being smaller, the denticles are evenly graded and all much smaller than the narrow primary cusp. There are approximately 14–17 denticles on the inner side of the much larger primary cusp and one short, triangular denticle on the outer side of the cusp. The older teeth have fewer denticles than the more recently developed ones. Reproductive system: The mature reproductive system is triaulic (Fig. 4F). The narrow pre-ampullary duct enters the elongate, saccate ampulla near the proximal end of the ampulla. The ampulla narrows again at the post-ampullary duct and divides into the short, narrow oviduct, which enters the female gland mass, and the vas deferens. The vas deferens gradually widens into the thick prostatic portion that loops prior to expanding distally, immediately before it narrows abruptly into the short, narrow ejaculatory segment. The ejaculatory segment widens into an elongate penial sac. The penial sac terminates adjacent to the vagina. The width of the vagina remains uniform along a long length until it enters the base of the large, irregularly shaped bursa copulatrix adjacent to the receptaculum duct. The short receptaculum duct enters the base of the smaller, smaller slightly pyriform receptaculum seminis. The moderately long uterine duct emerges from the receptaculum duct near the base of the receptaculum and enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: Our molecular phylogeny reveals that Trapania reticulata is strongly related to a weakly supported clade of T. darvelli and the ABGD analyses reveal a low genetic divergence of 1.2–1.4% in the COI gene and 1.6% in the 16S gene between T. reticulata and T. darvelli (Tables 3, 4). The COI ABGD analysis successfully recovers T. reticulata separately from T. darvelli when the prior maximal distance was smaller than 0.003, but when the prior maximal distance was greater than 0.005, the COI ABGD analysis was unable to recover T. reticulata separately from T. darvelli. The 16S ABGD analysis also fails to recover T. reticulata as a distinct species from T. darvelli at all prior maximal distances studied here. Additionally, the bPTP analysis also successfully recovers T. reticulata separately from T.darvelli. Despite the low genetic distances between these two species, there are strong morphological characteristics that support T. reticulata and T. darvelli as distinct species. Externally, the two species have dramatically different colour patterns. In T. reticulata the body colour is translucent brown with darker brown reticulations that surround yellow spots and opaque white areas with similarly coloured oral tentacles and extra-rhinophoral and extra-branchial appendages, while in T. darvelli the body colour is solid white with brown on the oral tentacles and extra-rhinophoral and extra-branchial appendages. Both species do have proportionately thick extra-rhinophoral and extra-branchial appendages and elongate radular teeth with many denticles on the inside of the large cusp near the outer border of the tooth. The radular teeth of T. reticulata (Fig. 10A–C; Rudman, 1987: fig. 10A– C) are distinctly straight with a narrow interior base that widens externally giving the tooth base a triangular appearance. In contrast the teeth of T. darvelli (Rudman, 1987: fig. 15 A–C; Fig. 10) are more broadly arched with a much wider primary cusp. The reproductive system of T. reticulata (Fig. 4F) differs in several key features from that of T. darvelli (Fig. 4G). The pre-ampullary duct of T. reticulata enters the proximal end of the ampulla below the proximal terminus of the ampulla, whereas it enters directly into the proximal terminus in T. darvelli. The prostate of T. reticulata is not as convoluted as that of T. darvelli. Most significantly, T. reticulata has a short, narrow ejaculatory segment that enters a long, wide penial sac, whereas T. darvelli has a long, narrow ejaculatory segment that enters a short bulbous penial sac. Both species have a long vagina, which is straight in T. reticulata and more curved in T. darvelli. The bursa copulatrix and receptaculum seminis of T. reticulata are more similar in size, whereas the bursa is much larger in T. darvelli., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 291-293, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Rudman WB. 1987. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53: 189 - 212.","Wagele H, Burghardt I, Anthes N, Evertsen J, Klussmann-Kolb A, Brodie GD. 2006. Species diversity of opisthobranch molluscs on Lizard Island, great barrier reef, Australia. Records of the Western Australian Museum 69: 33 - 59.","Herve JF. 2010. Guide des Nudibranches de Nouvelle- Caledonie et autres Opisthobranches. Noumea: Editions Catherine Ledru.","Gosliner TM, Behrens DW, Valdes A. 2015. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Neal L. 2009. Trapania reticulata from the Andaman Sea. In: Sea slug forum. Sydney: Australian Museum. Available at: http: // www. seaslugforum. net / find / 22804. Accessed 29 November 2021."]}

TRAPANIA SP. ‘ANILAO’ (FIGS 12C, 15, 16A) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. o r g: a c t: F 9 7 D E B F 9 - 5 E B B - 4 D C 8 - B C 9 6 - 5BEF74E77AF3. Material examined: NMP 041329, originally CASIZ 182907, one specimen, dissected and sequenced, Anilao Harbor (13.75993° N 120.92617°E), Balayan Bay, Batangas Province, Luzon Island, Philippines, May 2010, collected by Peri Paleracio. Geographical distribution: Known only from the Philippines. External morphology: Preserved animal (Fig. 12C) 15 mm. The preserved animal has an opaque grey body covered dorsally with speckled red-brown colour characterized by small, white specks and large, white blotches of pigmentation. Notable blotches reside laterally below the rhinophores, below the gill plume and at the posterior portion of the foot. Extrarhinophoral and extra-brachial appendages have red-brown and white speckling throughout most of their length. Both pairs of appendages have a white transverse band around their middle and an opaque white apex. Rhinophores with nine lamellae and a pointed tip are uniformly wide along their length and the same colour as the red-brown body, except for white tip. Oral tentacles are coloured at their base and opaque white at the tip. Similarly, the anterior margins of the foot, which extend laterally as elongate appendages, are coloured at their base and opaque grey at their tip. The gill plume is composed of three large and highly branched elements the same colour as the body. Buccal mass: The buccal mass is muscular with a moderately enlarged buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of well-developed jaws. The jaws contain two to three rows of rounded, stubby jaw elements (Fig. 15A). The radular formula of the holotype is 44–45 × 1.0.1 (Fig. 15B). The older teeth (Fig. 15C) are much smaller than the newer ones (Fig. 15D) and the older teeth have proportionately larger denticles than the newer ones. The radula widens gradually towards the more newly developed teeth. The teeth bear numerous relatively evenly graded, short denticles with the smallest ones being found on the inner edge of the tooth. There are approximately seven to 15 denticles on the inner side of the much larger, wide primary cusp and one denticle on the outer side of the cusp. The older teeth have fewer denticles than the more recently developed ones. Reproductive system: The mature reproductive system is triaulic (Fig. 16A). The narrow pre-ampullary duct enters the saccate ampulla at the distal portion of the ampulla, adjacent to the post-ampullary duct. The ampulla narrows again at the post-ampullary duct and divides into the short, narrow oviduct, which enters the female gland mass, and the vas deferens. The vas deferens gradually widens into the thick prostatic portion. At its proximal end, the vas deferens abruptly narrows into a short, thin ejaculatory segment, which again widens into the elongate, tubular penial sac.The penial sac terminates adjacent to the vagina. The vagina is relatively long and narrow throughout most of its entire length, except at the short, wider portion at the distal end where it joins with the penial sac. It has one short loop near its proximal end. The vagina enters the base of the large, spherical bursa copulatrix adjacent to the receptaculum duct. The receptaculum duct curves and enters the base of the smaller, spherical receptaculum seminis. The uterine duct emerges from the receptaculum duct near the base of the receptaculum and enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: This species is compared to the four other members of its species complex above. Despite the fact that this species is genetically distinct in the COI gene from other members of its species complex of four species and has distinct morphological features, it is known from only one preserved specimen. For that reason, we have not described this species until living specimens have been found and variation is better established., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 300-302, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746

MURPHYDORIS COBBI SP. NOV. (FIGS 2C, D, 3D–F, 4E–H, 5) ‘ Goniodoridella sp. 2 ’ – Debelius & Kuiter (2007: 29). ‘ Goniodoridella sp. 1 ’ – Cobb & Willian (2006: 79). ‘ Goniodoridella sp. 8 ’ – Gosliner et al. (2015: 134). ‘ Goniodoridella sp. 8 ’ – Gosliner et al. (2018: 51). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 399921BD-F4E3-4204-A76A-8DA92ED07636. Material examined: Holotype. QMMO 85978 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 29 June 2017), one preserved specimen, 2 mm long. Paratypes. QMMO 85979, QMMO 85981 - QMMO 85984, MNCN 15.05 / 200142 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41 ′ S, 153°08′E, 3–6 m depth; coll. G. Cobb). QMMO 85979 (Fig. 2D) (29 June 2017), one preserved specimen, 4 mm long, dissected (SEM: Radula). QMMO 85981 (29 September 2019), one preserved specimen 2 mm long, µCT. QMMO 85982 (Fig. 3F) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85983 (Figs 3E, 4F–H) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85984 (Fig. 4E) (29 September 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula, labial cuticle). MNCN 15.05 /200142 (29 September 2019), 18 preserved specimens. CASIZ 186114, CASIZ 186119, CASIZ 186123 (Philippines, Luzon, Batangas Province, Mabini, Balayan Bay, Matotonngil Point, 13°45′N, 120°54′E). CASIZ 186114 (coll. A. Hermosillo and P. Paleracio, 8 May 2011), one preserved specimen, 5 mm long, dissected (SEM: Radula). CASIZ 186119 (coll. T. M. Gosliner, 3 May 2011), one preserved specimen, 4 mm long, dissected (SEM: Radula). CASIZ 186123 (coll. T. M. Gosliner, 10 May 2011). CASIZ 186116, CASIZ 217417 (Philippines, Luzon, Batangas Province, Balayan Bay, Ligpo, 13°49′N, 120°54′E). CASIZ 186116 (coll. A. Hermosillo, 2 May 2011). CASIZ 217417 (coll. T. M. Gosliner, 17 April 2016). CASIZ 217284 (Philippines, Negros, Negros Oriental Province, Siaton, Siit; coll. T. M. Gosliner, 5 April 2016). CASIZ 206795 (Figs 1, 2C) (Philippines, Luzon, Batangas Province, Lobo, Malabrigo, 13°36′N, 121°15′E; coll. P.J. Aristorenas, 15 April 2015). Distribution: Indian and Western Pacific Oceans (Cobb & Willan, 2006; Gosliner et al., 2015, 2018), including New Caledonia: Noumea (Huaux, 2021); the Philippines: Mudjimba Island (Mullins, 2009) and Batangas, (De Marchini, 2021; present study); and Australia: Queensland (Cobb, 2005; Debelius & Kuiter, 2007; Mullins, 2009; present study). Ecology: Murphydoris cobbi is found in shallow sandy slopes among coral rubble (Gosliner et al., 2015). Etymology: Murphydoris cobbi is named after Gary Cobb, who collected, photographed and made numerous comments on the specimens studied, which greatly facilitated its description. External morphology (Figs 2C, D, 3D): Preserved specimens between 2 and 4 mm length. Body tall, limaciform and narrow. Foot tapered posteriorly, with slightly rounded edge at its front end. Foot narrow, does not protrude from sides of notum. Notal border well developed, reduced, ending in two lateral processes in posterior part of notum, one on each side of anal opening. Processes wide and elongated, wing-shaped. They are likely a modification of the branchial gill (Fig. 3D). No dorsal or lateral papillae. Rhinophores nonretractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, lacking oral tentacles. Reproductive opening located halfway up right side of body. Whole body, including lateral processes, covered by network of spicules. Colour pattern (Fig. 2C, D): Body hyaline white, with dark brown line surrounding the area behind rhinophores and widens towards sides. Second transverse band often present posterior to wingshaped processes. Some specimens with some small brown spots, dispersed randomly on body or in middle and behind lateral processes. Processes with same coloration of body. Rhinophores translucent white. Foregut anatomy (Figs 3E, 4E–G, 5A): Buccal bulb muscular. Elongated and small buccal pump located dorsally. Radular sac short, descending ventrally and expanding backwards. Oesophagus originates at buccal bulb, behind buccal pump. Nervous system surrounds this area. One small rounded salivary gland presents on each side at junction of oesophagus and buccal pump (Fig. 3E). Oesophagus continues and inserts into digestive-hermaphroditic gland (Fig. 5A). Oesophagus slightly wider at middle part, but oesophageal pump not differentiated (Fig. 5A). Stomach and oesophagus meet in diffuse chamber inside digestive-hermaphroditic gland. Elongated stomach, almost as wide as intestine at its beginning, left part of body. Intestine continues laterally towards right side of body and ends in dorsal, posterior anus (Fig. 5A). Labial cuticle surrounds lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 4E). Radular formula 14–19 × 1.1.0.1.1. Inner lateral tooth large and robust, with one upper cusp. Masticatory margin with small notches (Fig. 4F, G). Base of these teeth wide and straight (Fig. 4F, G). Outer lateral tooth much smaller, hook-shaped, with two sharp cusps. Upper cusp somewhat longer than lower (Fig. 4G). Base of outer tooth wide, robust and elongated. Reproductive system (Figs 3F, 4H, 5B): Reproductive system located in anterior-third of body. Thin and elongate hermaphroditic duct begins at ovotestis, located inside digestive-hermaphroditic gland. Hermaphroditic duct expands into kidney-shaped ampulla. Ampulla becomes thin proximally and continues as a short post-ampullary duct that divides into prostatic portion of vas deferens and oviduct. Oviduct enters female gland. Prostate wide and elongated, becomes narrow and continues as thin vas deferens, ending in penial sac at most distal part. Penial sac spherical. Penis armed. Penial spines small, thin, and pointed (Fig. 4H). Vagina short, curves and widens near origin of ducts connecting with bursa copulatrix, receptaculum seminis and uterine duct. Bursa copulatrix large and pyriform. From base of bursa copulatrix, short, narrow, curly duct merges into receptaculum seminis. Receptaculum seminis pyriform, smaller than bursa copulatrix. Thin and large uterine duct connects distal part of vagina with female gland mass. Remarks: Murphydoris cobbi differs from M. singaporensis and M. adusta in its colour pattern. Murphydoris cobbi has a general white body, whereas M. singaporensis is translucent white with many dark reddish-brown blotches and spots (Sigurdson, 1991; Swennen & Buatip, 2012) and M. adusta has a chocolate brown body with pale blue, iridescent white and yellow patches. Also, M. singaporensis and M. adusta have one gill branch on each end of the mantle edge (Paz-Sedano et al., 2021b), whereas these structures are modified in M. cobbi, having wing-shaped processes that likely have a respiratory function. Internally, the radula of M. adusta has only one row of teeth per side while M. cobbi has two. Moreover, the radula of M. singaporensis has inner teeth with a denticulate masticatory margin with small, thin denticles and outer teeth with one cusp, while the masticatory margin in M. cobbi has small, discrete notches and the outer teeth have two cusps (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Within the reproductive system, the size of the vagina is notably different in M. singaporensis and M. cobbi, being thin and elongated in the former and short and wider in M. cobbi (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Also, the uterine duct arises near the base of the receptaculum seminis in M. singaporensis, while in M. cobbi it arises from the vagina (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Murphydoris adusta has a rounded bursa copulatrix and elongated receptaculum seminis, whereas M. cobbi has a pyriform bursa copulatrix and receptaculum seminis (Table 3). The molecular analyses support the difference between M. adusta and M. cobbi (Fig. 1), with a p -distance of 14.5–15.1% for 16S (Table 2)., Published as part of Paz-Sedano, Sofía, Smirnoff, Dimitri, Candás, María, Gosliner, Terrence M. & Pola, Marta, 2022, Rediscovering the overlooked genus Murphydoris (Nudibranchia: Goniodorididae): the first phylogeny and addition of four new species, pp. 250-269 in Zoological Journal of the Linnean Society 196 on pages 259-260, DOI: 10.1093/zoolinnean/zlac002, http://zenodo.org/record/7037692, {"references":["Debelius H, Kuiter RH. 2007. Nudibranchs of the world. Frankfurt: IKAN.","Gosliner TM, Valdes A, Behrens D. 2015. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Gosliner TM, Valdes A, Behrens D. 2018. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Cobb G, Willan RC. 2006. Undersea jewels: a colour guide to nudibranchs. Canberra: Australian Biological Resources Study.","Huaux M. 2021. Goniodoridella sp 1. In: Flora and fauna of New Caledonia. Available at: https: // endemia. nc / en / faune / fiche 5971","Mullins D. 2009. Goniodoridella sp. from S. E. Queensland. In: Sea slug forum. Available at: http: // www. seaslugforum. net / find / 22015","De Marchini F. 2021. Available at: https: // skaphandrus. com / en / underwater-photography / photo / 12443","Cobb G. 2005. Goniodoridella sp 5. In: Nudibranchs of the Sunshine Coast, Queensland, Australia, Indo-Pacific. Available at: http: // nudibranch. com. au / pages / 9667 b. htm","Swennen C, Buatip S. 2012. Murphydoris singaporensis (Gastropoda: Nudibranchia: Goniodorididae) found in Thailand; with external and first internal description of the specimens. Strombus 19: 9 - 14.","Paz-Sedano S, Diaz-Agras G, Gosliner TM, Pola M. 2021 b. Revealing morphological characteristics of Goniodorididae genera (Mollusca: Nudibranchia). Organisms Biodiversity and Evolutionary Research. Available at: https: // link. springer. com / article / 10.1007 / s 13127 - 021 - 00508 - w citeas."]}

MURPHYDORIS PUNCTICULATA SP. NOV. (FIGS 2F, G, 3I–K, 6D–H) ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2008: 126). ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2018: 50). ‘ Goniodoridella sp. 11’ – Nakano (2018: 169). Z o o b a n k r e g i s t r a ti o n: u r n: l s i d: z o o b a n k. org:act: 8DB73463-292D-45AC-B0BB-EE3481B62319. Material examined: Holotype. WAM S72711 (Figs 1, 2F, 3J, K) (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 28 December 2018), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). Paratypes. MNCN 15.05 /200149– MNCN 15.05 /200150, WAM S72710, WAM S71781, WAM S71785, WAM S17182, WAM S72709, WAM S72089, WAM S72462 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb). MNCN 15.05 / 200149 (Fig. 6G) (11 December 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula, labial cuticle). MNCN 15.05 / 200150 (Fig. 6G) (2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S72710 (Fig. 2G) (15 December 2018), one preserved specimen, 3 mm long. WAM S71781 (May 2018), 1 preserved specimen, 4 mm long, dissected (SEM: Radula). WAM S71785 (Fig. 6D–F) (May 2018), one preserved specimen 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S17182 (May 2018), one preserved specimen, 3 mm long. WAM S72709 (15 December 2018), one preserved specimen, 5 mm long. WAM S72089 (20 November 2017), two preserved specimens. WAM S72462 (4 December 2016), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). CASIZ 222055 (Philippines, Visayas, Romblon Province, Alad Isand, Romblon Province, Visayas; coll. A. Fritts-Penniman, 7 April 2017), dissected (SEM: Radula). CASIZ 224656 (Philippines, Visayas, Romblon Province, Cobrador Island; col. K. L. Larkin, 19 March 2018). Distribution: Murphydoris puncticulata is found in the western and central Pacific Ocean (Gosliner et al., 2018), in Papua New Guinea, New Caledonia (Gosliner et al., 2008, 2015), the Philippines (Gosliner et al., 2018, present study), Hawaii (Gosliner et al., 2008, 2015; Pittman & Fiene, 2020), Japan: Shizuoka prefecture (Kakegawa, 2018) and Izu Island (Nakano, 2018), Indonesia: Bacan (Sali Bay Dive Resort, 2021) and Bali (Kakegawa, 2017) and Australia: Queensland (Cobb, 2020; present study). Ecology: The species has been found under coral rubble on reefs from the intertidal up to 10 m deep (Gosliner et al., 2018). Etymology: From Latin diminutive of punctum, point or dot. Murphydoris puncticulata is named after the brown notches formed by dots. External morphology (Figs 2F, G, 3I): Preserved specimens 2–7 mm length. Body limaciform and narrow. Foot acutely tapered at posterior end, with rounded edge at anterior part. Foot narrow, does not protrude from sides of notum. Notal border evident, reduced, continuing towards trilobate gill branches located at posterior part (Fig. 3I), same level of anal opening. Several small tubercles randomly located at sides of body. Tubercles supported by internal spicules. Dorsal or lateral papillae absent. Rhinophores nonretractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone without oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense net of spicules. Colour pattern (Fig. 2F, G): Body white with small dark brown spots scattered along body. One triangular band of concentrated brown spots at anterior portion of dorsum, behind rhinophores. Transverse dark brown patch at head, between rhinophores. Third horizontal band of concentrated spots behind gill. Rhinophores, gill branches and foot whitish translucent. Foregut anatomy (Figs 3J, 6D–G): Buccal bulb muscular. Dorsal buccal pump elongates posteriorly. Radular sac short descending ventrally. Thin oesophagus originating at buccal bulb, behind buccal pump. Nervous system surrounds this area. Oesophagus becoming wider and continuing to oesophageal pump. Small, rounded salivary gland present on each side, at junction of oesophagus and buccal pump (Fig. 3J). Labial cuticle surrounding lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 6D). Radular formula 13 × 1.1.0.1.1. Inner lateral tooth large and robust, with one cusp and masticatory margin bearing between five and seven sharped denticles (Fig. 6E, F). Base of these teeth wide and straight (Fig. 6E). Outer lateral tooth much smaller with two sharp cusps, the upper somewhat longer than the lower (Fig. 6G). Base of outer tooth wide and rectangular. Reproductive system (Figs 3K, 6H): Reproductive system located at anterior-third of body. Thin preampullary duct begins at ovotestis, located inside digestive-hermaphroditic gland. Pre-ampullary duct expands into large, bean-shaped ampulla. Thin postampullary duct arises from ampulla and enters in female gland mass. Inside female gland mass it divides into prostatic portion of vas deferens and oviduct. Prostate long and wide, becomes narrow and continuing as elongated vas deferens. Vas deferent as long as prostate, ends ending in a slightly wider penial sac in most distal part. Penis armed with small, hook-shaped penial spines at its base. Spines become longer, thinner and more pointed at most distal part of penis (Fig. 6H). Vagina short, as wide as vas deferens, connects with oval bursa copulatrix. From base of bursa copulatrix two thin ducts arise; one connects with small and elongated receptaculum seminis. Second duct corresponds to thin and long uterine duct, which enters in female gland mass. Remarks: Murphydoris puncticulata, M. cobbi and M. maracabranchia share having white body colour. Murphydoris puctulatum and M. cobbi have a hyaline white body with dark brown spots, one brown patch behind the rhinophores and one behind the gill. However, in M. cobbi this patch behind the rhinophores consists of a continuous line, while in M. puncticulata it is formed by several spots. Murphydoris maracabranchia has a brown line behind the rhinophores but lacks brown spots along the body. Moreover, in M. cobbi the gill is likely modified as two wing-shaped processes at the end of the notal border, while in M. maracabranchia the gill is narrow at the base with wider and rounded tip and M. puncutulatum has three distinguishable gill branches (Fig. 3). Murphydoris puncticulata has marked tubercles along the sides of the body that are absent in M. cobbi. With respect to the reproductive system, the uterine duct arises from the middle of the vagina in M. cobbi, from the middle of a duct between the bursa copulatrix and the receptaculum seminis in M. maracabranchia and near the base of the receptaculum seminis in M. puncticulata. Also, the receptaculum seminis and the bursa copulatrix are much larger in M. cobbi than in M. puncticulata. The receptaculum seminis is pyriform in M. cobbi, oval in M. maracabranchia and elongated in M. puncticulata (Table 3). Molecular results support M. puncticulata as different species (Fig. 1), having a COI p -distance of 10.7–11.2% and 19.4% with M. cobbi and M. maracabranchia, respectively; and a 16S p- distance of 13.2–14.5%, 7.9–8.2% and 18.9–19.2% with M. adusta, M. cobbi and M. maracabranchia, respectively (Table 2)., Published as part of Paz-Sedano, Sofía, Smirnoff, Dimitri, Candás, María, Gosliner, Terrence M. & Pola, Marta, 2022, Rediscovering the overlooked genus Murphydoris (Nudibranchia: Goniodorididae): the first phylogeny and addition of four new species, pp. 250-269 in Zoological Journal of the Linnean Society 196 on pages 262-265, DOI: 10.1093/zoolinnean/zlac002, http://zenodo.org/record/7037692, {"references":["Gosliner TM, Fahey SJ. 2008. Systematics of Trapania (Mollusca: Nudibranchia: Goniodorididae) with descriptions of 16 new species. Systematics and Biodiversity 6: 53 - 98.","Gosliner TM, Valdes A, Behrens D. 2015. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Gosliner TM, Valdes A, Behrens D. 2018. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Nakano R. 2018. Field guide to sea slugs and nudibranchs of Japan. Tokyo: Bun-ichi Co., Ltd.","Pittman C, Fiene P. 2020. Goniodoridella sp. 3. In: Sea slugs of Hawai'i. Available at: http: // seaslugsofhawaii. com / species / Goniodoridella-sp 3 - a. html","Kakegawa M. 2018. Goniodoridella sp 12. In: Seaslug. world. Available at: https: // en. seaslug. world / species / goniodoridella _ sp. 12 / 11941","Sali Bay Dive Resort. 2021. Goniodoridella sp 1. In: Sali Bay Dive Resort. Available at: https: // salibayresort. com / marinelife / molluscs / nudibranchia / goniodoridella / goniodoridella-sp- 1 /. Accessed on 19 January 2022.","Kakegawa M. 2017. Goniodoridella sp 12. In: Seaslug. world. Available at: https: // en. seaslug. world / species / goniodoridella _ sp. 12 / 6449","Cobb G. 2020. Goniodoridella sp 10. In: Nudibranchs of the Sunshine Coast, Queensland, Australia, Indo-Pacific. Available at: http: // nudibranch. com. au / pages / DSC 06516. htm","Sigurdsson JB. 1991. A nudibranch, Murphydoris singaporensis, new genus and species, from Singapore mangroves (Mollusca: Opisthobranchia: Goniodorididae). Raffles Bulletin of Zoology 39: 259 - 263."]}

TRAPANIA TATSULOK SP. NOV. (FIGS 2B, 4B, 5) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. o r g: a c t: 9 D F 2 5 E E 2 - 6 5 5 D - 4 8 5 B - 9 2 A 7 - 45B7D23B188D. Trapania sp. 15 Gosliner et al., 2015: 145, middle-right photograph. Trapania sp. 17 Gosliner et al., 2018: 63, bottom-left photograph. Typematerial: Holotype: NMP041333,originallyCASIZ 200530, one specimen, dissected and sequenced, Verde Island Passage (13.92816°N 120.6109°E), Calatagan, Batangas Province, Luzon Island, Philippines, 19 May 2014, collected by Shayle Matstuda. Type locality: Verde Island Passage (13.92816ºN 120.6109°E), Calatagan, Batangas Province, Luzon Island, Philippines. G e o g r a p h i c a l d i s t r i b u t i o n: K n o w n o n l y f r o m Calatagan, the Philippines. Etymology: Tatsulok is the Tagalog word for triangle and it refers to the black equilateral triangle found on the dorsum of this species. It is a noun in apposition. External morphology: The living animal (Fig. 2B) is 5 mm. The body coloration is opaque white with dark orange pigmentation on the outer three quarters of the extra-rhinophoral and extra-branchial appendages. There is a gradation of colour such that the basal portion is more yellow orange. The extra-rhinophoral and extra-branchial appendages are somewhat curved and cylindrical with a uniform diameter throughout. The conical, perfoliate rhinophores have five to six lamellae and are dark maroon/brown except for an opaque-white base. The oral tentacles are identical to the rhinophore coloration, with black spanning between the oral tentacles colouring the anterior margin of the head with a semicircle protruding posteriorly at the medial line. The anterior margins of the foot extend laterally as elongate appendages and are more translucent than the rest of the body. This specimen has a distinct black, equilateral triangle on its dorsum centred on the body midline, midway between the gill plume and rhinophores, with its tip pointing anteriorly. There are three other black markings adorning each side of the lateral flanks, starting at the middle of the body and extending to the posterior extension of the foot. There is another black spot situated medially, posterior to the gill plume. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of well-developed jaws (Fig. 5A). The jaw surface is covered with small semi-spheres, each covered with numerous short and blunt spikes resembling a medieval flail or mace (Fig. 5B). The radular formula of the holotype is 25 × 1.0.1 (Fig. 5C) in the specimen examined and the radula was fragmented into three pieces. The older teeth are much smaller than the newer ones and lack denticles on the outer side of the radular tooth (Fig. 5C). The teeth bear numerous denticles with the smallest ones being found on the inner edge of the tooth. In the newer, larger teeth, which were separated from the main part of the radula during preparation, there are approximately five to seven denticles on the inner side of the larger primary cusp with the ones nearest the primary cusp being the largest (almost the same size as the primary cusp) and a variable number of denticles on the outer side of the cusp with some teeth entirely lacking an outer denticle. There may be as few as three larger denticles or as many as eight smaller denticles, depending on the tooth (Fig. 5D). Reproductive system: The mature reproductive system is triaulic (Fig. 4B). The pre-ampullary is narrow at the distal end where it connects to the ovotestis. It then enters the saccate, elongate ampulla near its distal end. The ampulla narrows to the point where it divides into the short oviduct and the elongate vas deferens. The oviduct enters the female gland mass. The vas deferens widens into a curved prostatic portion that narrows into a short ejaculatory segment that is contiguous with the relatively narrow penial sac.The penial sac exits adjacent to the narrow vagina. The vagina is narrow throughout its length and joins the almost spherical bursa copulatrix adjacent to the long, narrow receptaculum duct. The receptaculum duct joins the spherical receptaculum seminist adjacent to the moderately long uterine duct, which enters the female gland mass. The bursa and receptaculum are roughly the same size. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: Our molecular phylogeny, the 16S ABGD analysis and the bPTP analysis reveal that Trapania tatsulok is a distinct species of Trapania and can be distinguished by its unique colour pattern of white pigment on the body with black pigment on the oral tentacles, anterior margin of the head, on the rhinophores, medially in front and behind the gill and laterally on the sides of the body. This species is a member of a clade that includes T. scurra, T. palmula, T. japonica and T. kahel (see Remarks section below). Both T. scurra and T. palmula have a stout body with proportionately thicker extra-rhinophoral and extra-branchial processes, whereas T. japonica has a more delicate, elongate body with thin, elongate extra-rhinophoral and extra-branchial as in T. tatsulok. These last two species also have a similar colour pattern of a white body with dark brown to black markings. In T. japonica, the extra-rhinophoral and extra-branchial process are yellow, while in T. tatsulok they are orange. The dark markings of T. japonica are rounded spots, whereas they are irregular in shape in T. tatsulok, with the exception of the triangular marking anterior to the gill. The gill is white in T. tatsulok, whereas the outer portions of the gill branches are black in T. japonica. Internally, T. tatsulok has several features that clearly distinguish it from other members of the clade of which it is a member. The jaw elements of T. tatsulok contain numerous minute spikes resembling a flail, while T. scurra, T. palmula and T. japonica have simple, acutely pointed jaw elements (Gosliner & Fahey, 2008, fig. 8a, fig. 10a; Fig. 5B). The radular teeth of T. tatsulok have one or two large denticles on the inner side of the primary cusp and numerous smaller denticles, whereas the teeth of T. scurra, T. palmula and T. japonica all have a single prominent cusp and numerous smaller denticles (Gosliner & Fahey, 2008: fig. 8b–d, fig. 10b–d; Fig. 5D). The reproductive system of T. tatsulok is similar to that of T. palmula, with a relatively elongate vagina and ejaculatory portion of the vas deferens, but in T. palmula (Gosliner & Fahey, 2008: fig. 9) both the vagina and ejaculatory duct are more elongate than in T. tatsulok. In T. scurra (Gosliner & Fahey, 2008: fig. 11) and T. japonica (Fig. 4A) the vagina and ejaculatory portion of the vas deferens are much shorter than in T. tatsulok. Three other described species of Trapania from the eastern Pacific have similar colour patterns to T. tatsulok. Trapania velox, T. goslineri Millen & Bertsch, 2000 and T. darwini Gosliner & Fahey, 2008, all have white body pigment with brown or black markings and orange pigment, but differ in the pattern and distribution of these pigments. In these three species, there is both brown and orange pigment on the extra-rhinophoral and extrabranchial processes, while in T. tatsulok there is only orange. These three species also have orange rhinophores, while in T. tatsulok the rhinophores are dark brown to black. In T. velox, the translucent, white body has darkbrown longitudinal lines, while T. goslineri has brown spots. In T. darwini there is a series of longitudinal brown markings interrupted by white spots. Bhave (2010) posted a photo of another species from India that has yet another CASIZ 208433), scale = 0.33 mm. E, Trapania aurata Rudman, 1987, CASIZ 186205, scale = 0.46 mm. F, Trapania reticulata Rudman, 1987, CASIZ 191431, scale = 0.50 mm. G, Trapania darvelli Rudman, 1987, CASIZ 222004, scale = 0.4 mm. H, Trapania tigger sp. nov., holotype, NMP 041328 (formerly CASIZ 180412), scale = 1.0 mm. I, Trapania tamaraw sp. nov., NMP 041334 (formerly CASIZ 208391), scale = 0.50 mm. J, Trapania stegodon sp. nov., holotype NMP 041332 (formerly CASIZ 186206), scale = 0.50 mm. Abbreviations: am, ampulla; bc, bursa copulatrix; fgm, female gland mass; p, penial sac; pr, prostatic portion of vas deferens; rs, receptaculum seminis; ud, uterine duct; v, vagina. 286 D. S. SMIRNOFF ET AL. combination of white, orange and brown markings and differs from the species discussed above. Rudman (2010) suggested that this specimen may be T. goslineri and that T. darwini may also be the same species as T. goslineri, despite the morphological distinctions clearly noted by Gosliner & Fahey (2008). Another feature clearly distinguishes the specimen photographed by Bhave. In this specimen from India, the base of the rhinophore club is broad and bulbous with an abruptly narrow apex. All of the other species of Trapania with similar colour patterns have narrow, evenly graded rhinophores and it is likely that the specimen from India represents another distinct, undescribed species., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 283-286, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Gosliner TM, Behrens DW, Valdes A. 2015. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Baba K. 1935. Nudibranchia of Mutsu Bay. Science Reports of Tohoku University 10: 331 - 360.","Gosliner TM, Fahey SJ. 2008. Systematics of Trapania (Mollusca: Nudibranchia: Goniodorididae) with descriptions of 16 new species. Systematics and Biodiversity 6: 53 - 98.","Millen SV, Bertsch H. 2000. Three new species of dorid nudibranchs from southern California, USA, and the Baja California Peninsula, Mexico. The Veliger 43: 354 - 366.","Bhave VJ. 2010. Trapania goslineri? from Ratnagiri, India. In: Sea slug forum. Sydney: Australian Museum. Available at: http: // www. seaslugforum. net / find / 23291. Accessed 29 November 2021.","Rudman WB. 1987. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53: 189 - 212.","Rudman WB. 2010. Comment on Trapania goslineri? from Ratnagiri, India by Vishal Bhave. In: Sea slug forum. Sydney: Australian Museum. Available at: http: // www. seaslugforum. net / find / 23291. Accessed 29 November 2021."]}

Trapania sp. 1 Pittman & Fiene, 2021. Type material: Holotype: CASIZ 189444, one specimen, dissected and sequenced, Airport Beach, Maui, Hawaiʻi, USA, 12–18 m depth, 19 February 2012, collected by Cory Pittman. Other material: Trapania sp. cf. kanaloa, CASIZ 199250, one specimen, sequenced, preserved length 1.5 mm, Koloa Landing, Kauaʻi, Hawaiʻi, USA, 8–14 m depth, 30 May 2014, collected by Cory Pittman. Type locality: Airport Beach, Maui, Hawaiʻi, USA. Geographical distribution: Known only from the Hawaiian Islands. Etymology: The Hawaiʻian god Kanaloa is often symbolized by a squid or octopus form. This species is also found in association with beds of the alga Halimeda kanaloana Vroom, 2006, which is named for the same deity. It is a noun in apposition. External morphology: The living animal is 3 mm in length (Fig. 12E, upper photo). The body is brown coloured with a dense frosting of small, white and darker brown punctations and brown patches. The rhinophores are thick and bulbous with approximately eight lamellae and a white apex. They have the same cream and brown pigment as the rest of the body. The extra-rhinophoral and extra-branchial appendages are thin with a clubshaped apex and are also brown and cream, with fine opaque white punctations. The gill is small consisting of three simply pinnate branches. The posterior end of the foot is short and blunt with brown and whitish mottling. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of well-developed jaws. The jaws contain several rows of acutely pointed, cylindrical jaw elements that are tightly packed together with a few gaps between them (Fig. 18A). The radular formula of the holotype is 19 × 1.0.1 (Fig. 18B). The older teeth (Fig. 18C) are much smaller than the newer ones (Fig. 18D) and the radula widens towards the more newly developed teeth. The teeth bear numerous relatively equal-sized denticles with the smallest ones being found on the inner edge of the tooth. There are approximately six to eight denticles on the inner side of the much larger primary cusp, and one to three much smaller denticles on the outer side of the cusp. The older teeth have fewer denticles than the more recently developed ones. Reproductive system: The mature reproductive system is triaulic (Fig. 16C). The pre-ampullary duct is narrow and joins the post-ampullary duct distally to the ampulla. The pyriform ampulla narrows to the point where it divides into the short oviduct and elongate vas deferens. The oviduct enters the female gland mass. The vas deferens is initially narrow and widens into a slightly convoluted, glandular prostatic portion and gradually transitions into the wide, muscular ejaculatory portion that abruptly narrows and makes a sharp turn prior to expanding into the wide penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The vagina is equally wide throughout its length. Immediately prior to its entrance into the large spherical bursa copulatrix is a short branch of the receptaculum duct that leads to the smaller, spherical receptaculum seminis. Emerging from the receptaculum duct is a uterine duct that enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: Our molecular phylogeny, the ABGD analyses and the bPTP analysis show that Trapania kanaloa is a distinct species separate from its similarly coloured sisterspecies T. euryeia. Externally, T. euryeia has a darker body colour than T. kanaloa with more brown pigment and much larger, more regular white punctations. In our ABGD and bPTP analyses, the holotype of T. kanaloa collected from Maui is potentially distinct from the specimen we call T. sp. cf. kanaloa (CASIZ 199250) due to a high genetic divergence of 4.3% in the mitochondrial gene COI (Table 3) and a genetic divergence of 2.6% in the mitochondrial gene 16S (Table 4). This specimen was collected from Kauaʻi and was previously considered to be a variant of T. euryeia by Pittman & Fiene (2021) due to its pale external coloration; however, this specimen is probably immature (preserved length = 1.5 mm) and the photo of the living animal (Fig. 12E, lower photo) indicates that it has less concentrated brownish pigment. Owing to its small size we were not able to dissect this specimen to compare its anatomy. In contrast, the interspecific variation between T. kanaloa and T. euryeia from the Philippines is 9.3% in the COI gene (Table 3) and 5.5% in the 16S gene (Table 4) The interspecific variation between T. sp. cf. kanaloa and T. euryeia is similar, i.e. 9.3% in the COI gene (Table 3) and 5.7% in the 16S gene (Table 4), indicating that the T. sp. cf. kanaloa specimen is not T. euryeia. Further study of T. kanaloa, with additional representatives from Hawaiʻi, is necessary to determine whether one or two species are present across the Hawaiʻian Archipelago. Similarly, comparisons with additional specimens of T. euryeia from a broader geographical area are necessary to better understand the genetic variation present within and between these two species. Present data indicate that pale specimens from Hawaiʻi should be regarded as T. kanaloa. Pittman & Fiene (2021) also show a photo of a darker specimen of T. euryeia from Hawaiʻi that more closely resembles specimens from elsewhere in the Indo-Pacific, and the question remains as to whether this is conspecific with other specimens of T. euryeia from the western Pacific. The radular teeth previously described for T. euryeia are similar to those found here in T. kanaloa. There appear to be significant differences in the reproductive systems of T. kanaloa and T. euryeia. In T. kanaloa, the prostatic portion of the vas deferens is less convoluted than in T. euryeia. Most significantly, the vagina of T. kanaloa is of uniform diameter throughout its length, whereas T. euryeia has a glandular collar present around the vagina near its junction with the penial sac, which is absent in T. kanaloa., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on page 305, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746

MURPHYDORIS ADUSTA SP. NOV. (FIGS 2A, B, 3A–C, 4A–D) ‘ Goniodoris aspersa ’ – Debelius & Kuiter (2007: 28). ‘ Goniodoridella sp. ’ – Su et al. (2009: 457, fig. 11B). ‘ Goniodoridella sp. 6 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 6 ’ – Gosliner et al. (2018: 51). ‘ Goniodoridella sp. 3 ’ – Nakano (2018: 166). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: A6372BF6-E947-4470-9B1D-D22A9AB5FE58. Material examined: Holotype. QMMO 85985 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 29 September 2019), one preserved specimen, 3 mm long, dissected. Paratypes. MNCN 15.05 /200143 – MNCN 15.05 /200148, WAM S72660, QMMO 85986 – QMMO 85988; QMMO 85995 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb). MNCN 15.05 /200143 (29 September 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula). MNCN 15.05 /200144 (Fig. 3B, C) (29 September 2019), one preserved specimen 2 mm long, dissected (SEM: Radula). MNCN 15.05 /200145 (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula). MNCN 15.05 /200146 (28 November 2019), one preserved specimen, 5 mm long. MNCN 15.05 /200147 (28 November 2019), one preserved specimen, 5 mm long, dissected (SEM: Radula, penis). MNCN 15.05 /200148 (5 December 2019) one preserved specimen, 10 mm long, dissected (SEM: Radula). WAM S72660 (Figs 1, 2A) (2 May 2018), one preserved specimen, 5 mm long. QMMO 85986 (Fig. 4D) (2019), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle, penis). QMMO 85987 (2019), one preserved specimen, 3 mm long, dissected (SEM: Radula). QMMO 85988 (29 September 2019), two specimens. QMMO 85995 (Figs 2B, 4A, C) (11 December 2019), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). CASIZ 177775 (Philippines, Batangas Province, Tingloy, Maricaban Island, Aphol’s Rock, 13°45′41.74′′N, 121°23′0.56′′E; coll. T. M. Gosliner, 23 April 2008). CASIZ 186117 (Philippines, Batangas Province, Calumpan Peninsula, Mabini, Murals dive site, 13°41′56.472′′N, 120°52′57.719′′E; coll. D. Raymundo, 4 May 2011). CASIZ 181290 (Fig. 4B) (Philippines, Luzon, Batangas Province, Mabini, Mainit Bubbles dive site, 13°41.155′N, 120°53.713′E; coll. T. M. Gosliner, 21 May 2009), dissected (SEM: Radula). Distribution: Indian Ocean to Indonesia, western Pacific Ocean; Madagascar (Gosliner et al., 2015, 2018); Indonesia: Palau Sangeang (Anderson, 2021) and Bali (Debelius & Kuiter, 2007); Philippines: Batangas (Femia Marzo, 2014; present study) and Negros Oriental (Tillen, 2015); Taiwan: Penghu (Su et al., 2009); Japan: Izu Peninsula (Nakano, 2018); Australia: New South Wales (Mayes, 2020) and Queensland (Cobb, 2017; present study). Ecology: The species is found from 1 m (Gosliner et al., 2015, 2018) to up to 21 m depth (Femia Marzo, 2014). It has been found on red algae (Mayes, 2020) and under coral rubble (Gosliner et al., 2015, 2018). Murphydoris adusta feeds on compound tunicates (Gosliner et al., 2015, 2018). Etymology: From Latin adustus, perfect participle passive of aduro, brown. Murphydoris adusta refers to the dark brown colour of the body. External morphology (Figs 2A, B, 3A): Preserved specimens between 2 and 3 mm length. Body elongated and narrow. Foot tapered at posterior end, with slightly rounded edge at anterior part. Foot narrow, does not protrude from sides of notum. Notal border well developed, with thickened edge at anterior part around head, continuing with slightly serrated margin, supported by internal spicules. Gill located at posterior two-thirds of body. Three bipinnate gill branches arise from common base. Middle branch larger with wider and rounded tip (Fig. 3A). Dorsal or lateral papillae absent. Rhinophores non-retractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, lacking oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense network of spicules. Colour pattern (Fig. 2A, B): Body chocolate brown with patches between pale blue and iridescent white. Edge of notum same colour as patches with discontinuous pale-yellow coloration at border. Middle dorsal part of the body with transversal iridescent white line and yellow patch on each side. Anteriormost part of head iridescent white with chocolate brown band in middle. Rhinophores with same coloration as body. Gill branches translucent white with yellow tips, sometimes more striking on larger middle branch. Thin line of iridescent white arises from each gill base that join at beginning of tail, continuing with wide iridescent white dorsal band until end of tail. Yellowish pigmentation may be present. Foot blueish with small brown spots. Foregut anatomy (Figs 3B, 4A–C): Buccal bulb muscular. Dorsal buccal pump elongated posteriorly (Fig. 3B). Radular sac short, descending ventrally and expanding posteriorly. Thin oesophagus originates at buccal bulb behind buccal pump. Nervous system surrounds this area. Oesophagus becomes wider and continues with oesophageal pump. One small and rounded salivary gland on each side, at the junction of oesophagus and buccal pump (Fig. 3B). Labial cuticle surrounds lips and expands within buccal pump. Radular formula 11–13 × 1.0.1. Lateral tooth large and robust, with two upper rounded cusps (Fig. 4A, B). Thick masticatory margin. Masticatory margin with few, small, thin denticles (Fig. 4C). Base of the teeth wide and straight (Fig. 4A, B). Outer lateral teeth absent. Reproductive system (Figs 3C, 4D): Reproductive system located at anterior-third of body. Thin, elongate hermaphroditic duct begins at ovotestis, located inside digestive-hermaphroditic gland. Hermaphroditic duct expands into large, bean-shaped ampulla. Thin post-ampullary duct enters in female gland mass and divides into the prostatic portion of vas deferens and oviduct. Prostate large, wide and elongated, narrowing and continuing as long vas deferens, ending in penial sac at most distal part. Penial sac short and spherical. Penis armed. Penial spines hooked at base, long and thin at most distal part (Fig. 4D). Vagina wide and short, followed by rounded bursa copulatrix. From base of bursa arises a thin duct that connects with elongated receptaculum seminis. Thin and long uterine duct arises at distal part of vagina and enters in female gland mass. Remarks: Murphydoris adusta and M. singaporensis share brownish coloration (Sigurdsson, 1991; Swennen & Buatip, 2012), but M. singaporensis is translucent with reddish-brown blotches (Sigurdsson, 1991; Swennen & Buatip, 2012), while M. adusta has brown colour with pale blue, iridescent white and yellow in different parts of the body. Moreover, M. adusta has a prolonged edge at the anterior part of the notal border, whereas M. singaporensis lacks this (Sigurdsson, 1991; Swennen & Buatip, 2012). Internally, M. adusta has only one row of teeth per side, which have two cusps, while M. singaporensis has one inner and one outer lateral tooth and the inner teeth have only a single cusp (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Moreover, the denticles of the masticatory margin are bigger and more numerous in M. singaporensis (PazSedano et al., 2021b) than in M. adusta. Regarding the reproductive system, in M. singaporensis there is a duct arising from the base of the bursa copulatrix connecting with the uterine duct (Swennen & Buatip, 2012; PazSedano et al., 2021b), whereas in M. adusta the uterine duct begins at the middle of the vagina (Table 3).

MURPHYDORIS ADUSTA SP. NOV. (FIGS 2A, B, 3A–C, 4A–D) ‘ Goniodoris aspersa ’ – Debelius & Kuiter (2007: 28). ‘ Goniodoridella sp. ’ – Su et al. (2009: 457, fig. 11B). ‘ Goniodoridella sp. 6 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 6 ’ – Gosliner et al. (2018: 51). ‘ Goniodoridella sp. 3 ’ – Nakano (2018: 166). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: A6372BF6-E947-4470-9B1D-D22A9AB5FE58. Material examined: Holotype. QMMO 85985 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 29 September 2019), one preserved specimen, 3 mm long, dissected. Paratypes. MNCN 15.05 /200143 – MNCN 15.05 /200148, WAM S72660, QMMO 85986 – QMMO 85988; QMMO 85995 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb). MNCN 15.05 /200143 (29 September 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula). MNCN 15.05 /200144 (Fig. 3B, C) (29 September 2019), one preserved specimen 2 mm long, dissected (SEM: Radula). MNCN 15.05 /200145 (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula). MNCN 15.05 /200146 (28 November 2019), one preserved specimen, 5 mm long. MNCN 15.05 /200147 (28 November 2019), one preserved specimen, 5 mm long, dissected (SEM: Radula, penis). MNCN 15.05 /200148 (5 December 2019) one preserved specimen, 10 mm long, dissected (SEM: Radula). WAM S72660 (Figs 1, 2A) (2 May 2018), one preserved specimen, 5 mm long. QMMO 85986 (Fig. 4D) (2019), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle, penis). QMMO 85987 (2019), one preserved specimen, 3 mm long, dissected (SEM: Radula). QMMO 85988 (29 September 2019), two specimens. QMMO 85995 (Figs 2B, 4A, C) (11 December 2019), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). CASIZ 177775 (Philippines, Batangas Province, Tingloy, Maricaban Island, Aphol’s Rock, 13°45′41.74′′N, 121°23′0.56′′E; coll. T. M. Gosliner, 23 April 2008). CASIZ 186117 (Philippines, Batangas Province, Calumpan Peninsula, Mabini, Murals dive site, 13°41′56.472′′N, 120°52′57.719′′E; coll. D. Raymundo, 4 May 2011). CASIZ 181290 (Fig. 4B) (Philippines, Luzon, Batangas Province, Mabini, Mainit Bubbles dive site, 13°41.155′N, 120°53.713′E; coll. T. M. Gosliner, 21 May 2009), dissected (SEM: Radula). Distribution: Indian Ocean to Indonesia, western Pacific Ocean; Madagascar (Gosliner et al., 2015, 2018); Indonesia: Palau Sangeang (Anderson, 2021) and Bali (Debelius & Kuiter, 2007); Philippines: Batangas (Femia Marzo, 2014; present study) and Negros Oriental (Tillen, 2015); Taiwan: Penghu (Su et al., 2009); Japan: Izu Peninsula (Nakano, 2018); Australia: New South Wales (Mayes, 2020) and Queensland (Cobb, 2017; present study). Ecology: The species is found from 1 m (Gosliner et al., 2015, 2018) to up to 21 m depth (Femia Marzo, 2014). It has been found on red algae (Mayes, 2020) and under coral rubble (Gosliner et al., 2015, 2018). Murphydoris adusta feeds on compound tunicates (Gosliner et al., 2015, 2018). Etymology: From Latin adustus, perfect participle passive of aduro, brown. Murphydoris adusta refers to the dark brown colour of the body. External morphology (Figs 2A, B, 3A): Preserved specimens between 2 and 3 mm length. Body elongated and narrow. Foot tapered at posterior end, with slightly rounded edge at anterior part. Foot narrow, does not protrude from sides of notum. Notal border well developed, with thickened edge at anterior part around head, continuing with slightly serrated margin, supported by internal spicules. Gill located at posterior two-thirds of body. Three bipinnate gill branches arise from common base. Middle branch larger with wider and rounded tip (Fig. 3A). Dorsal or lateral papillae absent. Rhinophores non-retractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, lacking oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense network of spicules. Colour pattern (Fig. 2A, B): Body chocolate brown with patches between pale blue and iridescent white. Edge of notum same colour as patches with discontinuous pale-yellow coloration at border. Middle dorsal part of the body with transversal iridescent white line and yellow patch on each side. Anteriormost part of head iridescent white with chocolate brown band in middle. Rhinophores with same coloration as body. Gill branches translucent white with yellow tips, sometimes more striking on larger middle branch. Thin line of iridescent white arises from each gill base that join at beginning of tail, continuing with wide iridescent white dorsal band until end of tail. Yellowish pigmentation may be present. Foot blueish with small brown spots. Foregut anatomy (Figs 3B, 4A–C): Buccal bulb muscular. Dorsal buccal pump elongated posteriorly (Fig. 3B). Radular sac short, descending ventrally and expanding posteriorly. Thin oesophagus originates at buccal bulb behind buccal pump. Nervous system surrounds this area. Oesophagus becomes wider and continues with oesophageal pump. One small and rounded salivary gland on each side, at the junction of oesophagus and buccal pump (Fig. 3B). Labial cuticle surrounds lips and expands within buccal pump. Radular formula 11–13 × 1.0.1. Lateral tooth large and robust, with two upper rounded cusps (Fig. 4A, B). Thick masticatory margin. Masticatory margin with few, small, thin denticles (Fig. 4C). Base of the teeth wide and straight (Fig. 4A, B). Outer lateral teeth absent. Reproductive system (Figs 3C, 4D): Reproductive system located at anterior-third of body. Thin, elongate hermaphroditic duct begins at ovotestis, located inside digestive-hermaphroditic gland. Hermaphroditic duct expands into large, bean-shaped ampulla. Thin post-ampullary duct enters in female gland mass and divides into the prostatic portion of vas deferens and oviduct. Prostate large, wide and elongated, narrowing and continuing as long vas deferens, ending in penial sac at most distal part. Penial sac short and spherical. Penis armed. Penial spines hooked at base, long and thin at most distal part (Fig. 4D). Vagina wide and short, followed by rounded bursa copulatrix. From base of bursa arises a thin duct that connects with elongated receptaculum seminis. Thin and long uterine duct arises at distal part of vagina and enters in female gland mass. Remarks: Murphydoris adusta and M. singaporensis share brownish coloration (Sigurdsson, 1991; Swennen & Buatip, 2012), but M. singaporensis is translucent with reddish-brown blotches (Sigurdsson, 1991; Swennen & Buatip, 2012), while M. adusta has brown colour with pale blue, iridescent white and yellow in different parts of the body. Moreover, M. adusta has a prolonged edge at the anterior part of the notal border, whereas M. singaporensis lacks this (Sigurdsson, 1991; Swennen & Buatip, 2012). Internally, M. adusta has only one row of teeth per side, which have two cusps, while M. singaporensis has one inner and one outer lateral tooth and the inner teeth have only a single cusp (Swennen & Buatip, 2012; Paz-Sedano et al., 2021b). Moreover, the denticles of the masticatory margin are bigger and more numerous in M. singaporensis (PazSedano et al., 2021b) than in M. adusta. Regarding the reproductive system, in M. singaporensis there is a duct arising from the base of the bursa copulatrix connecting with the uterine duct (Swennen & Buatip, 2012; PazSedano et al., 2021b), whereas in M. adusta the uterine duct begins at the middle of the vagina (Table 3)., Published as part of Paz-Sedano, Sofía, Smirnoff, Dimitri, Candás, María, Gosliner, Terrence M. & Pola, Marta, 2022, Rediscovering the overlooked genus Murphydoris (Nudibranchia: Goniodorididae): the first phylogeny and addition of four new species, pp. 250-269 in Zoological Journal of the Linnean Society 196 on pages 254-259, DOI: 10.1093/zoolinnean/zlac002, http://zenodo.org/record/7037692, {"references":["Debelius H, Kuiter RH. 2007. Nudibranchs of the world. Frankfurt: IKAN.","Su Y, Huang LJ, Chang YW, Mok HK. 2009. Temporal changes in nudibranch composition at a coastal site off Penghu (the Pescadores) in the Taiwan Strait. Zoological Studies 48: 448 - 459.","Gosliner TM, Valdes A, Behrens D. 2015. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Gosliner TM, Valdes A, Behrens D. 2018. Nudibranch & sea slug identification - Indo-Pacific. Jacksonville, Florida: New World Publications.","Nakano R. 2018. Field guide to sea slugs and nudibranchs of Japan. Tokyo: Bun-ichi Co., Ltd.","Anderson J. 2021. Goniodoridella sp 6. In: Indonesia nudibranchs & sea slugs. Available at: http: // www. nudibranch. org / Indonesia % 20 Sea % 20 Slugs / html / nudibranchs / goniodoridella-sp 6 - 01. html","Femia Marzo G. 2014. Goniodoridella sp. 01 Genus: PruvotFol, 1933. In: Sea slugs of the Mediterranean Sea and elsewhere. Available at: http: // www. medslugs. de / E / Pac-W / Goniodoridella _ sp _ 01 / Goniodoridella _ sp _ 01 _ 02. htm","Tillen G. 2015. Goniodoridella sp. 03 Genus: Pruvot-Fol, 1933. In: Sea-slugs from the Philippine Islands. Available at: https: // www. philippine-sea-slugs. com / Nudibranchia / Doridina / Goniodoridella _ sp _ 03 _ 10. htm","Mayes B. 2020. Nudibranch (Goniodoridella sp). In: Brian Mayes (Flickr photostream). Available at: https: // www. flickr. com / photos / brianmayes / 50884781246 /","Cobb G. 2017. Goniodoridella sp 7. In: Nudibranchs of the Sunshine Coast, Queensland, Australia, Indo-Pacific. Available at: http: // nudibranch. com. au / pages / Cap-Goni-sp 7 - 009 w. htm","Sigurdsson JB. 1991. A nudibranch, Murphydoris singaporensis, new genus and species, from Singapore mangroves (Mollusca: Opisthobranchia: Goniodorididae). Raffles Bulletin of Zoology 39: 259 - 263.","Swennen C, Buatip S. 2012. Murphydoris singaporensis (Gastropoda: Nudibranchia: Goniodorididae) found in Thailand; with external and first internal description of the specimens. Strombus 19: 9 - 14.","Paz-Sedano S, Diaz-Agras G, Gosliner TM, Pola M. 2021 b. Revealing morphological characteristics of Goniodorididae genera (Mollusca: Nudibranchia). Organisms Biodiversity and Evolutionary Research. Available at: https: // link. springer. com / article / 10.1007 / s 13127 - 021 - 00508 - w citeas."]}

TRAPANIA DARVELLI RUDMAN, 1987 (FIGS 4G, 10) Trapania darvelli Rudman, 1987: 206−209, figs 12A, 15; Rudman & Darvell, 1990; Ono, 2004: 97, middle photograph; Debelius & Kuiter, 2007: 19, middletop photograph; Gosliner et al. 2008: 138, second photograph from the top; Humann & Loach, 2010: 307, middle-right photograph; Gosliner et al., 2015: 141, bottom-right photograph; Gosliner et al., 2018: 59, bottom-left photograph. Material examined: CASIZ 186129, one specimen, sequenced, Devil’s Point (13.65084°N 120.84145°E), Maricaban Island, Batangas Province, Philippines, 07 May 2011, collected by Peri Paleracio. CASIZ 197303, one specimen, sequenced, Verde Island Passage (13.91000°N 120.60000°E), Calatagan, Batangas Province, Luzon Island, Philippines, 16 May 2014, collected by Shayle Matsuda. CASIZ 202112, one specimen, sequenced, Verde Island Passage (13.91422°N 120.60643°E), Calatagan, Batangas Province, Luzon Island, Philippines, 09 May 2013, collected by T. M. Gosliner. CASIZ 222004, one specimen, dissected and sequenced, Balayan 294 D. S. SMIRNOFF ET AL. Bay (13.72000°N 120.87000°E), Mabini (Calumpan Peninsula), Batangas Province, Luzon Island, Philippines, 5–17 m depth, 28 March 2017, collected by William Mendoza. Geographical distribution: Known from the Western Pacific Ocean, including Hong Kong, the Philippines, the Solomon Islands, Vanuatu and the Ryukyu Islands, Japan (Rudman, 1987; Ono, 2004; Gosliner et al. 2008, 2015, 2018; Humann & DeLoach, 2010; present study). Description: The external morphology, jaws and radula have been previously well described (Rudman, 1987). We add an additional description of the buccal apparatus and the first description of the reproductive system. Buccal mass: The buccal mass is thick and muscular with a slightly elevated buccal pump. The jaws are well developed and contain several rows of curved, hook-shaped denticles (Fig. 10A). The radular formula is 32 × 1.0.1 (Fig. 10B). The older teeth (Fig. 9C) are markedly smaller than the newer ones and the radula widens gradually towards the more newly developed teeth (Fig. 10D). All of the teeth are slightly curved and narrow at the inner base and broaden out at the outer edges. The teeth bear numerous denticles on the inner side of the primary cusp that are similar but increase in length slightly nearer the primary cusp. There are approximately five to nine denticles on the inner side of the much larger primary cusp in the older teeth, and one short, triangular denticle on the outer side of the cusp. The newer teeth have 12–18 denticles on the inner side of the fairly broad primary cusp and there does not appear to be an outer denticle. Reproductive system: The mature reproductive system is triaulic (Fig. 4G). The narrow pre-ampullary duct enters the elongate, saccate ampulla at the proximal end of the ampulla. The ampulla is widest proximally and narrows again at the post-ampullary duct and divides into the short, narrow oviduct, which enters the female gland mass, and the vas deferens. The vas deferens gradually widens into the thick prostatic portion that loops prior to expanding distally, immediately before it narrows abruptly into the long, narrow ejaculatory segment. The ejaculatory segment widens into a short penial sac. The penial sac terminates adjacent to the vagina. The width of the vagina remains uniformly narrow along a long length until it enters the base of the large, spherical bursa copulatrix adjacent to the receptaculum duct. The long receptaculum duct enters the base of the smaller, smaller pyriform receptaculum seminis. The moderately long uterine duct emerges from the receptaculum duct near the base of the receptaculum and enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 293-294, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Rudman WB. 1987. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53: 189 - 212.","Rudman WB, Darvell BW. 1990. Opisthobranch molluscs of Hong Kong. I. Goniodorididae, Onchidrodididae, Triophidae, Gymnodorididae, Chormodorididae (Nudibranchia). Asian Marine Biology 7: 31 - 79.","Ono A. 2004. Opisthobranchs of Ryukyu Islands. Tokyo: Rutles.","Debelius H, Kuiter RH. 2007. Nudibranchs of the world. Frankfurt: IKAN-Unterwasserarchiv.","Gosliner TM, Behrens DW, Valdes A. 2015. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Humann P, DeLoach N. 2010. Reef creature identification: tropical Pacific. Jacksonville: New World Publications, Inc."]}

MURPHYDORIS PUNCTICULATA SP. NOV. (FIGS 2F, G, 3I–K, 6D–H) ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2008: 126). ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 1 ’ – Gosliner et al. (2018: 50). ‘ Goniodoridella sp. 11’ – Nakano (2018: 169). Z o o b a n k r e g i s t r a ti o n: u r n: l s i d: z o o b a n k. org:act: 8DB73463-292D-45AC-B0BB-EE3481B62319. Material examined: Holotype. WAM S72711 (Figs 1, 2F, 3J, K) (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb, 28 December 2018), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). Paratypes. MNCN 15.05 /200149– MNCN 15.05 /200150, WAM S72710, WAM S71781, WAM S71785, WAM S17182, WAM S72709, WAM S72089, WAM S72462 (Australia, Queensland, La Balsa Park, Mooloolah River, 26°41′S, 153°08′E, 3–6 m depth; coll. G. Cobb). MNCN 15.05 / 200149 (Fig. 6G) (11 December 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula, labial cuticle). MNCN 15.05 / 200150 (Fig. 6G) (2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S72710 (Fig. 2G) (15 December 2018), one preserved specimen, 3 mm long. WAM S71781 (May 2018), 1 preserved specimen, 4 mm long, dissected (SEM: Radula). WAM S71785 (Fig. 6D–F) (May 2018), one preserved specimen 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S17182 (May 2018), one preserved specimen, 3 mm long. WAM S72709 (15 December 2018), one preserved specimen, 5 mm long. WAM S72089 (20 November 2017), two preserved specimens. WAM S72462 (4 December 2016), one preserved specimen, 4 mm long, dissected (SEM: Radula, labial cuticle). CASIZ 222055 (Philippines, Visayas, Romblon Province, Alad Isand, Romblon Province, Visayas; coll. A. Fritts-Penniman, 7 April 2017), dissected (SEM: Radula). CASIZ 224656 (Philippines, Visayas, Romblon Province, Cobrador Island; col. K. L. Larkin, 19 March 2018). Distribution: Murphydoris puncticulata is found in the western and central Pacific Ocean (Gosliner et al., 2018), in Papua New Guinea, New Caledonia (Gosliner et al., 2008, 2015), the Philippines (Gosliner et al., 2018, present study), Hawaii (Gosliner et al., 2008, 2015; Pittman & Fiene, 2020), Japan: Shizuoka prefecture (Kakegawa, 2018) and Izu Island (Nakano, 2018), Indonesia: Bacan (Sali Bay Dive Resort, 2021) and Bali (Kakegawa, 2017) and Australia: Queensland (Cobb, 2020; present study). Ecology: The species has been found under coral rubble on reefs from the intertidal up to 10 m deep (Gosliner et al., 2018). Etymology: From Latin diminutive of punctum, point or dot. Murphydoris puncticulata is named after the brown notches formed by dots. External morphology (Figs 2F, G, 3I): Preserved specimens 2–7 mm length. Body limaciform and narrow. Foot acutely tapered at posterior end, with rounded edge at anterior part. Foot narrow, does not protrude from sides of notum. Notal border evident, reduced, continuing towards trilobate gill branches located at posterior part (Fig. 3I), same level of anal opening. Several small tubercles randomly located at sides of body. Tubercles supported by internal spicules. Dorsal or lateral papillae absent. Rhinophores nonretractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone without oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense net of spicules. Colour pattern (Fig. 2F, G): Body white with small dark brown spots scattered along body. One triangular band of concentrated brown spots at anterior portion of dorsum, behind rhinophores. Transverse dark brown patch at head, between rhinophores. Third horizontal band of concentrated spots behind gill. Rhinophores, gill branches and foot whitish translucent. Foregut anatomy (Figs 3J, 6D–G): Buccal bulb muscular. Dorsal buccal pump elongates posteriorly. Radular sac short descending ventrally. Thin oesophagus originating at buccal bulb, behind buccal pump. Nervous system surrounds this area. Oesophagus becoming wider and continuing to oesophageal pump. Small, rounded salivary gland present on each side, at junction of oesophagus and buccal pump (Fig. 3J). Labial cuticle surrounding lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 6D). Radular formula 13 × 1.1.0.1.1. Inner lateral tooth large and robust, with one cusp and masticatory margin bearing between five and seven sharped denticles (Fig. 6E, F). Base of these teeth wide and straight (Fig. 6E). Outer lateral tooth much smaller with two sharp cusps, the upper somewhat longer than the lower (Fig. 6G). Base of outer tooth wide and rectangular. Reproductive system (Figs 3K, 6H): Reproductive system located at anterior-third of body. Thin preampullary duct begins at ovotestis, located inside digestive-hermaphroditic gland. Pre-ampullary duct expands into large, bean-shaped ampulla. Thin postampullary duct arises from ampulla and enters in female gland mass. Inside female gland mass it divides into prostatic portion of vas deferens and oviduct. Prostate long and wide, becomes narrow and continuing as elongated vas deferens. Vas deferent as long as prostate, ends ending in a slightly wider penial sac in most distal part. Penis armed with small, hook-shaped penial spines at its base. Spines become longer, thinner and more pointed at most distal part of penis (Fig. 6H). Vagina short, as wide as vas deferens, connects with oval bursa copulatrix. From base of bursa copulatrix two thin ducts arise; one connects with small and elongated receptaculum seminis. Second duct corresponds to thin and long uterine duct, which enters in female gland mass. Remarks: Murphydoris puncticulata, M. cobbi and M. maracabranchia share having white body colour. Murphydoris puctulatum and M. cobbi have a hyaline white body with dark brown spots, one brown patch behind the rhinophores and one behind the gill. However, in M. cobbi this patch behind the rhinophores consists of a continuous line, while in M. puncticulata it is formed by several spots. Murphydoris maracabranchia has a brown line behind the rhinophores but lacks brown spots along the body. Moreover, in M. cobbi the gill is likely modified as two wing-shaped processes at the end of the notal border, while in M. maracabranchia the gill is narrow at the base with wider and rounded tip and M. puncutulatum has three distinguishable gill branches (Fig. 3). Murphydoris puncticulata has marked tubercles along the sides of the body that are absent in M. cobbi. With respect to the reproductive system, the uterine duct arises from the middle of the vagina in M. cobbi, from the middle of a duct between the bursa copulatrix and the receptaculum seminis in M. maracabranchia and near the base of the receptaculum seminis in M. puncticulata. Also, the receptaculum seminis and the bursa copulatrix are much larger in M. cobbi than in M. puncticulata. The receptaculum seminis is pyriform in M. cobbi, oval in M. maracabranchia and elongated in M. puncticulata (Table 3). Molecular results support M. puncticulata as different species (Fig. 1), having a COI p -distance of 10.7–11.2% and 19.4% with M. cobbi and M. maracabranchia, respectively; and a 16S p- distance of 13.2–14.5%, 7.9–8.2% and 18.9–19.2% with M. adusta, M. cobbi and M. maracabranchia, respectively (Table 2).

TRAPANIA JAPONICA (BABA, 1935) (FIGS 2A, 3, 4A) Drepania japonica Baba, 1935: 336–338, fig. 4. Trapania japonica (Baba, 1935) Rudman, 1987: 197– 201, figs 3C, 7A, 8, 9; Baba, 1990: 8–9, figs 1–3; Rudman & Darvell, 1990; Nakano, 2004: 93, top photograph; Gosliner et al., 2018: 63, middle-right photograph. Trapania norakhalafae Khalaf & Khalaf, 2017:7–30. Material examined: CASIZ 182896 A, two specimens, one sequenced, Mabini, Batangas Province, Luzon Island, Philippines, May 2010, collected by Peri Paleracio. CASIZ 222116, one specimen, dissected, Mabini, Batangas, Philippines, May 2017, collected by Peri Paleracio. Geographical distribution: Widespread in the IndoPacific: known from Japan (Baba, 1935, 1990; Nakano, 2004), Australia (Rudman, 1987), Hong Kong (Rudman & Darvell, 1990), the United Arab Emirates (Khalaf-Prinz & Khalaf, 2017) and the Philippines (present study). External morphology: Living animal 3–5 mm in length (Fig. 2A). The body is opaque white, randomly covered in at least eight medium and small circles of dark-to-light brown pigment. Extra-rhinophoral and extra-branchial appendages are pointed with bright yellow from tip to the base, which is opaque white. Rhinophores, six to eight lamellae and a pointed tip. Rhinophores are uniformly wide along their length and the same colour as the brown circles. Oral tentacles are typically elongate and completely coloured the same brown as the rest of the body. The anterior margins of the foot, which extend laterally as elongate appendages, are opaque white adorned halfway down its length with a circle similar to the body. The gill plume branches are pigmented brown interspersed with areas of opaque white. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of weakly developed jaws. The jaws contain weakly developed triangular elements and have less cuticularization compared to other Trapania (Fig. 3A). The radular formula of one specimen (CASIZ 222116) is 28–30 × 1.0.1. The older teeth are much smaller than the newer ones and the radula widens towards the more newly developed teeth. The morphology of the individual teeth is atypical of Trapania. The teeth (Fig. 3B) bear numerous denticles on either side of the primary large cusp, which is pointed and slightly curved. Externally to the large cusp lie seven to nine evenly small denticles that resemble the teeth of a hair comb. Internally to the large cusp lie five to six small triangularly shaped denticles, diminishing in size toward the medial line. The older teeth have fewer denticles than the more recently developed ones. The interior denticles are larger than those found external to the large cusp in the middle of the width of the tooth. Reproductive system: The mature reproductive system is triaulic (Fig. 4A). The narrow pre-ampullary duct enters the posterior end of the elongated cylindrical ampulla. The ampulla terminates where the post-ampullary duct divides into the oviduct and vas deferens.The short oviduct enters the female gland mass. The vas deferens is initially narrow, widening into a glandular prostatic portion as it loops over itself. This gradually transitions into the narrower, muscular ejaculatory portion that is continuous with the wide, straight, elongate penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The vagina is moderately long and uniformly narrow throughout its length and joins the large, spherical bursa copulatrix and short, narrow receptaculum duct. The receptaculum duct is extremely short and connects with pyriform receptaculum seminis near the division of the short uterine duct, which enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: Our molecular phylogeny, the 16S ABGD analysis and the bPTP analysis support Trapania japonica as a distinct species and reveal that it is part of a clade composed of T. scurra, T. palmula, T. tatsulok and T. kahel. The external anatomy and radula have been previously described (Baba, 1935; Rudman, 1987), but not the reproductive system. Rudman noted the unique nature of the radular teeth, with a single, large denticle near the middle of the tooth. Khalaf (2017) described T. norakhalafae from specimens noting the similarity of this species to Trapania sp. 13 in Gosliner et al. (2015). Khalaf also noted the similarity of T. norakhalafae to T. japonica, but did not elaborate on this. His description did not include any internal anatomy and described only the living animal and its coloration. Subsequently, Gosliner et al. (2018) concluded that the specimen they had previously identified as T. sp. 13 was indeed T. japonica and noted that T. norakhalafae was likely a synonym. In this study, we confirm that the specimens identified as this species by Gosliner et al. (2018), have radular morphology consistent with T. japonica. No aspects of the external morphology of T. norakhalafae are inconsistent with those described for T. japonica and we can find no basis for the separation of T. norakhalafae as a distinct species. Therefore, we consider T. norakhalafae to be a junior synonym of T. japonica., Published as part of Smirnoff, Dimitri S., Donohoo, Samantha A. & Gosliner, Terrence M., 2022, Extra-branchial processes manifest extra diversity: systematics of the genus Trapania (Nudibranchia: Goniodorididae) and nine new species descriptions, pp. 270-313 in Zoological Journal of the Linnean Society 196 on pages 279-281, DOI: 10.1093/zoolinnean/zlac009, http://zenodo.org/record/7037746, {"references":["Baba K. 1935. Nudibranchia of Mutsu Bay. Science Reports of Tohoku University 10: 331 - 360.","Rudman WB. 1987. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53: 189 - 212.","Baba K. 1990. Notes on the rare genera Trapania and Ancula from Japan with the description of a new species (Nudibranchia: Goniodorididae). Venus (Japanese Journal of Malacology) 49: 8 - 18.","Rudman WB, Darvell BW. 1990. Opisthobranch molluscs of Hong Kong. I. Goniodorididae, Onchidrodididae, Triophidae, Gymnodorididae, Chormodorididae (Nudibranchia). Asian Marine Biology 7: 31 - 79.","Nakano R. 2004. Opisthobranchs of Japan Islands. Tokyo: Rutles.","Gosliner TM, Valdes A, Behrens DW. 2018. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc.","Khalaf N, Khalaf O. 2017. Trapania norakhalafae Khalaf, 2017: a new dorid nudibranch sea slug species from the Dibba Sea, east coast of the United Arab Emirates, Gulf of Oman, Arabian Sea, Indian Ocean. Gazelle: The Palestinian Biological Bulletin 153: 7 - 30.","Gosliner TM, Behrens DW, Valdes A. 2015. Indo-Pacific nudibranchs and sea slugs: a field guide to the world's most diverse fauna. Jacksonville: New World Publications, Inc."]}

MURPHYDORIS MARACABRANCHIA SP. NOV. (FIGS. 2E, 3G, H, 6A–C) ‘ Goniodoridella sp. 4 ’ – Gosliner et al. (2015: 133). ‘ Goniodoridella sp. 4 ’ – Gosliner et al. (2018: 50). Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 79EC6092-71AA-4222-9E61-1728A47A142D. Material examined: Holotype. CASIZ 186120 (Figs 1, 2E) (Philippines, Batangas Province, Maracaban Island, Sepok, 13°41′16.98′′N, 120°49′37.884′′E; coll. P. Paleracio, 25 May, 2011), one preserved specimen, 3 mm long. Paratypes. CASIZ 222056, CASIZ 224657 (Philippines, Romblon, Romblon Province, Logbon Reef). CASIZ 222056. (coll. P. Eschweiler, 3 April 2017). CASIZ 224657 (Figs 3H, 6 A–C) (coll. T. M. Gosliner, 8 March 2018), dissected (SEM: Radula). CASIZ 222015 (Philippines, Romblon Province, Alad Island, 0.0– 0.5 m depth; coll. T. M. Gosliner, 7 April 2017). Distribution: Papua New Guinea (Anderson, 2015, Gosliner et al., 2015, 2018), Japan (Gosliner et al., 2015, 2018) and the Philippines: Romblon Island, Batangas (Kim, 2021, present study). Ecology: The species is found on the underside coral rubble (Gosliner etal., 2008, 2015, 2018), from intertidal (present study) to 14 m deep (Anderson, 2015). Etymology: Murphydoris maracabranchia is named after the shape of the gill (Latin branchia), having the tips wide and rounded, which resemble the musical percussion instrument called maracas (from Guaraní mbaracás). External morphology (Figs 2E, 3G): Preserved specimens between 3 and 5 mm length. Body limaciform and narrow. Foot tipped at end, rounded at anterior part, with small tubercles in dorsal part. Notal border well developed, reduced and serrated due to presence of spicules. Spicules form small tubercles on dorsal part of tail. Notal border ends in two lateral processes in back-posterior part of notum, one on each side of anal opening. Processes narrow at base, ending in widened, rounded tip. They are likely a modification of the branchial gill (Fig. 3G). Dorsal or lateral papillae absent. Rhinophores non-retractile and smooth, located at anterior part of body. Rhinophoral sheath absent. Mouth displaced towards ventral zone, without oral tentacles. Reproductive opening located halfway up right side. Whole body, including lateral processes, covered by dense network of spicules. Colour pattern (Fig. 2E): Body uniformly opaque white with tips of rhinophores red or orange. A thin maroon line surrounds body behind rhinophores. Foregut anatomy (Fig. 6A–C): Buccal bulb muscular. Dorsal buccal pump elongates backwards. Radular sac short descending ventrally. Thin oesophagus originates at posterior end of buccal bulb, behind buccal pump. Nervous system surrounds this area. Oesophageal pump and salivary glands were not observed due to the small size of specimens. Labial cuticle surrounds lips and expands within buccal pump. Radula could not be opened due to the small size. Estimated radular formula 13 × 1.1.0.1.1. Inner lateral tooth large and robust, with masticatory margin bearing denticles (Fig. 6A, B). Outer lateral tooth with two thin and sharp cusps, the upper somewhat longer than lower (Fig. 6C). Base of outer tooth wide and rectangular. Reproductive system (Fig. 3H): Reproductive system located at anterior-third of body. Thin hermaphroditic duct begins at ovotestis, located inside digestivehermaphroditic gland. Hermaphroditic duct expands in wide, pear-shaped ampulla. Thin post-ampullary duct arises from ampulla and divides into prostatic portion of vas deferens and oviduct. Oviduct enters in female gland mass. Prostate not differentiated, same width as ejaculatory portion. Thin and elongate vas deferens terminates in wide penial sac in most distal part. Vagina wider than vas deferens, connecting with small, rounded bursa copulatrix. Short duct arises from base of bursa copulatrix and divides into two thin ducts; one connects with small, oval receptaculum seminis. Receptaculum seminis slightly smaller than bursa copulatrix. Second duct corresponds to thin and short uterine duct, which enters in female gland mass. Remarks: Murphydoris maracabranchia is the second white species of Murphydoris described. Murphydoris maracabranchia and M. cobbi have a white body with a brown line behind the rhinophores. However, M. cobbi has small brown spots along the body, whereas M.maracabranchia lacksthese.Inaddition, therhinophores of M. maracabranchia have red to orange tips, while the rhinophores of M. cobbi are entirely white. The gill of M.maracabranchia is narrow at base with a wide, rounded tip, whereas M. singaporensis and M. adusta have several branches arising from same stalk and M. cobbi has wingshaped gill. Murphydoris maracabranchia also differs from other species of Murphydoris in the internal anatomy. It is the only species of Murphydoris without a differentiated prostate. Moreover, the uterine duct emerges in the middle of a duct between the bursa copulatrix and seminal receptacle in M. maracabranchia, whereas in M. adusta and M. cobbi the uterine duct emerges at distal part of the vagina and at the base of the receptaculum seminis in M. singaporensis (Table 3). Molecular analyses support the difference among species (Fig. 1). Moreover, the p -distance for COI between M. cobbi and M. maracabranchia is 21.4–21.9%. The p -distance for 16S is 18.0–18.6% between M. maracabranchia and M. adusta and 17.7–18.6% between M. maracabranchia and M. cobbi (Table 2).

TRAPANIA TAMARAW SP. NOV. (FIGS 12A, 4I, 13) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: F4449CAF-D81F-41CF-8FA2-FD8BE7F3710C. Trapania sp. 13 Gosliner et al., 2018: 62, middle-right photograph. Type material: Holotype: NMP 041334, originally CASIZ 208391, one specimen, dissected and sequenced, Puerto Galera (13.51350°N 120.95833°E), Mindoro, Mindoro Oriental Province, Philippines, 15–20 m depth, 28 March 2015, collected by P. J. Aristorenas. Type locality: Puerto Galera (13.51350°N 120.95833°E), Mindoro, Mindoro Oriental Province, the Philippines. Geographical distribution: Known only from Puerto Galera, the Philippines. Etymology: A tamaraw is the native name for the dwarfed hoofed buffalo endemic to Mindoro, Bubalus mindorensis (Heude, 1888). The horns of this buffalo resemble the well-developed and large jaws of this species compared to other members of the genus Trapania. External morphology: Live animal 4 mm (Fig. 12A). Live animal coloured with alternating blotches of dark brown/ black and yellowish tan across the entire body.Rhinophores are cylindrical halfway up and conical to a blunt tip the rest of the way with five to six lamellae. Top two-thirds of the rhinophores have a purple-brown appearance, while the bottom third has light coloration, similar to other parts of the body. Gill plume consists of three branches that have a translucent yellow-tan appearance with the same coloration as the rhinophores along the gill plume tips. Extra-branchial and extra-rhinophoral appendages are elongated and curved, typical of Trapania. Each is coloured yellowish tan with a single spot of purple/brown coloration halfway along their length. Digitiform oral tentacles are dark brown/black at their base and yellowish tan at their distal halves. The anterior margins of the foot extend laterally as elongate appendages are dark brown/black at their base and translucent yellowish at their distal halves. Buccal mass: The buccal mass is muscular with a moderately developed buccal pump on the dorsal surface. Inside the anterior portion of the mass is a pair of extremely well-developed jaws.The jaws (Fig. 13A) contain a single row of massive, acutely pointed, slightly curved, triangular jaw elements, equally spaced with regularly sized gaps (Fig. 13B). The radular formula of the holotype is 20 × 1.0.1 (Fig. 13C). The older teeth are much smaller than the newer ones (Fig. 13D) and the radula widens towards the more newly developed teeth. The teeth bear numerous denticles with the smallest ones being found on the inner edge of the tooth. There are approximately six to eight denticles on the inner side of the much larger, narrow primary cusp, and either no denticle on the outer side of the cusp or a series of up to eight minute, almost vestigial, short denticles on a few teeth. The denticles on the inner side of the primary cusp are generally larger nearer the primary cusp, but are not evenly graded. Reproductive system: The mature reproductive system is triaulic (Fig. 4I). The saccate ampulla is narrow at the distal end where it connects to the ovotestis. From there, it abruptly widens before leading to a point where it divides into the short oviduct and vas deferens. The short, oviduct enters the female gland mass. The vas deferens gradually widens into a curved, thick prostatic portion and gradually transitions into the narrow, muscular ejaculatory portion that is continuous with the narrow, moderately short penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The narrow vagina maintains its width from its opening through a long region prior to its junction with the large, ovoid bursa copulatrix and short narrow receptaculum duct. The receptaculum duct joins the smaller spherical receptaculum seminis near the long division of the short uterine duct, which enters the female gland mass. The female gland mass is composed of the large mucous gland and the smaller albumen and membrane glands. Remarks: This species is compared to the four other members of its species complex below in the remarks following Trapania kamagong.