taxonID	type	description	language	source
0391879409774F6EFC65FA9AFBCCEBCF.taxon	materials_examined	Type species: Murphydoris singaporensis Sigurdson, 1991 by monotypy.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	description	(FIGS 2 A, B, 3 A – C, 4 A – D)	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	description	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: A 6372 BF 6 - E 947 - 4470 - 9 B 1 D-D 22 A 9 AB 5 FE 58.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	materials_examined	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 S 72660, 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. 3 B, 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 S 72660 (Figs 1, 2 A) (2 May 2018), one preserved specimen, 5 mm long. QMMO 85986 (Fig. 4 D) (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 2 B, 4 A, 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. 4 B) (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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	distribution	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	biology_ecology	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	etymology	Etymology: From Latin adustus, perfect participle passive of aduro, brown. Murphydoris adusta refers to the dark brown colour of the body. External morphology (Figs 2 A, B, 3 A): 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. 3 A). 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.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	description	Colour pattern (Fig. 2 A, 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 3 B, 4 A – C): Buccal bulb muscular. Dorsal buccal pump elongated posteriorly (Fig. 3 B). 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. 3 B). 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. 4 A, B). Thick masticatory margin. Masticatory margin with few, small, thin denticles (Fig. 4 C). Base of the teeth wide and straight (Fig. 4 A, B). Outer lateral teeth absent. Reproductive system (Figs 3 C, 4 D): 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. 4 D). 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.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409774F63FC52FA06FD71EA31.taxon	discussion	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., 2021 b). Moreover, the denticles of the masticatory margin are bigger and more numerous in M. singaporensis (PazSedano et al., 2021 b) 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., 2021 b), whereas in M. adusta the uterine duct begins at the middle of the vagina (Table 3).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	description	(FIGS 2 C, D, 3 D – F, 4 E – H, 5)	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	materials_examined	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. 2 D) (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. 3 F) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85983 (Figs 3 E, 4 F – H) (29 September 2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). QMMO 85984 (Fig. 4 E) (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, 2 C) (Philippines, Luzon, Batangas Province, Lobo, Malabrigo, 13 ° 36 ′ N, 121 ° 15 ′ E; coll. P. J. Aristorenas, 15 April 2015).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	distribution	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	biology_ecology	Ecology: Murphydoris cobbi is found in shallow sandy slopes among coral rubble (Gosliner et al., 2015).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	etymology	Etymology: Murphydoris cobbi is named after Gary Cobb, who collected, photographed and made numerous comments on the specimens studied, which greatly facilitated its description.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	description	External morphology (Figs 2 C, D, 3 D): 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. 3 D). 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. 2 C, 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 3 E, 4 E – G, 5 A): 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. 3 E). Oesophagus continues and inserts into digestive-hermaphroditic gland (Fig. 5 A). Oesophagus slightly wider at middle part, but oesophageal pump not differentiated (Fig. 5 A). 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. 5 A). Labial cuticle surrounds lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 4 E). 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. 4 F, G). Base of these teeth wide and straight (Fig. 4 F, G). Outer lateral tooth much smaller, hook-shaped, with two sharp cusps. Upper cusp somewhat longer than lower (Fig. 4 G). Base of outer tooth wide, robust and elongated. Reproductive system (Figs 3 F, 4 H, 5 B): 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. 4 H). 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.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097A4F60FEF2FB82FBC3EB8F.taxon	discussion	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., 2021 b), 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., 2021 b). 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., 2021 b). 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., 2021 b). 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 16 S (Table 2).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	description	(FIGS. 2 E, 3 G, H, 6 A – C)	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	description	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: 79 EC 6092 - 71 AA- 4222 - 9 E 61 - 1728 A 47 A 142 D.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	materials_examined	Material examined: Holotype. CASIZ 186120 (Figs 1, 2 E) (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 3 H, 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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	distribution	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	biology_ecology	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	etymology	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	description	External morphology (Figs 2 E, 3 G): 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. 3 G). 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. 2 E): Body uniformly opaque white with tips of rhinophores red or orange. A thin maroon line surrounds body behind rhinophores. Foregut anatomy (Fig. 6 A – 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. 6 A, B). Outer lateral tooth with two thin and sharp cusps, the upper somewhat longer than lower (Fig. 6 C). Base of outer tooth wide and rectangular. Reproductive system (Fig. 3 H): 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.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
0391879409794F66FC93FA4BFD02EBE3.taxon	discussion	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 16 S is 18.0 – 18.6 % between M. maracabranchia and M. adusta and 17.7 – 18.6 % between M. maracabranchia and M. cobbi (Table 2).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	description	(FIGS 2 F, G, 3 I – K, 6 D – H)	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	materials_examined	Material examined: Holotype. WAM S 72711 (Figs 1, 2 F, 3 J, 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 S 72710, WAM S 71781, WAM S 71785, WAM S 17182, WAM S 72709, WAM S 72089, WAM S 72462 (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. 6 G) (11 December 2019), one preserved specimen, 2 mm long, dissected (SEM: Radula, labial cuticle). MNCN 15.05 / 200150 (Fig. 6 G) (2019), one preserved specimen, 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S 72710 (Fig. 2 G) (15 December 2018), one preserved specimen, 3 mm long. WAM S 71781 (May 2018), 1 preserved specimen, 4 mm long, dissected (SEM: Radula). WAM S 71785 (Fig. 6 D – F) (May 2018), one preserved specimen 3 mm long, dissected (SEM: Radula, labial cuticle). WAM S 17182 (May 2018), one preserved specimen, 3 mm long. WAM S 72709 (15 December 2018), one preserved specimen, 5 mm long. WAM S 72089 (20 November 2017), two preserved specimens. WAM S 72462 (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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	distribution	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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	biology_ecology	Ecology: The species has been found under coral rubble on reefs from the intertidal up to 10 m deep (Gosliner et al., 2018).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	etymology	Etymology: From Latin diminutive of punctum, point or dot. Murphydoris puncticulata is named after the brown notches formed by dots.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	description	External morphology (Figs 2 F, G, 3 I): 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. 3 I), 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. 2 F, 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 3 J, 6 D – 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. 3 J). Labial cuticle surrounding lips and expands within buccal pump. It appears as a network similar to a honeycomb on inside buccal pump (Fig. 6 D). 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. 6 E, F). Base of these teeth wide and straight (Fig. 6 E). Outer lateral tooth much smaller with two sharp cusps, the upper somewhat longer than the lower (Fig. 6 G). Base of outer tooth wide and rectangular. Reproductive system (Figs 3 K, 6 H): 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. 6 H). 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.	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
03918794097F4F65FF0EF9D7FBE0EB0A.taxon	discussion	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 16 S 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).	en	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. Zoological Journal of the Linnean Society 196: 250-269, DOI: 10.1093/zoolinnean/zlac002
