taxonID	type	description	language	source
03E487E4FF91FB26EBF4FB77FF22F84E.taxon	materials_examined	Type species: Trapania fusca (Lafont, 1874), by typification of replaced name. Species treatments are arranged by clades in the order that species are listed in the phylogeny in Figure 1.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF91FB22EA00FA80FECAF908.taxon	description	(FIGS 2 A, 3, 4 A)	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF91FB22EA00FA80FECAF908.taxon	materials_examined	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. 2 A). 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. 3 A). 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. 3 B) 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. 4 A). 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 16 S 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF95FB3FEA11FB32FEA7FB45.taxon	description	(FIGS 2 B, 4 B, 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 - 45 B 7 D 23 B 188 D.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF95FB3FEA11FB32FEA7FB45.taxon	materials_examined	Type locality: Verde Island Passage (13.92816 ºN 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. 2 B) 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. 5 A). The jaw surface is covered with small semi-spheres, each covered with numerous short and blunt spikes resembling a medieval flail or mace (Fig. 5 B). The radular formula of the holotype is 25 × 1.0.1 (Fig. 5 C) 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. 5 C). 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. 5 D). Reproductive system: The mature reproductive system is triaulic (Fig. 4 B). 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 16 S 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. 8 a, fig. 10 a; Fig. 5 B). 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. 8 b – d, fig. 10 b – d; Fig. 5 D). 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. 4 A) 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF88FB3DEA39F96AFDDAFA24.taxon	description	(FIGS 1 C, 4 C, 6) 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: DEBA 67 C 6 - F 65 D- 418 D- 8 AE 6 - 40063 C 09 F 594.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF88FB3DEA39F96AFDDAFA24.taxon	materials_examined	Type material: Holotype: NMP 041330, originally CASIZ 186131, one specimen, dissected, Matotonngil Point (13.75528 ° N 120.90672 ° E), Mabini, Batangas Province, Luzon Island, Philippines, 3 May 2011, collected by Alicia Hermosillo. Paratype: CASIZ 186133, one specimen, dissected and sequenced, Matotonngil Point (13.75528 ° N 120.90672 ° E), Mabini, Batangas Province, Luzon Island, Philippines, 8 March 2011, collected by Alicia Hermosillo and Peri Paleracio. Type locality: Matotonngil Point (13.75528 ° N 120.90672 ° E), Mabini, Batangas Province, Luzon Island, the Philippines. Geographical distribution: Known only from Matotongil Point, the Philippines. Natural history: Feeds on an orange, encrusting bryozoan on small rocks surrounded by coarse sandy areas. Etymology: Kahel is theTagalog word for orange and refers to the encrusting, orange bryozoan on which this species has been observed feeding. It is a noun in apposition. External morphology: Living specimens are 5 – 6 mm (Fig. 2 C). They have an elongated body typical of Trapania, with an orange-red body coloration. Foot has a dorsal ridge coloured a dark brown / black running all the way to the posterior tip. The same intensity of dark pigmentation covers the three gill branches, the oral tentacles and the top two-thirds of the rhinophores, each with seven lamellae. The extra-branchial and extra-rhinophoral appendages have a slight curve typical of Trapania and are both coloured a bright opaque white. While the mantle between the extra-branchial appendages is the same orange-brown as the rest of the body, a continuous band of bright solid white runs tip-to-tip between the extra-rhinophoral appendages, creating the appearance of a white ribbon. The same white coloration extends up the first third of the rhinophores. Rhinophores are spearshaped with about ten lamellae and a pointed tip. The basal half of the rhinophores is white with a dark-brown to black apex. The oral tentacles are thin and elongate covered with black pigment along their entire length. The anterior margins of the foot extend laterally as elongate appendages are more colourless and are more translucent than the rest of the body. 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. 6 A). In the image, a membrane partially covers the elements, but the shape of the thin triangular elements is still visible. The radular formula of the holotype is 16 × 1.0.1 (Fig. 6 B). The teeth are all highly arched. The newer teeth (Fig. 6 C) are slightly larger than the older teeth (Fig. 6 D) and the radula widens somewhat 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 seven denticles on the inner side of the much larger primary cusp and one to three denticles on the outer side of the cusp. All of the denticles other than primary cusp are similar in size. The older teeth have a more pronounced primary cusp, but the smaller denticles are consistent in size across all teeth. Reproductive system: The mature reproductive system is triaulic (Fig. 4 C). The narrow pre-ampullary duct enters the saccate ampulla at its base, adjacent to the post-ampullary duct. The post-ampullary duct divides into the short oviduct and elongate vas deferens. The oviduct enters the female gland mass. The vas deferens is equally sized throughout its length and forms a loop that continues into a convoluted, glandular prostatic portion. This gradually transitions into the slightly wider, 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 large, spherical bursa copulatrix and short narrow receptaculum duct. The receptaculum duct joins the 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: In our molecular phylogeny, Trapania kahel is moderately supported as sister to T. japonica (PP = 0.94, BS = 54) in a clade composed of T. scurra, T. palmula, T. japonica and T. tatsulok. The ABGD analysis and the bPTP analysis reveal a strong genetic divergence of 14.9 % in the COI gene and 13.4 − 13.9 % in the 16 S gene between T. kahel and T. scurra and 15.1 % / 13.7 − 14.0 % between T. kahel and T. palmula (Tables 3, 4). The bright orange colour with white extra-branchial and extra-rhinophoral appendages also help distinguish this species from all described members of Trapania. No members of this clade have an orange body colour. Internally, T. kahel is similar to other members of the clade that have jaws with acutely pointed jaw elements but differs from T. tatsulok, which has jaw elements with numerous pointed spikes. As in T. scurra, T. palmula and T. japonica, the radular teeth have only a single prominent denticle, but have narrower teeth with fewer, coarser denticles. The vagina of T. kahel is much shorter than in other members of the clade and the male duct is elongate with little distinction or narrowing between the prostatic and ejaculatory portions of the duct. The other members of this clade have a distinctly narrower portion of the ejaculatory duct.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8AFB3BE8AFFF30FCD0F901.taxon	description	(FIGS 2 D, E, 4 D, 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: 172793 CC-C 647 - 44 F 6 - 8926 - 8 AB 7593490 B 2.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8AFB3BE8AFFF30FCD0F901.taxon	materials_examined	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. 2 D, 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. 7 A) 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. 7 B) are much smaller than the newer ones and the radula widens towards the more newly developed teeth (Fig. 7 C, 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. 4 D). 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 16 S 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8CFB3AEA12FB44FAA6FBDF.taxon	description	(FIGS 2 F, 4 E, 8)	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8CFB3AEA12FB44FAA6FBDF.taxon	description	Geographical distribution: Known from Hong Kong (Rudman, 1987; Rudman & Darvell, 1990; Ono, 2004), New Caledonia (Hervé, 2010) and the Philippines (present study). External morphology: The living animal is 6 mm (Fig. 2 F). The body coloration is opaque white with a powdery white appearance. The rhinophores are evenly wide throughout their length and have six lamellae and a pointed tip. They are translucent white at their base, opaque white along the lamellae and reddish orange at their tip and a bit along the front edge. The three gill plume branches are bipinnate and translucent, except for minor reddishorange fringing along the tips of the main branches. Both extra-rhinophoral and extra-branchial appendages are stocky with the same reddish orange coloration along the edges of their tips. The anterior margins of the foot extend laterally as elongate appendages and are more translucent than the rest of the body. Reddish orange coloration adorns their tips, as it does the tips of the elongate, digitiform oral tentacles. The dorsal ridge of the most posterior portion of the foot is tinted reddish orange. 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 at least two rows of acutely pointed jaw elements of various sizes that are tightly packed together with a few gaps between them (Fig. 8 A). The radular formula is 26 × 1.0.1 (Fig. 8 B). The older teeth (Fig. 8 C) are much smaller than the newer ones and the radula widens gradually towards the more newly developed teeth (Fig. 8 D). The base of the teeth is wide and expanded. The teeth bear numerous acutely pointed denticles with the smallest ones being found on the inner edge of the tooth. There are approximately nine to 12 denticles on the inner side of the much larger primary cusp and one wide, triangular denticle on the outer side of the cusp. The older teeth have fewer denticles than the more recently developed ones and the denticles appear more uniformly graded from smaller to large denticles in the more newly developed teeth. Reproductive system: The mature reproductive system is triaulic (Fig. 4 E). The narrow pre-ampullary duct enters the saccate ampulla near the distal end of the ampulla. The ampulla narrows again at the postampullary 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 narrowing briefly. Here it becomes a long smooth ejaculatory segment which transitions into a 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, spherical bursa copulatrix adjacent to the receptaculum duct. The receptaculum duct curves and enters the base of the smaller, slightly pyriform 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: The reproductive anatomy of T. aurata is described here for the first time. Following the publication of Gosliner & Fahey (2008), Rudman (2008) discussed difficulties in distinguishing T. aurata from T. vitta. Our molecular phylogeny, the ABGD analysis and the bPTP analysis suggest that these two species are sister to one another with a strong genetic divergence of 16.3 – 17.8 % in the COI gene and 6.8 − 7.3 % in the 16 S gene (Tables 3, 4). Due to the unavailability of additional specimens, T. aurata intraspecific variation was not studied here, but T. vitta has minimal intraspecific variation (0.0 – 0.8 %) between four specimens collected in Palau and the Philippines. There are also consistent external and internal morphological features that permit the distinction between the two species. Externally, T. aurata has orange lines along the edges of the extra-branchial and extra-rhinophoral appendages that are absent in T. vitta. Rudman noted variation in the other orange markings on the head, gill and foot between the two species, but the pigment differences on the extrabranchial and extra-rhinophoral appendages remains consistent. Internally, Gosliner & Fahey (2008) noted radular differences between the width of the outermost denticle of the radular tooth. With the addition of the material studied here, other radular differences are evident. The teeth of T. aurata are more sharply curved than those of T. vitta (Gosliner & Fahey, 2008: fig. 20 C, D). Additionally, the denticles on the inside of the primary denticle are more elongated in T. aurata and much shorter and evenly graded in size in T. vitta. The reproductive systems of the two are extremely similar with the possible exception that the prostatic portion of the vas deferens appears slightly more convoluted in T. aurata (Gosliner & Fahey, 2008: fig. 21; Fig. 4 E).	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8DFB38E886F910FA85F947.taxon	description	(FIGS 2 G, 4 F, 9)	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8DFB38E886F910FA85F947.taxon	materials_examined	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 2 G 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. 9 A). The radular formula is 34 × 1.0.1 (Fig. 9 B). The older teeth (Fig. 9 C) are somewhat smaller than the newer ones and the radula widens gradually towards the more newly developed teeth (Fig. 9 D). 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. 4 F). 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 16 S 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 16 S 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. 10 A – C; Rudman, 1987: fig. 10 A – 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. 4 F) differs in several key features from that of T. darvelli (Fig. 4 G). 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8FFB36E897FB79FDB2FE32.taxon	description	(FIGS 4 G, 10)	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF8FFB36E897FB79FDB2FE32.taxon	materials_examined	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. 10 A). The radular formula is 32 × 1.0.1 (Fig. 10 B). The older teeth (Fig. 9 C) are markedly smaller than the newer ones and the radula widens gradually towards the more newly developed teeth (Fig. 10 D). 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. 4 G). 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF81FB35EA20FC6EFB56F888.taxon	description	(FIGS 2 H, 4 H, 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: 3295 F 8 AD- 8 D 24 - 4204 - A 98 E- 8 F 6910 B 14367.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF81FB35EA20FC6EFB56F888.taxon	materials_examined	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. 2 H) 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. 11 A). The radular formula of the holotype is 50 × 1.0.1 (Fig. 11 B). 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. 11 C) have fewer denticles than the more recently developed ones (Fig. 11 D). 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. 4 H). 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 16 S 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF82FB34E8C6F9C8FB5EFFAF.taxon	description	(FIGS 12 A, 4 I, 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: F 4449 CAF-D 81 F- 41 CF- 8 FA 2 - FD 8 BE 7 F 3710 C.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF82FB34E8C6F9C8FB5EFFAF.taxon	materials_examined	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. 12 A). 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. 13 A) contain a single row of massive, acutely pointed, slightly curved, triangular jaw elements, equally spaced with regularly sized gaps (Fig. 13 B). The radular formula of the holotype is 20 × 1.0.1 (Fig. 13 C). The older teeth are much smaller than the newer ones (Fig. 13 D) 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. 4 I). 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF83FB31E8ADFCE6FE27F8BD.taxon	description	(FIGS 4 J, 12 B, 14) 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 5044129 - BF 78 - 4375 - 9 F 4 D- 2 A 941 FD 13 AF 6.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FF83FB31E8ADFCE6FE27F8BD.taxon	materials_examined	Type material: Holotype: NMP 041332, originally CASIZ 186206, one specimen, dissected and sequenced, Anilao Harbor (13.75993 ° N 120.92617 ° E), Balayan Bay, Batangas Province, Luzon Island, Philippines, 04 May 2011, collected by Alexis Principe. Ty p e l o c a l i t y: A n i l a o H a r b o r (1 3.7 5 9 9 3 ° N 120.92617 ° E), Balayan Bay, Batangas Province, Luzon Island, the Philippines. Geographic distribution: Known from the Philippines (present study) and Indonesia (Debelius & Kuiter, 2007). Etymology: This species is named for Stegodon luzonensis von Koenigwald 1956, a Pleistocene megafauna with ivory tusks that roamed Luzon, Mindanao and other Philippine islands. It refers to the white-tipped extra-rhinophoral and extrabranchial appendages. It is a noun in apposition. External morphology: Living animal (Fig. 12 B) 15 mm. It has an opaque grey body covered dorsally with speckled red-brown colour characterized by small and large blotches of missing pigmentation. Notable blotches reside laterally below the rhinophores, below the gill plume and at the posterior portion of the foot. Extra-rhinophoral and extra-brachial appendages have red brown at base but are light yellow to the tip. Rhinophores, eight to ten lamellae and a pointed tip, are uniformly wide along their length and the same colour as the red-brown body, except for the grey tip. Oral tentacles are coloured at their base and opaque grey 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 that are 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 several rows of acutely pointed and stubby jaw elements of various sizes that are tightly packed together with a few gaps between them (Fig. 14 A). The radular formula of the holotype is 44 – 45 × 1.0.1 (Fig. 14 B). The older teeth (Fig. 14 C) are much smaller than the newer ones (Fig. 14 D) and the radula widens gradually towards the more newly developed teeth. The teeth bear numerous denticles of irregular length, but with the smallest ones being found on the inner edge of the tooth. There are approximately nine to 16 denticles on the inner side of the much larger, narrow primary cusp, and one to two small 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. 4 J). The narrow pre-ampullary duct enters the saccate ampulla near the middle of its length. The ampulla narrows again at the postampullary 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, curved ejaculatory segment, which again widens into the relatively short, bulbous penial sac. The penial sac terminates adjacent to the vagina. The vagina is relatively short and wide throughout its entire length and 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 below in the remarks following Trapania kamagong.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFB8FB0CE8BDFAA4FD09FF1B.taxon	description	(FIGS 12 D, 16 B, 17) 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: E 8973 F 5 A- 119 C- 487 B- 9538 - 74 B 21 D 15 ACC 6.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFB8FB0CE8BDFAA4FD09FF1B.taxon	materials_examined	Type material: Holotype: NMP 041336, originally CASIZ 208585 A, one specimen, dissected and sequenced, Puerto Galera (13.51350 ° N 120.95833 ° E), Mindoro, Mindoro Oriental Province, Philippines, 6 – 20 m depth, 29 March 2015, collected by P. J. Aristorenas. Paratype: CASIZ 177526, one specimen, sequenced, Mainit Point (13.68640 ° N 120.895413 ° E), Mabini (Calumpan Peninsula), Batangas Province, Luzon Island, Philippines, 22.7 m depth, 21 March 2008, collected by T. M. Gosliner, A. Valdes, M. Pola, L. Witzel, B. Moore and A. Alejandrino. Ty p e l o c a l i t y: P u e r t o G a l e r a (1 3. 5 1 3 5 0 ° N 120.95833 ° E), Mindoro, Mindoro Oriental Province, the Philippines. Geographical distribution: Known only from the Philippines. Etymology: Kamagong is the Tagalog name for the velvet apple, Diospyros discolor Willd. (Ebenaceae), a species of persimmon tree found in the Philippines and Taiwan. It refers to the distinguishing dark-tipped, extra-rhinophoral and extra-branchial appendages of this species. External morphology: Living animal 15 mm (Fig. 12 D). It has an opaque grey body covered dorsally with speckled red-brown colour characterized by small and large blotches of missing pigmentation. Notable blotches reside laterally below the rhinophores, below the gill plum and at the posterior portion of the foot. Extra-rhinophoral and extra-branchial appendages lack coloration from base to the tip, which has the same red-brown coloration as the body. Rhinophores, eight to ten lamellae and a pointed tip, are uniformly wide along their length and the same colour as the red-brown body, except for grey tip. Oral tentacles are coloured at their base and opaque grey 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 plum is prominent and a lighter brown than 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 a row of acutely pointed and stubby jaw elements of various sizes that are tightly packed together with a few gaps between them (Fig. 17 A). The radular formula of the holotype is 23 × 1.0.1. The older teeth are much smaller (Fig. 17 B) than the newer ones (Figs. 17 C, D) and the radula widens gradually 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 four to 15 denticles on the inner side of the much larger primary cusp, and one to two 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. 16 B). The cylindrical ampulla is narrow and short from the distal end where it connects to the ovotestis to where it merges into post-ampullary duct. The post-ampullary divides at its base into a short oviduct and vas deferens. The oviduct enters the female gland mass. The vas deferens is initially narrow and widens into a convoluted, glandular prostatic portion with multiple turns before abruptly transitioning into the narrow, muscular ejaculatory portion that is continuous with the short, bulbous penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The vagina is equally narrow at its opening until its junction with the large, spherical bursa copulatrix. A narrow receptaculum duct leads from the bursa copulatrix to the spherical receptaculum at the 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: Our molecular phylogeny moderately (BI) or weakly (ML) places Trapania tamaraw at the base of a clade that includes a well-supported clade with T. stegodon, T. sp. ‘ Anilao’ and T. kamagong. However, our ABGD and bPTP analyses support T. tamaraw as a distinct species due to a strong divergence of 15.8 % in the COI gene between T. tamaraw and T. stegodon, 14.8 % / 7.6 − 7.8 % in the COI / 16 S genes between T. tamaraw and T. kamagong, and 13.6 % / 7.9 % between T. tamaraw and T. sp. ‘ Anilao’ (Tables 3, 4). The closest related species to T. stegodon include T. sp. ‘ Anilao’ and T. kamagong; which have a smaller COI genetic divergence of 6.3 % between T. stegodon and T. sp. ‘ Anilao’ and 6.6 – 7.0 % between T. stegodon and T. kamagong (Table 3). This species also has a distinctive patchy colour pattern of creamy yellowish to tan interspersed with dark-brown blotches. The colour pattern is similar to that of T. euryeia (see: Gosliner & Fahey, 2008) and Trapania toddi Rudman, 1987, with the exception that both of these species have lighter punctations in the dark blotches that are absent in T. tamaraw. Also, T. tamaraw is a member of a different clade than T. euryeia studied here. Internally, T. tamaraw has huge jaws, as compared to those found in T. euryeia. The reproductive system of T. tamaraw differs from that of T. euryeia and T. toddi. The pre-ampullary duct of T. tamaraw (Fig. 4 I) enters the distal end of the ampulla, whereas it enters subdistally in T. toddi (Rudman, 1987: fig. 4 D) and proximally in T. euryeia (Gosliner & Fahey, 2008: fig. 19). Also, T. euryeia has a vaginal collar (Gosliner & Fahey, 2008: fig. 19) that appears to be glandular and is absent in T. toddi and T. tamaraw. Trapania tamaraw, T. stegodon, T. sp. ‘ Anilao’ and T. kamagong form a weakly supported clade that represents a species complex. When T. tamaraw is compared to the other members of the clade in which it is placed, there are significant distinctive morphological characteristics for the four species in this clade. With respect to external coloration, T. tamaraw is smaller (about 4 mm, whereas the other three species are approximately 15 mm in length) and is unique in having well-separated dark-brown and yellowish patches and lacks the opaque white spotting found in T. stegodon, T. kamagong and T. sp. ‘ Anilao’. In T. stegodon, T. kamagong and T. sp. ‘ Anilao’, the brownish body pigmented areas are interrupted by small, irregularly shaped white patches. The primary external differences between T. stegodon, T. kamagong and T. sp. ‘ Anilao’ are in the pigment pattern found on the extra-rhinophoral and extra-branchial appendages. In T. stegodon, the extra-rhinophoral and extra-branchial appendages are uniformly whitish yellow, whereas the appendages of T. kamagong are the same colour but have a brown apex. The preserved pigmentation pattern of T. sp. ‘ Anilao’ are a uniform continuation of the body coloration with the exception of medial whitish transverse band and a white apex. The jaws of the four species also differ. In T. tamaraw, there are massive jaws that bear a single row of large, acutely-pointed denticles along the masticatory margin. In T. stegodon, the jaws bear several rows or acutely pointed rodlets over much of the surface of the jaws; in T. sp. ‘ Anilao’ there is a single row of rounded rodlets along the margin; in T. kamagong, there are two to three rows of rodlets on the margin of the jaws. Additional radular characters also separate the four members of this species complex. The radular teeth of T. tamaraw are more strongly arched along their anterior surface than in the other three species. Also, T. tamaraw has much reduced denticles on the outer side of the primary cusp, where the denticles are either entirely absent or are present as a series of up to eight minute denticles. Both T. stegodon and T. kamagong have denticles of irregular lengths on the inner side of the primary cusp, whereas they are more similar in size in T. sp. ‘ Anilao’. In T. stegodon the primary cusp is narrower and straighter than that of T. kamagong, where the primary cusp is broader at the base and is more curved inwardly. The reproductive systems of the four species forming this species complex bear significant differences. In T. tamaraw (Fig. 4 I), the pre-ampullary duct enters the distal end of the ampulla, while it enters far more proximally in T. stegodon (Fig. 4 J), T. sp. ‘ Anilao’ (Fig. 16 A) and T. kamagong (Fig. 16 B). In T. tamaraw the prostatic vas deferens does not narrow into a distinct ejaculatory segment, whereas the other three species all have a distinct narrowing between the prostate and penial sac. The penial sac of T. tamaraw (Fig. 4 I), T. stegodon (Fig. 4 J) and T. kamagong (Fig. 16 B) is relatively short, whereas it is much longer in T. sp. ‘ Anilao’ (Fig. 16 A).	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFBBFB0CEA1FFD6AFD37FE50.taxon	description	(FIGS 12 E, UPPER PHOTO, 16 C, 18) 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: FBDFFE 3 E- 81 DB- 4 FA 3 - A 05 F-ECD 10 A 728 B 88.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFBBFB0BEB94FCBCFBC5FB6D.taxon	materials_examined	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. 12 E, 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. 18 A). The radular formula of the holotype is 19 × 1.0.1 (Fig. 18 B). The older teeth (Fig. 18 C) are much smaller than the newer ones (Fig. 18 D) 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. 16 C). 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 16 S (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. 12 E, 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 16 S 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 16 S 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.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFBCFB09E8BEF98AFD01FA2B.taxon	description	(FIGS 12 F, 16 D, 19) 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: 9 A 854 BA 0 - 0 AEA- 4 D 98 - 9 B 38 - A 7 ABD 853 AE 90.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
03E487E4FFBCFB09E8BEF98AFD01FA2B.taxon	materials_examined	Type material: Holotype: NMP 041331, originally CASIZ 186132, one specimen, dissected and sequenced, Ligpo Island, Balayan Bay, Batangas Province, Luzon Island, Philippines, 22 May 2011, collected by Peri Paleracio. Type locality: Ligpo Island, Balayan Bay, Batangas Province, Luzon Island, the Philippines. Geographical distribution: Known only from Lipgo Island, Philippines. Etymology: From Latin word undulata, small wave, referring the notable undulating mantle margin of this species. External morphology: Living specimen 7 mm (Fig. 12 F). It has an elongated body typical of Trapania with a creamy white body throughout, including the extra-branchial and extra-rhinophoral appendages. Rhinophores are club-shaped with about nine lamellae and a pointed tip. Gill has a central, branching stalk flanked by two smaller branches. Both rhinophores and gills are reddish orange / brown becoming darker at the edges. The rhinophores are reddish orange basally and on the club the reddish orange is only found along the edges of the lamellae; the remainder of the rhinophores is the same colour as the body. Two black ridges run along the edge of the notal margin, approximately from the base of the rhinophores until just before the beginning of the extra-branchial appendages. The notal edge is thicker than the remainder of the body and varies in its width, creating the appearance of having an undulating margin. The anterior margins of the foot extend laterally as elongate appendages and are a more translucent white than the rest of the body. The oral tentacles are a more intense orange version of the rhinophore and gill coloration. The colour becomes lighter and more yellowish towards the tip, and darker and more dark-orange and black towards the base. This coloration continues across the oral veil creating a continuous mask that is mostly black with some reddish colour in the centre. The grey posterior foot had a dorsal indentation along its length creating two ridges along the lateral length. Slight bluish tint adorns the most posterior portion of foot. 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. 19 A). The radular formula of the holotype is 22 × 1.0.1 (Fig. 19 B). The older teeth (Fig. 19 C) are much smaller than the newer ones (Fig. 19 D) and the radula widens towards the more newly developed teeth. The teeth bear numerous evenly graded denticles with the smallest ones being found on the inner edge of the tooth and the longest ones adjacent to the primary cusp. There are approximately seven to 17 denticles on the inner side of the much larger, thick 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. 16 D). The pre-ampullary duct is short and enters the ampulla at its proximal end. The elongate, cylindrical ampulla is narrow at the distal end where it connects to the ovotestis. From there it widens and narrows again to the point where it divides into the short oviduct and long vas deferens. The oviduct enters the female gland mass. The vas deferens is initially narrow and widens into a convoluted, glandular prostatic portion and gradually transitions into the wide, muscular ejaculatory portion that is continuous with the wide penial sac. The penial sac terminates at the gonopore adjacent to the vagina. The vagina is widest at its basal opening and narrows abruptly as a short region prior to its junction with the large, spherical bursa copulatrix and short, narrow, receptaculum duct. The receptaculum duct joins the 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 ABGD analyses and the bPTP analysis successfully separates Trapania undulata from its similarly coloured sister-species T. gibbera. Furthermore, these two species are separated by a strong genetic divergence of 11.5 – 12.9 % in the COI gene and 9.6 − 9.8 % in the 16 S gene (Tables 3, 4). Both species have a white body colour and have red markings on the oral tentacles, rhinophores and gill branches and a black line on the anterior margin of the head. In T. undulata, the notal margin undulates and has a black marginal line, whereas T. gibbera (see: Gosliner & Fahey, 2008: fig 1 F) does not have a distinct notal margin and also lacks any lateral lines along the lateral edge of the notum. Another species with a similar colour pattern is Trapania caerulea Gosliner & Fahey, 2008. It also has a white body colour and has red markings on the oral tentacles, rhinophores and gill branches and a black line on the anterior margin of the head. In T. caerulea, there are black lines that extend posteriorly from the rhinophores to the lateral margins of the body near the extrabranchial appendages, whereas T. undulata has black lines along the notal margin. The notal margin of T. caerulea is indistinct. Additionally, T. caerulea has a blue Y-shaped marking on the notum between the rhinophores and gill and a second blue line posterior to the gill, running to the posterior end of the foot. In T. undulata, there is only a small blue patch at the posterior end of the foot. Unfortunately, no specimens of T. caerulea are available for genetic comparison. Internally, all three species have similar radular teeth but in T. caerulea, the denticles on the inside of the primary cusp seem to alternate between small and large denticles, whereas they are less regular in T. undulata and T. gibbera. All three species have a wide distal portion of the vagina. In T. caerulea and T. gibbera, there are two distinct vaginal widenings, whereas in T. undulata there is only one. Both T. caerulea and T. gibbera have a long vagina, whereas T. undulata has a short vagina.	en	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. Zoological Journal of the Linnean Society 196: 270-313, DOI: 10.1093/zoolinnean/zlac009
