taxonID	type	description	language	source
038E1B0DFFA2E97FD3891676B428F945.taxon	type_taxon	Type species: Doris coronata (Gmelin, 1791).	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	description	urn: lsid: zoobank. org: act: 212 CB 742 - C 51 E- 4 FE 7 - 80 AF-D 1485 B 615 BB 5 (Figs. 2 A – D, 3 A, 4, 5 A)	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	materials_examined	Type material. Holotype. CASIZ 182040, Cabrillo launch ramp 33.713231 ° N – 118.283415 ° E, San Pedro Harbor, 12 December 2009, Á. Valdés collector. Paratypes: CASIZ 056222, 15 specimens, mouth of Drake’s Estero Marin County, CA, 24 April 1984, Jeff Goddard, collector. CASIZ 184511, 7 specimens, one dissected, Ellwood boulder field, between Gaviota and Goleta, Santa Barbara County, California, 15 March 2010, Jeff Goddard collector. CASIZ 217592, one specimen, Santa Cruz boat harbor 35.964552 ° N – 122.001849 ° E, Santa Cruz, CA, 18 August 2017, Brenna Green, collector. CASIZ 217599, one specimen, Santa Cruz boat harbor 35.964552 ° N – 122.001849 ° E, Santa Cruz, CA, 18 August 2017, Brenna Green, collector.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	description	External morphology (Fig. 2 A – D). The living animals are 4 – 7 mm in length and the preserved holotype is 4.5 mm. The body color is translucent white with cream colored ovotestis visible through the body wall. There are a few to many small opaque white spots visible on the tips of the rhinophores not extending to the sheath or the head. The rhinophores extend just slightly above the sheath, which is well elevated from the notum. The apical margin of the rhinophore sheath is flared and of uniform width around the entire margin or wider anteriorly, and a few opaque white spots are occasionally present along the sheath margin. The digestive gland within each ceras ranges from a peach salmon to vibrant pinkish red. There are 4 – 7 pairs of cerata with around 15 tubercles on each ceras. The well-spaced tubercles are arranged in about 3 – 5 loosely organized whorls and are ovoid in shape. Each tubercle has an opaque white glandular apex. In some individuals (Fig. 2 C) cerata have been lost and are regenerating. The pseudobranch (Fig. 3 A) has up to five palmately lobed branches that are situated near the base of each ceras on the inner side. The anus is located on the right side between the first and second cerata. The genital opening is situated under the first ceras on the right side. The head is rounded with short, rounded extensions visible laterally. Buccal armature (Fig. 4). The buccal mass is small and muscular. The jaws are thin and membranous. The radula is elongate and narrow, consisting of a ribbon of at least 75 teeth. The asymmetrical teeth are simply arched posteriorly with a series of anterior denticles. Each tooth has an elongate central cusp that extends more anteriorly than the lateral denticles. The first two denticles on either side of the central cusp are offset, giving the tooth an asymmetrical appearance. There are two to three pairs of elongate denticles on either side of the central cusp. On the outer side of the tooth there are an additional 1 – 2 smaller denticles on either side of the tooth. Reproductive system (Fig. 5 A). The reproductive system is androdiaulic. The pre-ampullary duct is short and expands into the wide, ovoid ampulla. The ampulla again narrows distally and divides into the oviduct and vas deferens. The oviduct widens into a serial receptaculum seminis that narrows into an elongate convoluted vagina that terminates at the female gonopore adjacent to the penis. The vas deferens widens abruptly into a curved, thickened prostatic portion and narrows into a thin, undulating ejaculatory portion. The distal end of vas deferens enters the large, straight penial sac, which terminates adjacent to the opening of the vagina. The female glands are not well differentiated from each other, but the mucous gland comprises the bulk of the organs and terminates adjacent to the opening of the vagina and penis and has a separate aperture.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	etymology	Etymology. The species name urak comes from the southern Ohlone word for salmon, indicating the salmon pink to orange color of the cerata which is a distinctive feature of this species. The Ohlone people were the original inhabitants of much of the range of this species.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	distribution	Distribution. Known from San Diego County to Humboldt County, California, USA.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA2E97AD3891685B5C7FDD7.taxon	discussion	Remarks. Doto urak is sister to D. amyra, and both are sister to D. columbiana. Doto urak is externally most similar to D. amyra and has been confused with this species previously (D. Behrens, 1980, 1991; D. Behrens & Hermosillo, 2005). Both species lack dark pigment on the body. Doto amyr a is distinctly longer than D. urak and can reach 12 mm in length. The rhinophores on most D. amyra specimens are proportionately shorter than those of D. urak. The rhinophore sheaths of D. urak are wide and more flared than those of D. amyra. The ceratal cores and digestive glands of D. amyra are a much lighter white / beige color, while those of D urak are generally a salmon pink to orange or dark rust / red color. Additionally, D. urak has ceratal tubercles with opaque white glands that are not evident in D. amyra. In D. urak, the ceratal tubercles are more widely spaced than in D. amyra. Also, the tubercles in D. urak are more rounded and regular in shape than the flatter ovoid tubercles of D. amyra. The cerata of D. amyra are usually longer, with some exceptions noted (Agarwal, 2019) and also have more rows of tubercles. In D. amyra, the pseudobranch extends most of the length of the ceras with a few short lateral branches (Marcus, 1961 a, figs. 131, 132, 136) whereas D. urak has a palmately branched pseudobranch that is found basally on each ceras (Fig. 3 A). Previous literature on both species have indicated that D. amyra is more transparent than its counterpart, but some images suggest otherwise (McDonald, 2017). The most significant difference is in the developmental biology of the two species: Doto urak lays eggs averaging 76 µm in diameter and has planktotrophic larvae, while D. amyra lays eggs averaging 152 µm in diameter and has lecithotrophic larvae (J. H. R. Goddard, 1996; Shipman & Gosliner, 2015). Internally, the radular teeth of most of the northeastern Pacific Doto species are very similar in shape and appear to all have a certain degree of asymmetry (Marcus, 1961 a). The reproductive systems are also similar in form with a serial receptaculum seminis (Marcus, 1961 a), but do appear to differ in several details of their anatomy. In D. urak, the penial sac is more elongate than that of D. amyra (Marcus, 1961 a: figs. 134, 138 as D. wara, fig. 146 as D. ganda), while the prostate is not as convoluted as in D. amyra. Also, there is a distinct muscular sphincter at the distal end of the vagina in D. columbiana (Marcus, 1961 a: fig. 129) and D. amyra (Marcus, 1961 a) that is apparently absent in D. urak and D. kya (Marcus, 1961 a). Doto urak is molecularly distinct (BI = 1, ML = 100) and is a single distinct species according to each species delimitation method undertaken. D. urak has an intraspecific variation p - distance of 0.00 – 1.90 %. The interspecific p - distances between D. urak and the closest sister species are: 4.95 – 6.69 % to Doto amyra, 7.90 – 9.09 % to D. columbiana, 10.49 – 12.68 % to D. kya, and 6.29 – 7.54 % to D. sp. B.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	description	urn: lsid: zoobank. org: act: F 10 B 38 D 1 - ECA 4 - 4 BB 6 - BF 37 - 9 ECB 962834 CB (Figs. 2 E – F, 3 C, 5 B, 6)	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	materials_examined	Type material. Holotype. CASIZ 234657, 4.5 mm preserved, off Cuvier Park 32.843073 ° N – 117.281888 ° E, La Jolla, California, 27 February 2022, Chloe and Trevor Van Loon collectors.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	description	External morphology (Fig. 2 E – F). The living animal has a translucent white body color with a cream colored ovotestis visible through the body. There are small opaque white spots visible along the margins of the foot, along the margin of the rhinophore sheath, and on the entire length of the rhinophore. The dorsal notum and sides of the body have numerous scattered ovoid saffron yellowish orange spots, some of which have irregular borders and are slightly raised. The digestive gland within each ceras is light brownish with opaque white glands present within the tubercles on the cerata. There are 7 pairs of cerata with 4 whorls of elongate tubercles with bulbous tips. The size of the tubercles varies, but in general the more basal ones are smaller with the largest being the single terminal tubercle. The cerata often lack pseudobranchs but in one large ceras the pseudobranch consisted of a single linear axis without lateral branches. The rhinophoral sheaths are well-elevated from the notum with an extension on the anterior side. The head is broader than the rest of the body with a pair of curved lobes situated laterally. The anal papilla is situated dorsally, posterior to the second right ceras and genital openings on right side of body, immediately ventral to the first ceras. On the head of one specimen, photographed from off the east end of Santa Cruz Island, there is a brown patch of what appears to be a subcutaneous pigmentation (Klug, 2019). An additional specimen, from off the Palos Verdes Peninsula and included in Behrens et al. (2022) has subcutaneous black streaks extending from the head to near the posterior end of the body. Buccal armature (Fig. 6). The buccal mass is small and muscular. The jaws are thin and membranous. The radula is elongate and narrow, consisting of a ribbon of at least 69 teeth. The teeth are simply arched posteriorly with a series of anterior denticles. Each tooth has an elongate central cusp that extends more anteriorly than the lateral denticles. There are one to two pairs of elongate denticles on either side of the central cusp. On the outer side of the tooth there are an additional 1 – 2 smaller denticles on either side of the tooth. Reproductive system (Fig. 5 B). The reproductive system is androdiaulic. The pre-ampullary duct is short and expands into the wide, ovoid ampulla. The ampulla again narrows distally and divides into the oviduct and vas deferens. The oviduct widens into a serial receptaculum seminis that narrows into an elongate curved vagina that terminates at the female gonopore adjacent to the penis. The vas deferens widens into a curved, thickened prostatic portion and narrows into a thin, curved ejaculatory portion. The distal end of vas deferens enters the thick, curved penial sac, which terminates adjacent to the opening of the vagina. The female glands are not well differentiated from each other, but the mucous gland comprises the bulk of the organs and terminates adjacent to the opening of the vagina and penis and has a separate aperture.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	etymology	Etymology. The species epithet comes from the Kumeyaay indigenous name kwakwak, meaning yellow, signifying the yellow-orange spots on the notum that distinguish this species. The Kumeyaay are the original inhabitants of the region around the type locality in San Diego County.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	distribution	Distribution. Known intertidally from La Jolla, and subtidally from off the Palos Verdes Peninsula, Isla Vista, and Anacapa and Santa Cruz Islands, California, USA (Behrens et al., 2022, present study).	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
038E1B0DFFA7E965D3891233B745FD2B.taxon	discussion	Remarks. This species is known only from a few photographs and the only documented specimen collected is the holotype, from La Jolla, CA (Fig. 2 E) (Van Loon, 2022). Six other specimens have been photographed: three from Anacapa Island in the Channel Islands by Johnson (2018) (Fig. 2 F) and Klug (2017, 2020); one from Santa Cruz Island in the Channel Islands (Klug, 2019); one by Margaret Webb from off Haggerty’s, Palos Verdes, CA (Behrens et al., 2022); and one from the shallow subtidal off Coal Oil Point, Isla Vista, CA by Mazza (2019). In our phylogeny, Doto kwakwak and Doto kya are sister to most of the other members of the Atlantic / eastern Pacific clade (Fig. 1). Of the species found along the Pacific coast of the United States and Canada, it can be readily distinguished externally by the presence of yellowish-orange spots on the body. Similarly, D. kwakwak, like D. amyra and D. urak, generally lacks the scattered brown pigment on the body that is found in D. kya and D. columbiana, but some specimens may have subcutaneous dark pigment (Klug, 2019). In D. amyra, the lateral extensions of the foot are elongate, whereas, they are much shorter in D. columbiana, D. kya, D. lancei Marcus & Marcus, 1967, D. urak, and D. kwakwak. In D. urak and D. kwakwak, the rhinophore sheaths are flared apically while they are largely cylindrical in D. amyra, D. columbaiana, D. kya, and D. lancei. The ceratal tubercles of D. kwakwak, D. urak, D. lancei, and D. kya are spherical to elongate apically, while those of D. columbiana and D. amyra are much lower to flattened or irregular in shape (Fig. 3 B). Also, the opaque white glandular apices of the ceratal tubercles of D. kwakwak, D. urak, and D. kya differ from those of D. columbiana, D. lancei, and D. amyra, which lack opaque white on the tubercle apices. Doto kwakwak has many tubercles of different sizes shapes (some bulbous and some longer), in contrast to D. amyra, D. urak, D. lancei, D. columbiana, and D. kya, which have uniformly tuberculate cerata. The cerata or D. kwakwak are elongate like those of D. amyra, D. kya, D. lancei, and D. columbiana, but tend to get longer towards the posterior end of the body. The cerata of D. urak are shorter and more rounded. The digestive gland within the ceratal core of D. kwakwak appears to be a much more vibrant rust / red color, than in D. kya, D. lancei, and D. columbiana, which have beige to light orange cores. In D. kwakwak, a pseudobranch is either absent from the inner side of the cerata or if present is only an unbranched vertical line. In all other eastern Pacific species of Doto a well-developed, branched pseudobranch is present (Fig. 3 B). As stated in the remarks regarding D. urak, the species of Doto from the Pacific coast of North America all have similar radular tooth morphology, and there is nothing distinctive about the radular teeth of D. kwakwak. The reproductive system of D. kwakwak is similar to that of D. columbiana (Marcus, 1961 a: fig. 129) with a relatively short, curved ejaculatory segment and a relatively short penial sac. However, the vaginal duct of D. columbiana has a distinct sphincter as in D. amyra, in contrast to that of D. kwakwa k (Fig. 5 B), D. urak (Fig. 5 A), and D. kya (Marcus, 1961 a: fig. 140). Doto kwakwak is molecularly distinct and is a single distinct species according to each species delimitation method undertaken. D. kwakwak interspecific p - distances to the closest species are: 12.51 – 12.99 % to Doto amyra, 14.27 % to D. columbiana, 14.09 – 14.46 % to D. kya, 13.54 – 14.64 % to D. lancei, 12.47 – 12.84 % from D. sp. B, 14.09 % to D. sp. C, 14.83 % to D. sp. I, and 13.90 – 14.93 % to D. urak. Little is known about the natural history of this species and its larval biology and development mode remains unknown.	en	Adayapalam, Sneha, Bonomo, Lynn J., Shipman, Carissa, Gosliner, Terrence M. (2025): The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California. Zootaxa 5609 (4): 553-577, DOI: 10.11646/zootaxa.5609.4.6, URL: https://doi.org/10.11646/zootaxa.5609.4.6
