THE NAUTILUS 129(l):31-42, 2015 Page 31 Reviving a cold case: two northeastern Pacific dendrodorid nudibranchs reassessed (Gastropoda: Opisthobranchia) Jeffrey H.R. Goddard Angel Valdes Marine Science Institute Department of Biological Sciences University of California California State Polytechnic University Santa Barbara, CA 93106-6150 USA 3801 West Temple Avenue [email protected] Pomona, CA 91768-4032 USA [email protected] ABSTRACT ing the Gulf of California (Behrens and Hermosillo, 2005): Dendrodoris azineae Behrens and Valdes, 2004; Ten nominal species of dendrodoridid nudibranchs are known Dendrodoris behrensi Millen and Bertsch, 2005; Dendrodoris from the NE Pacific Ocean, including the Gulf of California. fumata (Riippell and Leuchart, 1831); Dendrodoris stohleri However, disagreement surrounds the taxonomic status of Millen and Bertsch, 2005; DoriojtsiUa albopunctata (Cooper, Doriopsilla nigromaculata (Cockerell in Cockerell and Eliot, 1905) and Doriopsilla rowena Marcus and Marcus, 1967, includ¬ 1863); Doriopsilla gemela Gosliner, Schaefer and Millen, ing the correct generic placement of the former. To resolve 1999; Doriopsilla janaina Marcus and Marcus, 1967; this disagreement, we examined type specimens, the original Doriopsilla nigromaculata (Cockerell in Cockerell and descriptions, and unpublished materials in the James Lance Eliot, 1905); Doriopsilla rowena Marcus and Marcus, Collection at the California Academy of Sciences and conclude 1967; and Doriopsilla spaldingi Valdes and Behrens, that Doriopsilla nigromaculata is: (1) a member of Dendrodoris; 1998. However, an important taxonomic disagreement (2) not synonymous widi the valid species Doriopsilla rowena-, remains in the modern literature. Camacho-Garcia et al. and (3) a senior synonym of Dendrodoris behrensi Millen and (2005) considered Doriopsilla rowena a valid species, Bertsch, 2005. Like other members of the genus, Dendrodoris whereas Behrens and Hermosillo (2005) regarded it as a nigromaculata has a centered anus, smooth dorsum, delicate wavy mande edge, and possesses both ptyaline and esophageal junior synonym of Doriopsilla nigromaculata. The sug¬ glands. It is translucent white widi chocolate brown blotches, the gested synonymy of D. nigromaadata and D. rowena was larger of which are usually clustered into three or four groups raised as a possibility by James Lance in Keen (1971: 830; centered mid-dorsally. In contrast, Doriopsilla rowena has an cited as Doriopsilla nigromaadata) and again by McDonald off-center anus, a papillate and densely spiculate dorsum with a (1983: 171; cited as Dendrodoris nigromaadata). How¬ stiff margin, and lacks both ptyaline and esophageal glands. Its ever, if Camaeho-Gareia et al. (2005) were correct, and scattered brown decks and larger, round concentrations of Doriopsilla rowena is valid, then D. nigromaculata, which opaque white distinguish it dorsaOy, and notal spicules include was originally described based on a single specimen col¬ rods and forks. Doriopsilla rmuena grows to 12 mm long and has lected by Cockerell in La Jolla, California, has either been found in La Jolla, California, the Pacific coast of Baja California, and the northern Gulf of California soudi to Panama. remained unknown since its original description and Dendrodoris nigromaculata grows to 27 mm and is known from should be regarded as a nomen dubium, or has since been Monterey, California soudi to the San Benitos Islands, Baja described under another name. California. Bodi species have large eggs and ametamoqihic A contributing factor to this disagreement is that direct development, but small eggs indicating planktotropbic historically the distinctions between Dendrodoris and development have also been observed in D. rcnvena from Jalisco, Doriopsilla have been confusing, including for species Mexico, suggesting D. rowena may constitute a cryptic species from the northeast Pacific Ocean (reviewed by Steinberg complex or display poecilogony. 1961; Valdes and Ortea 1997; Gosliner et al. 1999). How¬ Additional Keywords: Dendrodorididae, Nudibranchia, ever, Valdes et al. (1996) and Valdes and Ortea (1997) nomenclature provided clear anatomical and morphological criteria separating the two genera, and recent phylogenetic anal¬ yses support this separation (Valdes and Gosliner 1999, Valdes 2003). Here, we use these criteria and the exam¬ ination of type specimens to first establish the correct generic placement of D. rowena and D. nigromaculata, INTRODUCTION ruling out the synonymy of these two species. Then we Ten nominal species of dendrodoridid nudibranchs are compare their original descriptions with those of other currently recognized from the NE Pacific Ocean, includ¬ species of dendrodoridids known from the region to Page 32 THE NAUTILUS, Vol. 129, No. 1 show that (1) Doriopsilla rowena is valid, and (2) that research.calacademy.org/redirect?url = http://research one species described recently from the region is in fact archive.calacademy.org/research/izg/iz_coll_db/index.asp) a junior synonym of Dendrodoris nigromaculata. In both to obtain additional and otherwise unpublished locality steps we also draw from extensive materials in the James information to further document the known geographic R. Lance Collection at the California Academy of Sci¬ distribution of these species. Here, we used information ences in San Francisco (hereafter, Lance Collection). for specimens identified by established authorities famil¬ These constitute new evidence not available prior to iar with the nudibranch fauna of the region. These are Lance’s death in 2006 and shed light on what had effec¬ referenced below by CASIZ and the corresponding cata¬ tively become a taxonomic cold case. log numbers. RESULTS MATERIALS AND METHODS To establish the generic placement of Doriopsilla Description oe Relevant Materials in the Lance Collection nigromaculata and Doriopsilla rowena, we extracted from their original descriptions information on four mor¬ 1. Species Folder: Doriopsilla rowena contains: phological and two anatomical characters which taken (1) A hand-written description, accompanied by pencil together can be used to separate Dendrodoris from sketches, of this species with the manuscript name Doriopsilla (Table 1). We corroborated this information “Doriopsilla puertecitensis” (later, Lance wrote the name by examining the type specimens of both species, which D. rowena in red ink on this description). The pencil were obtained, respectively, from The Natural History sketches are based on living specimens (CASIZ 182606) Museum, London (NHMUK) and the U.S. National collected 20 March 1965 from 4.4 mi south of Puertecito, M useum of Natural History (USMNH). We also exam¬ Baja California, and include dorsal and ventral views of an ined Lance’s specimens of D. rowena in the Invertebrate adult, details of the notum and notal spicules, and penis. Zoology and Geology collection at the California Acad¬ The description also includes a sketch and brief descrip¬ emy of Sciences (CASIZ) and used his description of D. tion of an egg mass laid by a 12 mm adult. (2) Six separate rowena in Keen (1971), as well as his unpublished notes, pen and ink illustrations, all labeled as Doriopsilla rowena 35mm photographic slides, and illustrations in the Lance and clearly based on the above pencil sketches. (3) A Collection. Lance kept many of these materials orga¬ sheet labeled ‘'Doriopsilla rowena - Field Account Data” nized by species in folders, which contain information for sites in the Gulf of California. The dates listed include on the morphology, color, anatomy, egg masses, and years from 1954 to 1979, and out of 10 sites listed record development of living specimens of most of the D. rowena only from Puerto Penasco. (4) One clear plastic dendrodoridids known from the northeast Pacific Ocean. sheet holding 35 mm slides from 1966 and 1969, the The folder for D. rowena in particular supplements earliest labeled first as “Dendrodoris sp.” and later as Marcus and Marcus’s original (1967) description of “Doriopsilla rowena or D. nigromacidata?” D. rowena, which was based on preserved material. Where appropriate, we refer to species folders in the 2. Species Folder: Doriopsilla nigromaculata con¬ Lance Collection by their names; field accounts by num¬ tains: (1) A list of specimens found at South Casa Reef ber, date, and locality; and 35mm slides by date and and Windansea Reef, both in La Jolla, California. (2) A locality when possible. To compare D. nigromaculata lined sheet with dorsal and ventral sketches of an adult with other species of Dendrodoris from the northeast from La Jolla. The notation “F.A. 181”on this sheet refers Pacific Ocean, we used information from our examina¬ to Field Account 181, which was for 25 June 1967 at tion of the type specimen, as well as from the Lance South Casa Reef, La Jolla. This sheet is first labelled as Collection, Behrens and Hermosillo (2005), Millen and “Brown spotted Doriopsilla,” with “? D. nigromaculata” Bertsch (2005), and Goddard (2005). Finally, we used later added in red ink. (3) A note stating “D. nigromaculata the online searchable database of the California Acad¬ Published sketch in Opis. News. 14(8): 29 of hatching.” emy of Sciences Invertebrate Zoology collection (http:// (4) A three page typed description of this species with the Table 1. Select diagnostic characters distinguishing Dendrodoris from Doriopsilla. Based on Valdes et al. (1996) and Valdes and Ortea (1997). Character Dendrodoris Doriopsilla Mantle Smooth and soft, rarely with large tubercles Hard and tubereulate Mantle spicules Absent or isolated Strong network, including in tubercles Mantle margin Delicate, wavy Stiff Position of anus relative to midline of body Centered Off-center Ptyaline glands Present Absent Esophageal glands Present Absent J.H.R. Goddard and A. Valdes, 2015 Page 33 Figures 1-6. Type specimens. 1. Dendrodoris nigromaculata (NHMUK 1904.7.7.1). Photo by Harry Taylor. 2-6. Doriopsilla roxvena (USMNH 678413). Photos by Chris Meyer. heading “Doriopsilla nigromaculata (Cockerell and Eliot, San Diego. (2) A hand-written description of this species 1905) Figs. 1-7.” The figures for this description and with the manuscript name “Dendrodoris barbarensis,” legend are on a separate sheet and are a composite of based on a single specimen collected from 8 m depth at copies of the six pen and ink illustrations from the Species Naples Reef, Santa Barbara County 30 Oct 1966, two Folder lor Doriopsilla rowena, plus a map of California specimens collected at 30 m depth 1.6 km south of South with a single red ink dot showing “Distribution in California.” Coronado Island by Nan Limbaugh on 22 Apr 1961, and The description is based on the four specimens Lance two specimens collected intertidally at Point Loma by found in La Jolla on 25 June 1967 but refers to the speci¬ Wesley Farmer on 29 Oct 1963. (3) A typed sheet with men from La Jolla described by Cockerell and Eliot information on three specimens collected intertidally at 70 years earlier. (5) Two sheets with sketches and notes Lunada Bay, Palos Verdes Peninsula, by William Jaeekle, on egg masses laid by specimens collected at Windansea 20 Apr 1983. A note indicates that one of these speci¬ Reel in 1968, San Quintin in 2001, and South Casa Reef mens laid an egg mass on 4 May 1983. (4) Pencil notes in 1998. (6) Two sheets containing 35 mm photographic and sketches of an egg mass laid by an individual col¬ slides, one with images of adult specimens, the other with lected from South Casa Reef, La Jolla, 20 July 1974, and images of egg masses and hatching juveniles. sketches of the subsequent direct development to hatch¬ ing juveniles from that egg mass. 3. Species Folder: White Porostome Spotted con¬ tains: (1) A sheet titled “Crenulate dorid” with sketches 4. Field Account 181 (June 25, 1967, South Casa Reef, in pencil of two specimens and a brief description, La Jolla) contains, in addition to the annotated list of opis- including dimensions of 24 x 5 mm and 27 x 5 mm. thobranchs found on that date by Lance (and Barbara This sheet is undated but the dimensions of the larger Good), contains notes, from three separate dates, on die specimen, combined with information in (2) below indi¬ development of embryos in an egg mass deposited by cate that these specimens were collected in either 1961 Doriopsilla rowena (cited as “doriopsillids,” with a later nota¬ from the Coronados Islands or in 1963 from Point Loma, tion as “Doriopsilla nigromaculata?”) collected on that date. Page 34 THE NAUTILUS, Vol. 129, No. 1 Figures 7-10. Living adult Doriopsilla rowena. 7. Percebu, Gulf of California, Baja California, 7 April 1966 (35 mm slide in Lance Collection, Species Folder: Doriopsilla rowena). 8. Total length 9.2 mm, San Quintin, Baja California, 16 November 2001 (35 mm slide in Lance Collection, Species Folder: Doriopsilla nigromaculata). 9. South Casa Reef, La Jolla, California, 25 June 1967 (35 mm slide in Lance Collection, Folder: Doriopsilla nigromaculata). 10. Total length (in MgCl2) 6.7 mm, Lindo Mar, Bahia de Banderas, Jalisco, Mexico, 26 Feb. 2006 (CASIZ 174055; Goddard and Hermosillo, 2008, as Doriopsilla nigromaculata). Genus-Level Taxonomy gills, respectively, in the type specimen, are consistent with contemporary descriptions of species of Dendrodoris and Doriopsilla nigromaculata. In the original descrip¬ clearly indicate the correct generic of D. nigromaculata is tion of D. nigromaculata Cockerell and Eliot (1905) Dendrodoris (Table 1). described the mantle of a single preserved specimen as “smooth, not tubereulate,” with a “rather narrow” mar¬ Doriopsilla rowena. Marcus and Marcus (1967) gin. The latter is consistent with the delicate margin char¬ described D. rowena based on seven preserved syntypes, acteristic of living specimens of Dendrodoris (Table 1) and five of which remain (USMNH 678413), and notes by is visible in the type specimen (NHMUK 1904.7.7.1) as the collector on the color of the living animals, which upturned, crenulate and soft in appearance (Figure 1). were collected in Puerto Penasco, Sonora, Mexico, in Cockerell and Eliot (1905) mentioned “a number of glis¬ the northern Gulf of California (Figures 2-6). They tening white spicules.. .imbedded in the skin” of the man¬ described the mantle as “smooth,” with evidence of tle and described them as “mostly fairly straight long rods, large, but dissolved, diagonally crossed spicules in the but some of the smaller ones are bent and have an irreg¬ “connective tissue of the back." They noted that “the ular outline.” The spicules are not described as being thick anal papilla and the renal pore in front of it lie to regularly or densely arrayed and are visible in the type the left of the branchial tuft,” clearly indicating the specimen where the mantle tissue had been carefully eccentric position of the anus. The anterior part of the scraped away, presumably by Eliot in his original exami¬ alimentary tract is illustrated in their figure 62B, and nation of the specimen (Figure 1). The position of the does not show either ptyaline or esophageal glands. Else¬ anus can be inferred as centered based on their descrip¬ where, Marcus and Marcus (1967: 99) state that the tion of the gill plume as “set in a semicircle open absence of a ptyaline gland is one characteristic of behind.” Internally, they described a “follieulate” mouth Doriopsilla. Marcus and Marcus (1967) did not describe gland “with a fairly long duct,” which is clearly a ptyaline any dorsal tubercles in D. roivena. Plowever, James Lance, gland (e.g., illustrations in Valdes et ah, 1996; Millen and who found and observed living specimens of this species Bertsch, 2005). No esophageal glands were mentioned, from Puerteeitos, Baja California, and Puerto Penasco, but these are minute (Valdes et al. 1996; Millen and both in the northern Gulf of California (Figures 11—12), Bertsch, 2005, Figure 2) and likely would not have gar¬ illustrated minute, spiculate dorsal tubercles, which was nered attention. All of these traits, as well as the extreme confirmed with examination of Lances original specimens anterior and posterior position of the rhinophores and (CASIZ 182606). Lance also illustrated the pattern and J.H.R. Goddard and A. Valdes, 2015 Page 35 Species-Level Taxonomy 11 Doriopsilla roivena. Based on the above morpho¬ logical and anatomical differences, Doriopsilla roivena cannot be a synonym of Dendrodoris nigromaculata. Doriopsilla roivena differs externally from all other con¬ geners known from the region in its small adult size (up to 12 mm total length) and unique color pattern, consisting of small reddish-brown flecks scattered over the dorsum, as well as larger, round concentrations of opaque white, frequently arranged in longitudinal series, against a ground color of off white to pale yellow, pink or orange (Marcus and Marcus, 1967; Lance in Keen 1971; Behrens and Ilermosillo, 2005, cited as Doriopsilla nigromaculata; Camaeho-Garefa et al., 2005) (Figures 2-10). Doriopsilla roivena is therefore a valid species. Dendrodoris nigromaculata. Cockerell and Eliot (1905) based their original description of D. mgromaculata on a single specimen, 10 mm long preserved, that was collected in July 1901 from La Jolla, California (Figure 1). The color of die preserved specimen was described as “yellowish-drab with a slight inclination to lilac in places,” with “a double border of black spots round the dorsal margin, and a few larger black blotches symmetiieally dis¬ posed, one in front of the rhinophores, two behind them, two in die middle of die back, and five in front of the branchiae.” These larger blotches are still visible in the type specimen (Figure 1). Two nominal species of Dendrodoris from die northeast Pacific Ocean have dark spots against a pale background: D. behrensi Milieu and Bertsch, 2005 (Figure 17) and D. .stohleri Millen and Bertsch, 2005. In D. stohleri die black spots are relatively uniform in size and scattered over the entire dorsum, save its margin (Millen and Bertsch, 2005). However, compari¬ son of the color patterns originally described for D. nigronuicidata and D. behrensi reveals a virtually identical match (Table 2). Indeed, the only discrepancy between the descriptions of these two species is that Millen and Bertsch (2005: 195) state that D. behrensi lacks mantle spicules. However, in the companion paper to Millen and Bertsch (2005), Goddard (2005) illustrated and described “slightly Figures 11-13. Doriopsilla roivena, details of notum; all curved, spindle-shaped spicules” arranged in a lattice Irom Lance Collection, Species Folder: Doriopsilla roivena. in D. behrensi recently hatched (Figure 21) from egg 11. 1x1 mm detail of notal surface. 12. Notal tubercle with masses hud by adults (Figure 17) included as paratypes spicule detail. 13. Notal spicule pattern. of D. behrensi by Millen and Bertsch (2005). Lance was also familiar with this species and observed egg masses and hatching juveniles virtually identical to those described by Goddard (2005) (Lance Collection, Spe¬ shapes of the notal spicules (Figure 13), which consist of cies Folder: White Porostome Spotted) (Figures 18-20). diagonally crossed rods and forks (Lance Collection, Spe¬ Moreover, Lance described the adult body of this cies Folder: Doriopsilla roivena). Taken together, these species as “very slightly spiculose with notal margins traits indicate that D. roivena is indeed a Doriopsilla, non spiculose” (Lance Collection, Species Folder: White as originally described by Marcus and Marcus (1967) Porostome Spotted). The density of notal spicules there¬ (Table 1). This was further confirmed with the examination fore decreases as juveniles grow into adults, which of the syntypes (Figures 2-6), which are relatively flat and could make the spicules easily overlooked in living adult wide animals covered with small spiculate tubercles and an specimens. A similar phenomenon was recently docu¬ eccentric anus. All these traits are consistent widi the diag¬ mented by Sanehez-Tocino et al. (2014) for some chromo- nosis of Doriopsilla by Valdes and Gosliner (1999). dorid nudibranchs. Page 36 THE NAUTILUS, Vol. 129, No. 1 Table 2. Comparison of original descriptions of external color of a single preserved Dendrodoris nigromaculata and living Dendrodoris behrensi by Cockerell and Eliot (1905) and Millen and Bertsch (2005), respectively. Dendrodoris nigromaculata Dendrodoris behrensi (Cockerell and Eliot, 1905) Millen and Bertsch, 2005 Background color “yellowish-drab with a slight inclination “semi-translucent white or pale cream” to lilac in places” Larger spots Black, “symmetrically disposed:” Dark reddish brown or chocolate brown, “clustered in four areas in the mid line:” Position Group 1 “in f ront of the rhinophores” “in front of the rhinophores” 2 “behind [rhinophores]” “behind the rhinophores” 3 “middle of back" “middle of the body” 4 “in front of the branchiae” “in front of the gills” Smaller spots “double border... round the dorsal margin” “scattered towards the edges of the dorsum” Other color elements None None With no significant differences between their original Anatomy: The anatomy of D. rowena was described descriptions, Dendrodoris behrensi is therefore a junior by Marcus and Marcus (1967), with additional details synonym of Dendrodoris nigromaculata. presented by Valdes and Gosliner (1999). Further, as noted and illustrated by Lance, the notum is densely spieulate with rods and forks (Figures 12-13, 16). SYSTEM ATICS External Morphology: The external morphology of D. roivena was described by Marcus and Marcus (1967), Based on the taxonomic results described above, a new with additional details presented by Lance (Lance systematic arrangement and list of synonyms is proposed Collection, Species Folders: Doriopsilla rowena and for D. rowena and D. nigromaculata. Doriopsilla nigromaculata) and Lance in Keen (1971). Family Dendrodorididae O’Donoghue, 1924 Living adults reach 12 mm in length. The dorsum is flecked with dark brown, has larger, round concentra¬ Genus Doriopsilla Bergh, 1880 tions of opaque white, typically in longitudinal series, and is covered by minute, spieulate tubercles (Figures 7- Doriopsilla rowena Marcus and Marcus, 1967 12). The ground color varies from whitish to yellow to (Figures 2-16) pinkish orange. Doriopsilla rowena Marcus and Marcus, 1967: 205- Development: Lance obtained egg masses and 207; Keen, 1971: 830; Poorman and Poorman, 1978: 373; observed development of Doriopsilla rowena from Bertsch and Kersbtch, 1984: 267; Valdes and Ortea, 1997: La Jolla (Figures 14-16). The egg ribbons were tightly 253; Gosliner et al., 1999: 209; Valdes and Gosliner, 1999: coiled and laid flat, rather than on edge like most other 338-340; Camacho-Garcia et al., 2005: 80; Goddard and dorid nudibranchs, and contained large eggs, encap¬ Hermosillo, 2008: 87; Angulo-Campillo, 2005: Table 2. sulated singly, that developed into hatching juveniles Dendrodoris (?) nigromaculata [non Cockerell in (Figure 15) in more than 26, but less than 38 days, at an Cockerell and Eliot, 1905] - Behrens, 1980: 58. ambient temperature of approximately 20° C (Lance Dendrodoris nigromaculata [non Cockerell in Collection, Species Folder: Doriopsilla nigromaculata-. Cockerell and Eliot, 1905],—Steinberg 1961: 59; Sphon Field Account 181, South Casa Reef, 25 June 1967). 1972: 61; McDonald and Nybakken, 1980: 52; Lance, Recently hatched juveniles possess the same notal spic¬ 1982: 29; McDonald, 1983: 170-171; Behrens, 1991: 71; ule complement of rods and forks as adults (Figures 13 Angulo-Campillo, 2003: Table 2; Goddard, 2004: 1959, and 16). Lance sketched another egg mass, laid by a 1963; Goddard, 2005: 206. specimen 9.5 mm long collected from Windansea Reef Doriopsilla nigromaculata [non Cockerell in Cockerell in June 1968. This egg mass was also laid flat and had 4 and Eliot, 1905],—Behrens and Hermosillo, 2005: whorls in a closed spiral and a total diameter of 5 mm, as 88; Lance Collection, Species Folder: Doriopsilla indicated by a scale bar. Based on that scale bar, the eggs nigromaculata. California Academy of Sciences and illustrated by Lance measured approximately 215 pm in Goddard, 2013: worksheets for Ladera St., False Point, diameter. A note on Lance Field Account 181 indicates Windansea, and So. Casa Reef (data from unpublished that the embryos did not develop a shell. Taken together, field accounts, no page numbers). this information indicates that D. rowena from La Jolla have ametamorphic direct development (Bonar 1978; Type Material: Doriopsilla roivena - Syn types: 5 Goddard 2004). Lances sketch and notes on an egg mass specimens, Puerto Penasco, Sonora, Mexico (USMNH laid by a D. roivena from San Quintin, Baja California 678413). are also consistent with this mode of development J.H.R. Goddard and A. Valdes, 2015 Page 37 Figures 14-16. Doriopsilla rowena, egg mass and hatching juveniles; all from 35 mm slides in Lance Collection, Species Folder: Doriopsilla nigromaculata. 14. Egg mass from South Casa Reef, 26 Apr. 1998. 15. Hatching juveniles, no date or locality (35 mm slide processed Sep. 1974). 16. Spicule detail, hatched juvenile. South Casa Reef, no date (35 mm slide processed Sep. 1974). (Lance Collection, Species Folder: Doriopsilla Geographic Distribution: Doriopsilla roivena is nigromaculata). known from the northern Gulf of California to Panama, A specimen of D. rowena (Figure 10; CASIZ 174055), (Marcus and Marcus 1967; Keen 1971; Poorman and 6.7 mm long, collected from Nayarit, Mexico laid a flat Poorman 1978; Bertsch and Kerstitch 1984; Camacho- egg ribbon of 6 turns in a closed spiral 6 mm across, Gareia et al. 2005; Goddard and Hermosillo 2008; virtually identical to the egg masses described above Angulo-Campillo 2005; CASIZ 171209, 171229), the from La Jolla (JG, unpublished data). However, the eggs Galapagos Islands (CASIZ 78390, 78408), and La Jolla, from Nayarit averaged 97.4 pm in diameter, indicating California to El Campo, near Punta Eugenia, Baja planktotrophic development (Goddard and Hermosillo, California Sur (Lance Collection, Species Folder: 2008). Lance described similar egg masses laid by speci¬ Doriopsilla nigromaculata; CASIZ 71519; Behrens anti mens of D. roivena from the northern Gulf of California Hermosillo 2005) (Figure 22). but mode of development cannot be inferred based on any of the information included (Lance Collection, Spe¬ Remarks: As evidenced in the Lance Collection (Spe¬ cies Folder: Doriopsilla rowena). cies Folder: Doriopsilla roivena), Jim Lance started to Page 38 THE NAUTILUS, Vol. 129, No. 1 describe D. rowena under the manuscript name “D. Eliot, 1905)”, the name used by California workers dur¬ puerteeitensis,” based on three specimens he collected at ing this time period for what we have shown here to be Puertecitos, on the Baja California side of the northern Doriopsilla roivena. Lance noted publication of this illus¬ Gulf of California. Sometime following the publication of tration on a separate sheet in his Species Folder Marcus and Marcus (1967), Lance added in red ink the “Doriopsilla nigromaculata”. However, comparison of name D. rowena on his original pencil sketches of these this illustration with his sketches of direct development specimens. Later he executed a set of undated pen and in his species folder “White Porostome Spotted" reveals ink illustrations, some of which are reproduced here in it to be copied from a sketch for that species, which we Figures 11-13 and labeled them all as Doriopsilla rowena demonstrate here to be Dendrodoris nigromaculata. (these illustrations were likely made for the monograph Although their egg masses and spicule complement on Panamic opisthobranehs that Lance intended but as hatching juveniles are different, both species have never completed). In the drawing of the dorsal view of ametamorphic direct development (see below), and an adult (not reproduced here) Lance depicted slx Lance (1982) was probably mainly making a statement thin, widely spread, bipinnate gills and a centered anus. about developmental mode in the one. The unexpected However, examination of his original specimens from twist is that the binomial used in the figure caption turns Puertecitos (CASIZ 182606) reveal more tightly clustered out to be accurate for the species actually illustrated. gills and an eccentric anus, indicating that Lances draw¬ The specimens described and identified as D. rowena ing in these respects was purely schematic. by Bertseh and Aguilar-Rosas (1984) from El Tomatal, On 25 June 1967 at South Casa Reef in La Jolla, Lance on the Pacific coast of Baja California, are, based on their found four specimens of a small (up to 9 mm long) larger size (up to 30 mm long) and color pattern, actually “doriopsillid” that he first referred to in his notes and Diaulula aurila Marcus and Marcus (1967), which is illustrations as the “Brown-spotted Doriopsilla” but later common in that region (Bertseh et al. 1999, cited as “Sal changed in red ink to “?D. nigromaculata” (Lance Collec¬ y pimenta” (Salt and pepper [dorid]); Goddard and tion, Field Account 181 and Species Folder: Doriopsilla Schickel 2000, cited as Discodoris sp. 1 of Behrens nigromaculata). He eventually found this species at three 1991; personal observations). more sites on the outer coast of San Diego County, as well as in Bahia San Quintin, Baja California, and in his field Genus Dendrodoris Ehrenberg, 1831 accounts for these sites consistently used the name Dendrodoris nigromaculata (Cockerell in Cockerell Doriopsilla nigromaculata for this species (Lance Collec¬ and Eliot, 1905) tion; California Academy of Sciences and Goddard 2013). (Figures 1, 17-21) Lance’s Species Folder for Doriopsilla nigromaculata includes a typed description, titled Doriopsilla Doridopsis vidua (?) [non Bergh, 1878],—Cockerell and nigromaculata (Cockerell and Eliot, 1905), which was Eliot, 1905: 40^1. based on the four specimens from La Jolla, California in Doris nigromaculata Cockerell in Cockerell and Eliot, June 1967. This description is accompanied by and ref¬ 1905: 40-41. erences a composite figure, also labeled as Doriopsilla Doridopsis nigromaculata (Cockerell in Cockerell and nigromaculata, but which was comprised solely of his Eliot, 1905).—Cockerell, 1908: 106. earlier illustrations (eventually labelled as D. rowena) of Dendrodoris vidua [non Bergh, 1878] .—O'Donoghue, “D. puerteeitensis” from the Gulf of California (some of 1926: 212. which are reproduced here in Figures 11-13). In the text Dendrodoris nigromaculata (Cockerell in Cockerell and of this description Lance states “it is likely that the pres¬ Eliot, 1905).—O’Donoghue, 1926: 213; Steinberg, ent species and an unnamed one, frequently encoun¬ 1961: 59. Lance, 1982: 29. tered in the Gulf of California, will eventually prove to Dendrodoris sp. Lee and Brophy, 1969: 20. be nonspecific.” This indicates that Lance wrote this Dendrodoris sp. Behrens, 1980: 100; Behrens and description after June 1967, but prior to receiving a copy Gatewood, 1986: 139, 142. of Marcus and Marcus (1967), and also suggests that Dendrodoris sp. b McDonald and Nybakken, 1980: 54; after receiving the latter, he considered D. rowena a McDonald, 1983: 171. junior synonym of D. nigromaculata. Further, as men¬ Dendrodoris sp. 3 Behrens, 1991: 72; Goddard, 2004: tioned above, hatching juveniles from La Jolla possess 1957, 1959, 1963. the same notal spicule complement of rods and forks Dendrodoris behrensi Millen and Bertseh, 2005: 189- as adults from the Gulf of California (Figures 13 and 199; Goddard, 2005: 201-211; Behrens and Hennosillo, 16). Although morphologically the specimens from La 2005: 86; California Academy of Sciences and Goddard, Jolla are very similar to D. rowena from the Gulf of 2013: worksheets for Hill St. and So. Casa Reef. California, Lance never applied the name D. nigromaculata to the latter, and in conversation with at least one colleague, maintained that the two were probably distinct (T.M. Type Material: Gosliner, personal communication to JG, 12 Dec 2014). Doris nigromaculata - Holotype: La Jolla, California, Lance (1982) illustrated a hatching juvenile, labeled as July 1901 (NHMUK 1904.7.7.1), dissected by Sir C. “Dendrodoris nigromaculata (Cockerell in Cockerell and Eliot in 1905. J.H.R. Goddard and A. Valdes, 2015 Page 39 Figures 17-21. Dendrodoris nigromaculata. 17. Living adult, 22 imn long, from Bahia Falsa, Baja California, May 2001. Included as a paratype of Dendrodoris behrensi by Millen and Bertsch (2005). 18. Adult, no date or locality (Lance Collection, Species Folder: White Porostome Spotted). 19. Egg mass, sketch labelled as South Casa Reef, La Jolla, 20 July 1974, but Lance Field Accounts indicate adults were from Hill Street, San Diego, 19 July 1974 (Lance Collection, Species Folder: White Porostome Spotted). 20. Recently hatched juvenile, Aug. 1974 (pencil sketch in Lance Collection, Species Folder: White Porostome Spotted). 21. Juve¬ nile, 570 pm long, four days after hatching, June 2001. From egg mass laid by adult from Bahia Falsa, Baja California, May 2001 (Goddard 2005, as Dendrodoris behrensi). Dendrodoris behrensi - Holotype: California, 14 Feb are sparse and consist of straight to slightly curved rods 1988 (CASIZ 69303). Paratypes: California, 14 Feb only, some of which are irregular in outline (Figures 1, 1988 (CASIZ 171658); San Quintfn, Mexico, 27 May 20-21). 2001 (CASIZ 171659), San Quintrn, Mexico, 27 May 2001 (CASIZ 171660). External Morphology: The external morphology of D. nigronmculata was described by Cockerell and Eliot Anatomy; The anatomy of Dendrodoris nigromaculata (1905) and Millen and Bertsch (2005), with additional was described by Cockerell and Eliot (1905) and Millen details presented by Lance (Lance Collection, Species and Bertsch (2005). As noted by Cockerell and Eliot Folder: White Porostome Spotted). Adults grow to (1905), Lance (Lance Collection, Species Folder: White 27 mm long and are distinguished externally by their trans¬ Porostome Spotted), and Goddard (2005), notal spicules lucent white ground color and chocolate brown blotches. Page 40 THE NAUTILUS, Vol. 129, No. 1 the larger of which are usually clustered into three or four groups centered mid-dorsally, and the smaller of which are scattered toward the edges of the dorsum (Figures 17-18). Pacific Grove, CA i' Jk Development: As described by Goddard (2005) and Carmel Bay, CA Point Dume, Malibu, CA Naples Reef. Santa Barbara Co.. CA Lance (Lance Collection, Species Folder: Wliite Porostome — Santa Monica Bay, CA San Miguel Island, CA— Redondo Beach, CA Spotted) Dendrodoris nigromaculata has arnetamorphie Anacapa Island, CA W .A-Palos Verdes Peninsula, CA direct development in short, stout egg ribbons laid on edge Santa Cruz Island, CA t Jy—Laguna Beach, CA Santa Catalina Island, CA y La Jolla, CA in a loose coil of only a turn or two (Figure 19). Juveniles San Clemente Island, CA Point Loma, San Diego, CA (Figures 20-21) hatch after an embryonic period of 38 days Bahia Falsa, San Quintin, BC Islas Coronados, BC at 16-19° C and are about 510 microns long. Islas San Benito, BC- \ Geographic Distribution: Dendrodoris nigromaculata is known from the Monterey Peninsula, California south V/' to the San Benitos Islands, Baja California (McDonald, PACIFIC OCEAN 1983; Behrens and Gatewood, 1986; Millen and Bertseh, 2005) (Figure 23). Remarks: Bergh (1878) described the species Doiiojms Figure 23. Map showing collection localities of Dendrodoris vidua based on specimens collected from Tahiti, French nigromaculata. Polynesia. The illustrations of the live animal (Bergh, 1878: pi. 1, figs. 17-20) represent an elongate Dendrodoris with a white background color and numerous black spots all over the dorsum, larger near the center of the animal, in La Jolla. Cockerel and Eliot (1905) noted some differ¬ and small dorsal tubercles. Based on the body shape and ences between their specimen and Bergh s (1878) original coloration, D. vidua is most likely a senior synonym of description in several details, including the coloration Dendrodoris elongate Baba, 1936. Cockerel and Eliot and mantle margin width. Cockerel and Eliot (1905) also (1905) tentatively reported this species from California as inchoated that their record was “suspicious” considering Doridopsis vidua (?), based on a single specimen collected the geographic distance between the type locality and California, thus they introduced Cockerell’s new name Doridopsis nigromaculata in case the specimen was ulti¬ mately proven to belong to a distinct species. As men¬ Jj tioned above, examination of the specimen studied by If v Cockerel and Eliot (1905) and therefore the holotype I, -Puerto Pertasco, Son. & of D. nigromaculata (NHMUK 1904.7.7.1) revealed San Quintin. BC' —-ipercebu, qc -PuerteCitos. BC that it is a species of Dendrodoris. The holotype of El Campo. BCS—^ ' - . Bahia de Los Angeles. BC • A D. nigromaculata has three pairs of large dark spots and Guaymas. Sort. several smaller spots irregularly distributed (Figure 1), <\ -v>- iS.la CerTalvo. BCS I \ very different from the original description of Doriopsis Punta Pcrico. BCS-—' . T—Mazatten. S_ in GULF OF \ . ( vidua by Bergh (1878) and references to D. nigromaculata MEXICO % 7 by other authors, but nearly identical to the original description of D. behrensi by Millen and Bertseh (2005). Lance was aware of this species as early as 1961 (or possibly even 1953) and referred to it in his notes and illustrations first as the “erenulate dorid,” then as the CARIBBEAN PACIFIC OCEAN SEA “white dendrodorid” or “brown-spotted Dendrodoris” (Lance Collection, Species Folder: White Porostome Spotted), and in his field accounts as “Dendrodoris sp. true Pacific coast of Costa Rica IslasI sSlae Ucavsa., PPaannaammaa /x ‘ dendrodoris” (19 July 1974, Hill St) and “Dendrodoris sp. 3 [following Behrens 1991] white porostome” (26 Apr. 1998, So Casa Reef). As evidenced by an undated, hand¬ written description and pen and ink illustration of an adult (Figure 18), Lance started to formally describe it under the manuscript name “Dendrodoris barbarensis,” based on five specimens collected in the 1950s and 60s from Naples Reef, Santa Barbara County; Point Lima, San Diego; and tire Coronado Islands. Lances folder for this species con¬ Figure 22. Map showing collection localities of Doriopsilla tains no evidence diat he ever associated Cockerell and rowena. Eliots (1905) description of D. nigromaculata widi it.