Records of the Western Australian Museum Supplement No. 69: 83-93 (2006).

Marine collected by the Steamer Albatross from the Philippines in 1908

Terrence M. Gosliner Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103 USA. E-mail: [email protected]

From 1887 until 1925, the United States Fish generally striking and the majority of the fifty-three Commission's Steamer Albatross conducted illustrations can be identified to (Table 1). pioneering marine explorations in the Pacific and Several of these taxa have been described only Indian . Most of the results of these recently and at least two of the identifiable taxa expeditions have been published in accounts of the remain undescribed. The significance of these expeditions and through numerous scientific papers paintings has only been briefly documented dealing with particular marine taxa. The most (Gosliner and Tyler 1990) and the only illustration ambitious expedition of the steamer's service was a published was presented by Elwood et al. (2000) in two-and-a-half year expedition to the Philippine their original description of Aldisa albatrossae. Islands, beginning in 1907, in the wake of the Included in this paper are many of Ito's paintings United States occupation of the Philippines of species that are readily identifiable together with following the Spanish American War. All of the colour photographs of living of these collections of the gastropods illustrated here species. Also included are some of the remaining occurred during 1908. paintings of Ito's that could not be readily Six civilian scientists participated in this identified. Only thirteen of the species depicted expedition (http://www.nmnh.sLeduI vertIfishesI cannot be linked with described, or known but albatross lexped.html), including the noted undescribed, species. Of these only two cannot be malacologist Paul Bartsch of the Smithsonian identified or tentatively placed in known genera. Institution's National Museum of Natural History. The vast majority of taxa represented are While the National Museum holds many collections opisthobranch gastropods, but also included is one from this expedition, there is only a fragmentary species of vetigastropod, three species of record of the opisthobranch and other slug-like caenogastropod and one pulmonate. Members of mollusks from the expedition and it appears no these vetigastropod and caenogastropod taxa have other molluscan collections from the expedition are internal shells and are often included in collections resident in the Smithsonian's collections, based on of opisthobranchs, as are the unshelled pulmonates extensive searching of those collections. None of the of the Onchidiidae. Of the opisthobranch preserved opisthobranch specimens are currently gastropods one is a species of cephalaspidean, three housed in the collections and are presumably lost. are anaspideans, three are notaspideans and the While I was conducting a postdoctoral fellowship remainder are nudibranchs. The tentative in the Division of Mollusks at the Smithsonian identification of each taxon is included in Table l. Institution in 1982, Diane Tyler, then a curatorial In order to demonstrate the diagnostic nature of assistant in the Division, brought to my attention a the illustrations a series of them are reproduced series of original paintings, kept in the Mollusk here together with photographs of living animals of Division, of opisthobranchs and other mollusks the same species from different localities collected in 1908 as part of the Albatross Expedition throughout the tropical Indo-Pacific (Figures. 1-6). to the Philippines. These paintings were undertaken Figures 6K and 6L depict two species that cannot be by the expedition artist Kumataro Ito. While many readily identified with any described or known of Ito's fish paintings have been published (Springer undescribed species. The first one appears to be an 1999) and are accessible via the internet (http: II undescribed species of Tritonia while the second www.nmnh.sLedu/vert/fishes lalbatross I appears to be a species of Dendrodoris. It is illusito.html), these molluscan illustrations have interesting to note that the illustration of the never been published. In the absence of preserved Dendrodoris includes a drawing of the egg mass of specimens, the mollusc illustrations constitute the this species. From the presence of relatively few, only record of gastropod molluscs from the large eggs, it appears that this species has direct expedition that are known to exist. rather than planktonic development. Direct The scientific accuracy of these paintings is development appears to be very uncommon in 84 T.M. Gosliner

Table 1 List of marine gastropods illustrated by Ito from the Albatross expedition to the Philippines.

Vetigastropoda Rostanga sp. unidentified dorid Scutus unguis (Linnaeus, 1758) orange dorid Dendrodorididae Dendrodoris denisoni (Angas, 1864) sp. Dendrodoris nigra (Stimpson, 1855) nigra Blainville, 1824 Dendrodoris sp. Coriocella sp. Phyllidiidae Pulmonata Phyllidiella nigra (van Hasselt, 1824) Onchidiidae Phyllidiella pustulosa (Cuvier, 1804) Onchidium sp. gracillimum Semper in Bergh, 1876 Cephalaspidea Ceratosoma tenue Abraham, 1876 Aglajidae (Gray, 1827) Chelidonura varians Eliot, 1903 Chromodoris annae Bergh, 1877 Anaspidea Chromodoris aureopurpurea Collingwood, 1881 Aplysiidae Chromodoris magnifica (Quoy and Gaimard, 1832) Aplysia oculifera A. Adams and Reeve, 1850 Chromodoris striatella Bergh, 1876 Bursatella leachii leachii de Blainville, 1817 Glossodoris cincta (Bergh, 1888) Stylocheilus longicauda (Quoy and Gaimard, 1825) Hypselodoris bullocki (Collingwood, 1881) Sacoglossa Polyceratidae Caliphyllidae Nembrotha chamberlaini Gosliner and Behrens, 1997 Cyerce sp. Roboastra luteolineata (Baba, 1936) Plakobranchidae Aegiridae Elysia sp. serenae (Gosliner and Behrens, 1997) Plakobranchus ocellatus van Hasselt, 1824 Pleurobranchoidea Dendronotoidea Pleurobranchidae Berthellina delicata (Pease, 1861) pelagica Linnaeus, 1758 Pleurobranchaea sp. Tritoniidae Pleurobranchus peroni Cuvier, 1804 Tritonia sp. Nudibranchia Arminoidea Anthobranchia Arminidae Armina sp. 1 Hexabranchidae Armina sp. 2 Hexabranchus sanguineus (Riippell and Leuckart, 1828) Dermatobranchus sp. 1 Dorididae Dermatobranchus sp. 2 Aldisa albatrossae Elwood, Valdes and Gosliner 2000 Aeolidoidea Facelinidae Asteronotus caespitosus (van Hasselt, 1824) Pteraeolidia ianthina (Angas, 1864) Atagema intecta (Kelaart, 1858) Atagema spongiosa (Kelaart, 1858) Atagema sp. Discodoris boholiensis Bergh, 1877 Halgerda carlsoni Rudman, 1978 Jorunna funebris (Kelaart, 1858)

tropical Indo-Pacific taxa (personal observation). margin. The outline of the internal shell can clearly Inclusion of life-history data increases the value of be seen in Ito's illustration. these important illustrations that document a small Lamellaria sp. This species is depicted in two sample of the marine opisthobranch biodiversity of paintings, one of a dorsal view and the second of the Philippine Islands. While many of the taxa the ventral aspect. The pink mantle with a included here are relatively common in the longitudinal band is distinctive. The ventral view Philippine Islands, others have not been collected with a broad foot and deeply divided elongate oral there since these illustrations were made. tentacles clearly indicate that this is a species of Remarks for each species indicate the basis for the Lamellaria. The systematics of the Indo-Pacific identifications that have been determined within members of this taxon remain poorly known and the present study. the degree of variability and species limits require Scutus unguis (Linnaeus, 1758). (Figures lA, B). This additional detailed study. vetigastropod is characterized by having a Coriocella nigra Blainville, 1824. (Figures le, D). The uniformly black pigmented mantle with a frilled uniformly velvety black color of this species with a Gastropods of the 1908 Albatross Expedition 85

Figure 1 A., B. Scutus l/Ilgl/is (Linnaeus, 1758), photo: Bagabag Island, Madang, Papua ew Guinea. c., D. Coriocelll/ Iligra Blainville, 1824, photo: Jerry Alien, Batangas, Luzon Philippines. E., F. Chelidolll/ra vl/ril/lls Eliot, 1903, photo: Olango, Cebu, Philippines. G.,H. Stylocheill/s longicl/udl/ (Quoy and Gaimard, 1825), photo: Madang, Papua ew Guinea. 1.,]. Bl/rsatelll/ leachii lel/chii de Blainville, 1817, photo: Salt Rock, atal, South Africa. 86 T.M. Gosliner tuberculate mantle is distinctive. The four dorsal There are no external characteristics that distinguish lobes of the mantle are also characteristic of this species of Berthellina. As such, the identification species. made here is based exclusively on the fact that all Coriocella sp. The lighter mantle color and shorter specimens of Berthellina from the Philippines mantle lobes differ from the above species. The examined to date are B. delicata, based on their degree of variation present in species of Coriocella genitalia. requires further investigation. Pleurolmmchus peroni Cuvier, 1804. (Figures 2K, Onchidium sp. The tuberculate body with elongate L). The finely reticulate orange mantle with tentacular eye-stalks and bilobed velum indicate scattered opaque white spots is characteristic of that this figure illustrates a species of Onchidiidae, Pleurobranchus peroni. most likely a species of Onchidium. Pleurobranchaea sp. The illustration of this species Chelidonura varians Eliot, 1903. (Figures 1E, F). The with an extruded buccal mass, a pallid color and a black body color with blue lines on the parapdodia wrinkled mantle indicate the specimen was dead or and dorsum and elongate posterior mantle lobes moribund at the time it was illustrated. The widely readily identify this species as Chelidonura varians. separated and lateral gill clearly Stylocheilus longicauda (Quoy and Gaimard, 1825). indicate that it is a species of Pleurobranchaea. The (Figures 1G, H). The pale yellowish body with short most common member of this in Philippine papillae and elongate foot indicate that this is likely waters is P. brockii Bergh, 1897 (present study) and Stylocheilus longicauda. Typically living specimens this is most likely the species depicted by Ito. have prominent blue ocelli that are not readily Hexabranchus sanguineus (Rlippell and Leuckart, evident in Ito's illustration. 1828). (Figures 3A, B). The swimming posture of Bursatella leachii leachii de Blainville, 1817. this large orange dorid with fimbriate oral tentacles (Figures 1I, J). This sea hare is readily distinguished indicate that it can only be the common Spanish by its mantle ocellae and dense covering of ramified Dancer, Hexabranchus sallguineus. papillae. Aldisa albatrossae Elwood, Valdes and Gosliner Aplysia oculifera A. Adams and Reeve, 1850. 2000. (Figures 3C D). Ito's drawing of this species (Figures 2A, B). This is one of the few Indo-Pacific has been previously identified as Aldisa albatrossae species of Aplysia that is capable of swimming. The (Elwood, Valdes and Gosliner 2000). widely open mantle lobes in Ito's left illustration Asteronotus cespitosus (van Hasselt, 1824). (Figures indicate that the is in swimming posture. 3E, F). The brownish body with mid-dorsal ridge, The black ocellae of the living animal are not strong, irregular tubercles and concentric rings of evident in Ito's illustration. marginal tubercles readily identify this illustration Elysia sp. (Figures 2C, D). The animal depicted in as Asteronotus cespitosus. Ito's illustration represents an undescribed species Atagema in tecta (Kelaart, 1858). (Figures 3G, H). of £lysia that has been found sporadically in the The brown, tuberculate body with mid-dorsal, Philippines and Papua New Guinea (present study). longitudinal opaque white stripe is characteristic of This species is similar to the common Atagema intecta. circumtropical species Elysia omata (Swainson, Atagema spongiosa (Kelaart, 1858). (Figures 31, J). 1840) and like that species has white and black The finely granular surface of the body with distinct parapodial margins. Elysia ornata has an additional holes resembling the oscula of indicates orange marginal line that is absent in E. sp. Elysia that this species is most likely Atagema spongiosa, sp. also has a y-shaped opaque white marking on although the possibility that it represents an the head that is absent in E. ornata. undescribed species cannot be completely Cyerce sp. (Figures 2E, F). This represents an eliminated. undescribed species of Cyerce. The yellow to Atagema sp. (Figures 3K, L). This species with a orange tips with stippled circles of brown reticulate brownish to pink notum and a smooth pigment are characteristic of this species. longitudinal band probably represents the Plakobranchus ocellatus van Hasselt, 1824. (Figures undescribed species of Atagema shown in the 2 G, H). The widely separated rhinophores and adjacent photo. ocellae on the mantle are characteristic of Discodoris boholiensis Bergh, 1877. (Figures 4A, B). Plakobranclws ocellatus. A second specimen is also The brownish color with branching, lighter, represented in Ito's paintings, but this individual radiating lines near the mantle margins suggest that has less prominent ocellae. Ito's drawing represents Discodoris bol1Oliellsis. Berthellina delicata (Pease, 1861). (Figures 2I, J). The Halgerda carlsoni Rudman, 1978. (Figures 4C D). uniformly smooth orange mantle with darker The white body with ridges punctuated by large viscera visible through the mantle is characteristic orange dots and smaller orange specks is of species of Berthellina. Berthellina delicata is the characteristic of Halgerda carlsoni. The gill and common member of the genus found in the western rhinophores are also ornamented with dark brown Pacific (Cervera and Gosliner, in preparation). to black specks. Gastropods of the 1908 Albatross Expedition 87

Figure 2 A., B. Aplysia oClIlifera A. Adams and Reeve, 1850, photo: Midway Atoll. c., D. Elysia sp., photo: Madang, Papua New Guinea. E., F. Cyerce sp., photo: Madang, Papua New Guinea. G., H. Plakobranclws ocellallls van Hasselt, 1824, photo: Madang, Papua ew Guinea. 1., J. Berthe/lina delicata (Pease, 1861), photo: Madang, Papua ew Guinea. K., L. Plellrobranchlls peroni Cuvier, 1804, photo: Tulamben, Bali, Indonesia. 88 T.M. GosIiner

Figure 3 A., B. Hexabranchus sanguineus (Ruppell and Leuckart, 1828), photo: Koloa, Kauai, Hawaii. c., D. Aldisa albatrossae Elwood, Valdes and Gosliner 2000, photo: Robert Bolland, Okinawa, Ryukuyu Islands, Japan. E., F. Asteronotus cespitosus (van Hasselt, 1824), photo: Madang, Papua New Guinea. G., H. Atagema in tecta (Kelaart, 1858), photo: Batangas, Luzon, Philippines. I., J. Atagema spongiosa (Kelaart, 1858), photo: Madang, Papua New Guinea. K., 1. Atngema sp., photo: Batangas, Luzon Philippines. Gastropods of the 1908 Albatross Expedition 89

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Figure 4 A., B. Discodoris boholiensis Bergh, 1877, photo: Batangas, Luzon Philippines. c., D. Halgerda carlsoni Rudman, 1978, photo: Madang, Papua New Guinea. E., F. jorUlllla ftl11ebris (Kelaart, 1858), photo: Louisiade Archipelago, Papua ew Guinea. G., H. Dendrodoris dellisolli (Angas, 1864), photo: Salt Rock, atal, South Africa. I., J. Dendrodoris nigra (Stimpson, 1855), photo: Madang, Papua ew Guinea. 90 T.M. Gosliner

Figure 5 A, B. Chromodoris annae Bergh, 1877, photo: Batangas, Luzon, Philippines. c., D. Chromodoris magnifica (Quoy and Gaimard, 1832), photo: Batangas, Luzon, Philippines. E., F. Chromodoris striatella Bergh, 1876, photo: Pauline Fiene, Adonara, Indonesia. G., H. Glossodoris cincta (Bergh, 1888), photo: Tulamben, Bali, Indonesia. 1., J. Hypselodoris bullocki (Collingwood, 1881), photo: Puerto Galera, Mindoro, Philippines. K., L. Ceratosoma gracillimum Semper in Bergh, 1876, photo: Batangas, Luzon, Philippines. Gastropods of the 1908 Albatross Expedition 91

10rIllllla fllllcbris (Kclaart, 1858). (Figures 4E, F). base, orange and gilL opaque white The white body with black rings, black gill and marginal band and reddish purple body color. rhinophores is characteristic of 101'11111111 jilllclJris. Ccmtosolilll gmcillillllllll Semper in Bergh, 1876. Rosfllllga sp. The uniformly red-orange granular (Figures 5K, L). The relatively uniform reddish body indicates that this dorid is most likely a body color with numerous spots of mottling and species of Rostallga. color indicate that Ito's illustration of this species is unidentified dorid. This dorid with a granular most likely Ceratosollla gracillilillllil. texture, unipinnate gill branches and anastanlosing Cemtosollla tClllle Abraham, 1876. The reddish reddish lines cannot readily be placed in any mottled body with relatively little expansion of the specific group of cryptobranch dorids. mantle and a purple marginal line suggest that this DClldrodoris dCllisolli (Angas, 1864). (Figures 4C, species is most likely Ceratosollla tell Ill'. H). The brownish body with bright blue spots, large Cemtos0111a trilolJtltulII (Cray, 1827). The reddish gill and rhinophores strongly suggest that Ito's body, wide mantle lobes suggest that this species is drawing represents DClldrodoris dCII isoll i. most likely Ceratosolilil trilobatlllil. DClldrodoris Iligra (Stimpson, 1855). (Figures 41, 1). Aegires sercllae (Cosliner and Behrens, 1997). Ito's drawing of an animal with a black notum and (Figures 6A, B). The distinctive greenish color, large a lighter marginal band, sparsely pinnate gill and branchial lobes and smooth rhinophores are white apices of the rhinophores strongly suggests characteristic for Aegires sercllac. that this is DClldrodoris lIigra. NC111brotlIa clIalllberlailli Cosliner and Behrens, DClldrodoris sp. (Figure 6L). It appears that this 1997. (Figures 6C. D). The white body color with mottled dorid with large rhinophores represents a brownish pigment on the center of the notum and a species of DClldrodoris. The egg mass (drawn by Ito red gill are characteristic for the common Philippine and shown for this species in Figure 6L) indicates species, NClllbrotha clllllllbcrlailli. an egg mass with few, large eggs suggesting that ?Roboastra IlItcolillcata (Ba ba, 1936). This this species has direct development. illustration of Ito's suggests that the black animal orange dorid. This reddish orange dorid with a with numerous longitudinal lines of yellow may be pustulose notum and no readily evident dorsal gill Roboastra IlIteolillmta. The large oral tentacles that may be a species of Phyllidiidae. are characteristic of this species are not evident in PlItfllidiella lIigra (van Hasselt, 1824). The black Ito's drawing, suggesting it may also represent a body with large rounded lighter tubercles suggests species of Talllbja or that this characteristic was that this species is likely Phyllidiella lIigra. omitted from the drawing. PlIyl/idiel/a pIIstlllosa (Cuvier, 1804). The low Scyllaea pclagica Linnaeus, 1758. (Figures 6E, F). irregular, pinkish tubercles of this black animal This species is quite distinctive with its four large suggest that it is likely PlIyllidiella pIIstlllosa. mantle lobes and expanded rhinophore sheaths. ClIro1110doris allllae Bergh, 1877. (Figures SA, B). There is little doubt that this is SCljllaca pelagica. The distinctive pattern of a blue body without Tritollia sp. (Figure 6K). This white species with longitudinal black bands with orange margin is darker pigment on the head, appears to be a species characteristic of Chr01110doris a111 liIe. of Tritollia, but cannot be identified with any ClIrolllOdoris 111agllifiCl7 (Quoy and Caimard, 1832). known species. (Figures 5C, D). The white body with black A 1'111 ilia sp. 1. (Figures 6(;, H). This species with a longitundinal lines, orange margin, orange gill and red body and numerous opaque white longitudinal rhinophores is distinctive for Chrolllodoris lIIagllifiCl7. lines appears to be undescribed. The specimen Chrolllodoris striatella Bergh, 1876. (Figures SE, F). depicted by Ito closely matches a specimen A white body with a series of brown lines or collected from Anilao, Batangas, Philippines varying intensity is a distinguishing feature of (Figures 6H). ClIrolilOdoris striatclla, as are the orange marginal Arlllilla sp. 2. Ito illustrated a second species of spots and rhinophores with opaque white spots. Arlllilla with uniform orange pigment and low ClIrolllodoris allrcopllrpllrca Collingwood, 1881. longitudinal ridges. It cannot be readily identified This species with a white body and brownish notal with any known species. spots and a blue margin is similar to that of OCrJllatolmmclllls sp. 1. Ito illustrated a dark gray Chrolllodoris allreopllllctata and C. rllfo111aclllata species of Dcrlllatobmllclllls with lighter Pease, 1871. The egg mass is also illustrated. longitudinal lines. This species cannot be readily Glossodoris cillcta (Bergh, 1888). (Figures. 5C, H). identified with any known species. The brownish body with blue and orange marginal Oerlllatobrallchlls sp. 2. A second species of bands and deeply indented central portions of the DCl'llliItolmmchlls was illustrated bv Ito. It is also mantle are characteristic of Glossodoris cillcta. gray in color with lighter longitudinal lines. It llypsclodoris blll/ock} (Collingwood, 1881). differs from the previous species in having more (Figures. 51, J). Ito's drawing of Hypselodoris prominent rhinophores that are ornamented with bllllocki is very characteristic with the elevated gill- yellow pigment. The same yellow pigment is found 92 T.M. Gosliner

Figure 6 A, B. Aegires serenae (Gosliner and Behrens, 1997), photo: Madang, Papua New Guinea. c., D. Nembrotha chamberlaini Gosliner and Behrens, 1997, photo: Batangas, Luzon Philippines. E., F. SC1Jllaea pelagica Linnaeus, 1758, photo: Madang, Papua New Guinea. G., H. Armina sp., photo: Batangas, Luzon, Philippines. 1., J., Pteraeolidia ianthina (Angas, 1864), photo: Batangas, Luzon, Philippines. K. Tritonia sp. L. Dendrodoris sp. Gastropods of the 1908 Albatross Expedition 93 along the margin of the foot. This species cannot be knowledge of Philippine opisthobranchs and readily identified with any known species. assisted in our various field expeditions. I also wish Pteraeolidia ianthina (Angas, 1864). (Figures 61, J). to thank Bob Bolland who has advanced our The elongate body shape, elevated ceratal clusters knowledge of the opisthobranch fauna from the and elongate oral tentacles make Ito's illustration nearby Ryukuyu Islands of Japan. This work was immediately identifiable with Pteraeolidia iantlzina. supported by a National Science Foundation PEET While tentative identifications are made here for grant DEB 0329054 Phylogenetic systematics of all of the material illustrated by Ito during the dorid nudibranchs to the author and Angel Valdes. Albatross Expedition, only 35 of Ito's 53 illustrations are included in the plates presented here. All of the images, including those not included in this paper REFERENCES will be posted on the PEET webpage: Elwood, H. A. Valdes and T. Gosliner 2000. Two new http://www.calacademy.org/research/izg/ species of Aldisa Bergh, 1878 (, nudibranchsl . Nudibranchia) from the tropical lndo-Pacific. Proceedings ofthe California Academy ofSciences 52: 171­ 181. ACKNOWLEDGEMENTS Gosliner, T. and D. Tyler. 1990. Marine Gastropods collected by the Steamer Albatross in 1908. Western I wish to thank Diane Tyler for first bringing the Society of Malacologists Annual Report 22: 15. existence of Ito's paintings of mollusks to my Springer, V. 1999. Kumataro Ito, Japanese Artist on Board attention and for her enthusiastic interest in the the u.s. Bureau of Fisheries Steamer Albatross During historical importance of these surviving documents the Philippine Expedition, 1907-1910. Marine Fisheries of the Albatross expedition. I also wish to thank Review 61: 42-57. many individuals who have contributed to the