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

Marine Gastropoda collected by the Steamer Albatrossfrom the Philippines in 1908

Terrence M. GoslinerDepartment of Invertebrate Zoology and Geology, California Academy of Sciences,

875 Howard Street, San Francisco, CA 94103 USA. E-mail: tgosliner@calacademy.org

From 1887 until 1925, the United States FishCommission's Steamer Albatross conductedpioneering marine explorations in the Pacific andIndian Oceans. Most of the results of theseexpeditions have been published in accounts of theexpeditions and through numerous scientific papersdealing with particular marine taxa. The mostambitious expedition of the steamer's service was atwo-and-a-half year expedition to the PhilippineIslands, beginning in 1907, in the wake of theUnited States occupation of the Philippinesfollowing the Spanish American War. All of thecollections of the gastropods illustrated hereoccurred during 1908.

Six civilian scientists participated in thisexpedition (http://www.nmnh.sLedu I vert Ifishes Ialbatross lexped.html), including the notedmalacologist Paul Bartsch of the SmithsonianInstitution's National Museum of Natural History.While the National Museum holds many collectionsfrom this expedition, there is only a fragmentaryrecord of the opisthobranch and other slug-likemollusks from the expedition and it appears noother molluscan collections from the expedition areresident in the Smithsonian's collections, based onextensive searching of those collections. None of thepreserved opisthobranch specimens are currentlyhoused in the collections and are presumably lost.While I was conducting a postdoctoral fellowshipin the Division of Mollusks at the SmithsonianInstitution in 1982, Diane Tyler, then a curatorialassistant in the Division, brought to my attention aseries of original paintings, kept in the MolluskDivision, of opisthobranchs and other molluskscollected in 1908 as part of the Albatross Expeditionto the Philippines. These paintings were undertakenby the expedition artist Kumataro Ito. While manyof Ito's fish paintings have been published (Springer1999) and are accessible via the internet (http: I Iwww.nmnh.sLedu/vert/fishes lalbatross Iillusito.html), these molluscan illustrations havenever been published. In the absence of preservedspecimens, the mollusc illustrations constitute theonly record of gastropod molluscs from theexpedition that are known to exist.

The scientific accuracy of these paintings is

generally striking and the majority of the fifty-threeillustrations can be identified to species (Table 1).Several of these taxa have been described onlyrecently and at least two of the identifiable taxaremain undescribed. The significance of thesepaintings has only been briefly documented(Gosliner and Tyler 1990) and the only illustrationpublished was presented by Elwood et al. (2000) intheir original description of Aldisa albatrossae.

Included in this paper are many of Ito's paintingsof species that are readily identifiable together withcolour photographs of living animals of thesespecies. Also included are some of the remainingpaintings of Ito's that could not be readilyidentified. Only thirteen of the species depictedcannot be linked with described, or known butundescribed, species. Of these only two cannot beidentified or tentatively placed in known genera.The vast majority of taxa represented areopisthobranch gastropods, but also included is onespecies of vetigastropod, three species ofcaenogastropod and one pulmonate. Members ofthese vetigastropod and caenogastropod taxa haveinternal shells and are often included in collectionsof opisthobranchs, as are the unshelled pulmonatesof the Onchidiidae. Of the opisthobranchgastropods one is a species of cephalaspidean, threeare anaspideans, three are notaspideans and theremainder are nudibranchs. The tentativeidentification of each taxon is included in Table l.

In order to demonstrate the diagnostic nature ofthe illustrations a series of them are reproducedhere together with photographs of living animals ofthe same species from different localitiesthroughout the tropical Indo-Pacific (Figures. 1-6).Figures 6K and 6L depict two species that cannot bereadily identified with any described or knownundescribed species. The first one appears to be anundescribed species of Tritonia while the secondappears to be a species of Dendrodoris. It isinteresting to note that the illustration of theDendrodoris includes a drawing of the egg mass ofthis species. From the presence of relatively few,large eggs, it appears that this species has directrather than planktonic development. Directdevelopment 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.

VetigastropodaFissurellidae

Scutus unguis (Linnaeus, 1758)Caenogastropoda

VelutinidaeLamellaria sp.Coriocella nigra Blainville, 1824Coriocella sp.

PulmonataOnchidiidae

Onchidium sp.OpisthobranchiaCephalaspidea

AglajidaeChelidonura varians Eliot, 1903

AnaspideaAplysiidae

Aplysia oculifera A. Adams and Reeve, 1850Bursatella leachii leachii de Blainville, 1817Stylocheilus longicauda (Quoy and Gaimard, 1825)

SacoglossaCaliphyllidae

Cyerce sp.Plakobranchidae

Elysia sp.Plakobranchus ocellatus van Hasselt, 1824

PleurobranchoideaPleurobranchidae

Berthellina delicata (Pease, 1861)Pleurobranchaea sp.Pleurobranchus peroni Cuvier, 1804

NudibranchiaAnthobranchia

DoridoideaHexabranchidae

Hexabranchus sanguineus (Riippell and Leuckart, 1828)Dorididae

Aldisa albatrossae Elwood, Valdes and Gosliner 2000Discodorididae

Asteronotus caespitosus (van Hasselt, 1824)Atagema intecta (Kelaart, 1858)Atagema spongiosa (Kelaart, 1858)Atagema sp.Discodoris boholiensis Bergh, 1877Halgerda carlsoni Rudman, 1978Jorunna funebris (Kelaart, 1858)

tropical Indo-Pacific taxa (personal observation).Inclusion of life-history data increases the value ofthese important illustrations that document a smallsample of the marine opisthobranch biodiversity ofthe Philippine Islands. While many of the taxaincluded here are relatively common in thePhilippine Islands, others have not been collectedthere since these illustrations were made.

Remarks for each species indicate the basis for theidentifications that have been determined withinthe present study.Scutus unguis (Linnaeus, 1758). (Figures lA, B). Thisvetigastropod is characterized by having auniformly black pigmented mantle with a frilled

Rostanga sp.unidentified doridorange dorid

DendrodorididaeDendrodoris denisoni (Angas, 1864)Dendrodoris nigra (Stimpson, 1855)Dendrodoris sp.

PhyllidiidaePhyllidiella nigra (van Hasselt, 1824)Phyllidiella pustulosa (Cuvier, 1804)

ChromodorididaeCeratosoma gracillimum Semper in Bergh, 1876Ceratosoma tenue Abraham, 1876Ceratosoma trilobatum (Gray, 1827)Chromodoris annae Bergh, 1877Chromodoris aureopurpurea Collingwood, 1881Chromodoris magnifica (Quoy and Gaimard, 1832)Chromodoris striatella Bergh, 1876Glossodoris cincta (Bergh, 1888)Hypselodoris bullocki (Collingwood, 1881)

PolyceratidaeNembrotha chamberlaini Gosliner and Behrens, 1997Roboastra luteolineata (Baba, 1936)

AegiridaeAegires serenae (Gosliner and Behrens, 1997)

CladobranchiaDendronotoidea

ScyllaeidaeScyllaea pelagica Linnaeus, 1758

TritoniidaeTritonia sp.

ArminoideaArminidae

Armina sp. 1Armina sp. 2Dermatobranchus sp. 1Dermatobranchus sp. 2

AeolidoideaFacelinidae

Pteraeolidia ianthina (Angas, 1864)

margin. The outline of the internal shell can clearlybe seen in Ito's illustration.Lamellaria sp. This species is depicted in twopaintings, one of a dorsal view and the second ofthe ventral aspect. The pink mantle with alongitudinal band is distinctive. The ventral viewwith a broad foot and deeply divided elongate oraltentacles clearly indicate that this is a species ofLamellaria. The systematics of the Indo-Pacificmembers of this taxon remain poorly known andthe degree of variability and species limits requireadditional detailed study.Coriocella nigra Blainville, 1824. (Figures le, D). Theuniformly 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

tuberculate mantle is distinctive. The four dorsallobes of the mantle are also characteristic of thisspecies.Coriocella sp. The lighter mantle color and shortermantle lobes differ from the above species. Thedegree of variation present in species of Coriocellarequires further investigation.Onchidium sp. The tuberculate body with elongatetentacular eye-stalks and bilobed velum indicatethat this figure illustrates a species of Onchidiidae,most likely a species of Onchidium.Chelidonura varians Eliot, 1903. (Figures 1E, F). Theblack body color with blue lines on the parapdodiaand dorsum and elongate posterior mantle lobesreadily identify this species as Chelidonura varians.Stylocheilus longicauda (Quoy and Gaimard, 1825).(Figures 1G, H). The pale yellowish body with shortpapillae and elongate foot indicate that this is likelyStylocheilus longicauda. Typically living specimenshave prominent blue ocelli that are not readilyevident in Ito's illustration.Bursatella leachii leachii de Blainville, 1817.(Figures 1I, J). This sea hare is readily distinguishedby its mantle ocellae and dense covering of ramifiedpapillae.Aplysia oculifera A. Adams and Reeve, 1850.(Figures 2A, B). This is one of the few Indo-Pacificspecies of Aplysia that is capable of swimming. Thewidely open mantle lobes in Ito's left illustrationindicate that the animal is in swimming posture.The black ocellae of the living animal are notevident in Ito's illustration.Elysia sp. (Figures 2C, D). The animal depicted inIto's illustration represents an undescribed speciesof £lysia that has been found sporadically in thePhilippines and Papua New Guinea (present study).This species is similar to the commoncircumtropical species Elysia omata (Swainson,1840) and like that species has white and blackparapodial margins. Elysia ornata has an additionalorange marginal line that is absent in E. sp. Elysiasp. also has a y-shaped opaque white marking onthe head that is absent in E. ornata.Cyerce sp. (Figures 2E, F). This represents anundescribed species of Cyerce. The yellow toorange cerata tips with stippled circles of brownpigment are characteristic of this species.Plakobranchus ocellatus van Hasselt, 1824. (Figures2 G, H). The widely separated rhinophores andocellae on the mantle are characteristic ofPlakobranclws ocellatus. A second specimen is alsorepresented in Ito's paintings, but this individualhas less prominent ocellae.Berthellina delicata (Pease, 1861). (Figures 2I, J). Theuniformly smooth orange mantle with darkerviscera visible through the mantle is characteristicof species of Berthellina. Berthellina delicata is thecommon member of the genus found in the westernPacific (Cervera and Gosliner, in preparation).

T.M. Gosliner

There are no external characteristics that distinguishspecies of Berthellina. As such, the identificationmade here is based exclusively on the fact that allspecimens of Berthellina from the Philippinesexamined to date are B. delicata, based on theirgenitalia.Pleurolmmchus peroni Cuvier, 1804. (Figures 2K,L). The finely reticulate orange mantle withscattered opaque white spots is characteristic ofPleurobranchus peroni.Pleurobranchaea sp. The illustration of this specieswith an extruded buccal mass, a pallid color and awrinkled mantle indicate the specimen was dead ormoribund at the time it was illustrated. The widelyseparated rhinophores and lateral gill clearlyindicate that it is a species of Pleurobranchaea. Themost common member of this genus in Philippinewaters is P. brockii Bergh, 1897 (present study) andthis is most likely the species depicted by Ito.Hexabranchus sanguineus (Rlippell and Leuckart,1828). (Figures 3A, B). The swimming posture ofthis large orange dorid with fimbriate oral tentaclesindicate that it can only be the common SpanishDancer, Hexabranchus sallguineus.Aldisa albatrossae Elwood, Valdes and Gosliner2000. (Figures 3C D). Ito's drawing of this specieshas been previously identified as Aldisa albatrossae(Elwood, Valdes and Gosliner 2000).Asteronotus cespitosus (van Hasselt, 1824). (Figures3E, F). The brownish body with mid-dorsal ridge,strong, irregular tubercles and concentric rings ofmarginal tubercles readily identify this illustrationas Asteronotus cespitosus.Atagema in tecta (Kelaart, 1858). (Figures 3G, H).The brown, tuberculate body with mid-dorsal,longitudinal opaque white stripe is characteristic ofAtagema intecta.Atagema spongiosa (Kelaart, 1858). (Figures 31, J).The finely granular surface of the body with distinctholes resembling the oscula of sponges indicatesthat this species is most likely Atagema spongiosa,although the possibility that it represents anundescribed species cannot be completelyeliminated.Atagema sp. (Figures 3K, L). This species with areticulate brownish to pink notum and a smoothlongitudinal band probably represents theundescribed species of Atagema shown in theadjacent photo.Discodoris boholiensis Bergh, 1877. (Figures 4A, B).The brownish color with branching, lighter,radiating lines near the mantle margins suggest thatIto's drawing represents Discodoris bol1Oliellsis.Halgerda carlsoni Rudman, 1978. (Figures 4C D).The white body with ridges punctuated by largeorange dots and smaller orange specks ischaracteristic of Halgerda carlsoni. The gill andrhinophores are also ornamented with dark brownto 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 ocellalllsvan 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. Aldisaalbatrossae 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

Cl'

~ Q Q <> .0~. 4 0 <:> ._,,--. 0 4) • /

E .--._._~- ------~

G

89

Figure 4 A., B. Discodoris boholiensis Bergh, 1877, photo: Batangas, Luzon Philippines. c., D. Halgerda carlsoniRudman, 1978, photo: Madang, Papua New Guinea. E., F. jorUlllla ftl11ebris (Kelaart, 1858), photo: LouisiadeArchipelago, Papua ew Guinea. G., H. Dendrodoris dellisolli (Angas, 1864), photo: Salt Rock, atal, SouthAfrica. 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

10rIllllla fllllcbris (Kclaart, 1858). (Figures 4E, F).The white body with black rings, black gill andrhinophores is characteristic of 101'11111111 jilllclJris.Rosfllllga sp. The uniformly red-orange granularbody indicates that this dorid is most likely aspecies of Rostallga.unidentified dorid. This dorid with a granulartexture, unipinnate gill branches and anastanlosingreddish lines cannot readily be placed in anyspecific group of cryptobranch dorids.DClldrodoris dCllisolli (Angas, 1864). (Figures 4C,H). The brownish body with bright blue spots, largegill and rhinophores strongly suggest that Ito'sdrawing represents DClldrodoris dCII isoll i.DClldrodoris Iligra (Stimpson, 1855). (Figures 41, 1).Ito's drawing of an animal with a black notum anda lighter marginal band, sparsely pinnate gill andwhite apices of the rhinophores strongly suggeststhat this is DClldrodoris lIigra.DClldrodoris sp. (Figure 6L). It appears that thismottled dorid with large rhinophores represents aspecies of DClldrodoris. The egg mass (drawn by Itoand shown for this species in Figure 6L) indicatesan egg mass with few, large eggs suggesting thatthis species has direct development.orange dorid. This reddish orange dorid with apustulose notum and no readily evident dorsal gillmay be a species of Phyllidiidae.PlItfllidiella lIigra (van Hasselt, 1824). The blackbody with large rounded lighter tubercles suggeststhat this species is likely Phyllidiella lIigra.PlIyl/idiel/a pIIstlllosa (Cuvier, 1804). The lowirregular, pinkish tubercles of this black animalsuggest that it is likely PlIyllidiella pIIstlllosa.ClIro1110doris allllae Bergh, 1877. (Figures SA, B).The distinctive pattern of a blue body withoutlongitudinal black bands with orange margin ischaracteristic of Chr01110doris a111 liIe.ClIrolllOdoris 111agllifiCl7 (Quoy and Caimard, 1832).(Figures 5C, D). The white body with blacklongitundinal lines, orange margin, orange gill andrhinophores is distinctive for Chrolllodoris lIIagllifiCl7.Chrolllodoris striatella Bergh, 1876. (Figures SE, F).A white body with a series of brown lines orvarying intensity is a distinguishing feature ofClIrolilOdoris striatclla, as are the orange marginalspots and rhinophores with opaque white spots.ClIrolllodoris allrcopllrpllrca Collingwood, 1881.This species with a white body and brownish notalspots and a blue margin is similar to that ofChrolllodoris allreopllllctata and C. rllfo111aclllataPease, 1871. The egg mass is also illustrated.Glossodoris cillcta (Bergh, 1888). (Figures. 5C, H).The brownish body with blue and orange marginalbands and deeply indented central portions of themantle are characteristic of Glossodoris cillcta.llypsclodoris blll/ock} (Collingwood, 1881).(Figures. 51, J). Ito's drawing of Hypselodorisbllllocki is very characteristic with the elevated gill-

91

base, orange rhinophore and gilL opaque whitemarginal band and reddish purple body color.Ccmtosolilll gmcillillllllll Semper in Bergh, 1876.(Figures 5K, L). The relatively uniform reddishbody color with numerous spots of mottling andcolor indicate that Ito's illustration of this species ismost likely Ceratosollla gracillilillllil.Cemtosollla tClllle Abraham, 1876. The reddishmottled body with relatively little expansion of themantle and a purple marginal line suggest that thisspecies is most likely Ceratosollla tell Ill'.

Cemtos0111a trilolJtltulII (Cray, 1827). The reddishbody, wide mantle lobes suggest that this species ismost likely Ceratosolilil trilobatlllil.Aegires sercllae (Cosliner and Behrens, 1997).(Figures 6A, B). The distinctive greenish color, largebranchial lobes and smooth rhinophores arecharacteristic for Aegires sercllac.NC111brotlIa clIalllberlailli Cosliner and Behrens,1997. (Figures 6C. D). The white body color withbrownish pigment on the center of the notum and ared gill are characteristic for the common Philippinespecies, NClllbrotha clllllllbcrlailli.?Roboastra IlItcolillcata (Ba ba, 1936). Thisillustration of Ito's suggests that the black animalwith numerous longitudinal lines of yellow may beRoboastra IlIteolillmta. The large oral tentacles thatare characteristic of this species are not evident inIto's drawing, suggesting it may also represent aspecies of Talllbja or that this characteristic wasomitted from the drawing.Scyllaea pclagica Linnaeus, 1758. (Figures 6E, F).This species is quite distinctive with its four largemantle lobes and expanded rhinophore sheaths.There is little doubt that this is SCljllaca pelagica.Tritollia sp. (Figure 6K). This white species withdarker pigment on the head, appears to be a speciesof Tritollia, but cannot be identified with anyknown species.A 1'111 ilia sp. 1. (Figures 6(;, H). This species with ared body and numerous opaque white longitudinallines appears to be undescribed. The specimendepicted by Ito closely matches a specimencollected from Anilao, Batangas, Philippines(Figures 6H).Arlllilla sp. 2. Ito illustrated a second species ofArlllilla with uniform orange pigment and lowlongitudinal ridges. It cannot be readily identifiedwith any known species.OCrJllatolmmclllls sp. 1. Ito illustrated a dark grayspecies of Dcrlllatobmllclllls with lighterlongitudinal lines. This species cannot be readilyidentified with any known species.Oerlllatobrallchlls sp. 2. A second species ofDCl'llliItolmmchlls was illustrated bv Ito. It is alsogray in color with lighter longitudinal lines. Itdiffers from the previous species in having moreprominent rhinophores that are ornamented withyellow 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. Nembrothachamberlaini 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

along the margin of the foot. This species cannot bereadily identified with any known species.Pteraeolidia ianthina (Angas, 1864). (Figures 61, J).The elongate body shape, elevated ceratal clustersand elongate oral tentacles make Ito's illustrationimmediately identifiable with Pteraeolidia iantlzina.

While tentative identifications are made here forall of the material illustrated by Ito during theAlbatross Expedition, only 35 of Ito's 53 illustrationsare included in the plates presented here. All of theimages, including those not included in this paperwill be posted on the Nudibranch PEET webpage:http:// www.calacademy.org/research/izg/nudibranchsl .

ACKNOWLEDG EMENTS

I wish to thank Diane Tyler for first bringing theexistence of Ito's paintings of mollusks to myattention and for her enthusiastic interest in thehistorical importance of these surviving documentsof the Albatross expedition. I also wish to thankmany individuals who have contributed to the

93

knowledge of Philippine opisthobranchs andassisted in our various field expeditions. I also wishto thank Bob Bolland who has advanced ourknowledge of the opisthobranch fauna from thenearby Ryukuyu Islands of Japan. This work wassupported by a National Science Foundation PEETgrant DEB 0329054 Phylogenetic systematics ofdorid nudibranchs to the author and Angel Valdes.

REFERENCES

Elwood, H. A. Valdes and T. Gosliner 2000. Two newspecies of Aldisa Bergh, 1878 (Mollusca,Nudibranchia) from the tropical lndo-Pacific.Proceedings of the California Academy of Sciences 52: 171­181.

Gosliner, T. and D. Tyler. 1990. Marine Gastropodscollected by the Steamer Albatross in 1908. WesternSociety of Malacologists Annual Report 22: 15.

Springer, V. 1999. Kumataro Ito, Japanese Artist on Boardthe u.s. Bureau of Fisheries Steamer Albatross Duringthe Philippine Expedition, 1907-1910. Marine FisheriesReview 61: 42-57.