From the Marshall Islands, Including 57 New Records 1

Home , Aegires, Aldisa, Aplustridae, Berghia, Berthellina, Berthellina citrina

Notes on Some Opisthobranchia (Mollusca: Gastropoda) from the Marshall Islands, Including 57 New Records 1

SCOTT JOHNSON2 and LISA M. BOUCHER2

ABSTRACT: The rich opisthobranch fauna of the Marshall Islands has re mained largely unstudied because of the geographic remoteness of these Pacific islands. We report on a long-term collection ofOpisthobranchia assembled from the atolls of Bikini, Enewetak, Kwajalein, Rongelap, and Ujelang . Fifty-seven new records for the Marshall Islands are recorded, raising to 103 the number of species reported from these islands. Aspects ofthe morphology, ecology, devel opment, and systematics of 76 of these species are discussed.

THE OPISTHOBRANCH FAUNA OF THE Marshall viously named species are discussed, 57 of Islands, a group of 29 atolls and five single which are new records for the Marshall islands situated 3500 to 4400 km west south Islands (Table 1). west of Honolulu, Hawaii, is rich and varied but has not been reported on in any detail. Pre vious records of Marshall Islands' Opistho METHODS branchia record only 36 species and are largely restricted to three studies. Opisthobranchs The present collections were made on inter collected in the northern Marshalls during the tidal reefs and in shallow water by snorkeling period of nuclear testing (1946 to 1958) and and by scuba diving to depths of 25 m, both now in the U.S. National Museum, along with by day and night. additional material from Micronesia, were Descriptions, measurements, and color studied by Marcus (1965). Marcus and Burch photographs of living specimens were re (1965) and Young (1967) reported additional corded. When possible, prey items were col species from Enewetak Atoll based on short lected and identified. Ova were obtained by term collections at the Enewetak Marine Bio holding specimens in individual containers logical Laboratory (presently the Mid Pacific placed in an aquarium provided with flow Research Laboratory). through seawater at ambient temperatures This study is the result of a long-term (27.5° to 29°C). We have followed conven collection assembled in the Marshall Islands tion in using Thompson's (1961) larval shell over a period of 3 years. Collections were classification. Developmental features of the made virtually daily at sites throughout chromodorid species recorded here are found Enewetak Atoll (11033 /N, 162°20/E) for a in Boucher (in press). period ofmore than a year. Shorter term col Initial identifications based on external lections were made at Rongelap (11°19/ N, morphological characteristics were verified 166°49/E), Bikini (11035/N, 165°20/E), by dissections of the radulae and reproduc Kwajalein (8°58/ N, 167°42/ E) and Ujelang tive systems when appropriate. Descriptions (9°51/N, 160058/E). Additional records were of coloration, external morphology, and in obtained from collections and from photo ternal anatomy are provided only for those graphs taken by several residents ofKwajalein. species not adequately described in previous In this paper 76 readily identifiable, pre- studies, or in those cases in which our material varied significantly from other descriptions. Locality data are cited such that the atolls are simply named followed by specific local 1 Manuscript accepted 3 January 1983. 2 Mid Pacific Research Laboratory, Enewetak, Mar ities within the atoll , for example, Jedrol shall Islands, P. O. Box 1768, APO San Francisco 96555. Island, Medren Pier, Enewetak. 251 252 PACIFIC SCIENCE, Volume 37, July 1983

TABLE 1

S UMMARY OF OPISTHOBRANCH S PECIES R EPORTED FROM THE M ARSHALL ISLANDS

OPISTHOBRANCH SPECIES ATOLL SOURCE

Order Soleolifera

Onchidiella evelinae Mar cus and Burch, 1965 E Marcus and Bur ch 1965 Peronia peronii (Cuvier, 1804) A, B, E Mar cus 1965

Order Cepha laspidea

Acteon variegatus (Bruguiere, 1789) E Ekdale et al. 1979 Pupa sulcata (Gmelin, 1791) E Ekdale et al. 1979 Retusa sp. B M arcus 1965 Phanerophthalmus smaragdinus (Riippell and Leuckart, 1828) E Marcus and Burch 1965 Smaragdinella calyculata (Broderip and Sowerby, 1829) E Mar cus and Burch 1965 Hydatina amplustre (Linnaeus, 1758) E, K Brost and Coale 1971 (K) Hydatina physis (Linnaeus, 1758) K Brost and Co ale 1971 E *M icrom elo guamensis (Quoy and Gairnard , 1825) E, K Atys cylindricus (Hebling, 1779) E Ekdale et al. 1979 Haminoea cymbalum (Quoy and Gairnard, 1835) E Marcus and Burch 1965 Hamino ea galba Pease, 1861 E Ekda1e et al. 1979 Ham inoea linda Marcus and Burch, 1965 E Marcus and Burch 1965 Hamin oea musetta Marcus and Burch, 1965 E Marcus and Bur ch 1965 *Bulla vernicosa Gould, 1859 E Chelidonura hirudin ina (Quoy and Gairnard, 1833) B Marcus 1965 K " Chelldonura inornata Baba, 1949 E, K Philinopsis gardineri (Eliot, 1903) E Switz er 1971 (?) K *Sagaminopteron psyc hedelicum Carlson and Hoff, 1974 K

Order An aspidea

*Aplysia parvula Guilding in March, 1863 E, K Aplysia pulm onica Gould, 1852 E Marcus 1965 *Dolabella auricularia (Lightfoot, 1786) E,K Dolabrifera dolabrifera (Cuvier, 1817) B Mar cus 1965 E Marcus and Bur ch 1965 K Stylocheilus longicaudus (Quoy and Gaim ard, 1825) B Ma rcus 1965 E M arcu s and Burch 1965 K

Order Sacoglossa

*Stiliger smaragdinus Baba, 1949 E *Plak obranchus ocellatus van Hasselt , 1824 B, E,K, U Elysia bayeri Marcus, 1965 B Marcus 1965 E, K *Elysia halimedae MacNae, 1954 E *Elysia livida Baba, 1955 E Elysia marginat a (Pea se, 1871) B Marcus 1965 E, R *Elysia obtusa Baba, 1938 B,E *Elysia ratna M arcus, 1965 E,K *Elysia vatae Ri sbec , 1928 E, K * Oxynoe viridis (Pease , 1861) R Marshall Islands Opisthobranchia-s-Jonxsox AND BOUCHER 253

OPISTHOBRANCH SPECIES ATOLL SOURCE

Order Notaspidea

*Berthellina citrina (Ruppell and Leuckart , 1828) E, K

Order Nudibranchia

Hexabran chus sanguineus (Riippell and Leuckart , 1828) E Marcus 1965 K Doriopsis pecten (Collingwood, 1881) E Young 1967 B,K, U Doriopsis viridis Pease, 1861 E Young 1967 *P/atydoris cruenta (Quoy and Gaimard, 1832) E,K,U P/atydoris scabra (Cuvier, 1804) B Marcus 1965 E,K Jorunna alisonae Marcus, 1976 E Young 1967 *Jorunna Junebris (Kelaart , 1858) E,K *Discodoris fra gi/is (Alder and Hancock, 1864) E *Trippa intecta (Kelaart, 1858) R "Halgerda e/egans Bergh" Marcus , 1965 R Marcus 1965 *Ha/gerda e/egans Bergh, 1905 K *Ha/g erda wasinensis Eliot, 1904 E *Selerodoris paliensis Bertsch and John son, 1982 E *A/disa pikokai Bertsch and Johnson, 1982 K Casella atromarginata (Cuvier, 1804) B Marcus 1965 E,K *Chromodoris a/bopunctatus (Garrett, 1879) E Chromodoris a/bopustu/osa (Pease, 1860) E Young 1967 K Chromodoris briqua Marcus and Burch, 1965 (?) E Marcus and Burch 1965 *Chromodoris decora (Pease, 1860) B,E,K *Chromodoris elisabethina Bergh, 1877 E,K Chromodoris fidelis (Ke1aart, 1858) E Marcus and Burch 1965 B,K Chromodoris geometrica Risbec, 1928 E Young 1967 K Chromodoris inornata Pease, 1871 E Young 1967 B,K *Chromodoris marginata (Pease, 1860) B,E *Chromodoris tryoni (Garrett, 1873) E,K *Hypse/odoris australis (Risbec , 1928) E,K *Hypse/odoris decorat a (Risbec , 1928) E,K *Hypse/odoris infu cata (Ruppel! and Leuckart, 1828) E Hypse/odoris ka yae Young , 1967 E Youn g 1967 Hypse/odoris mouaci (Risbec, 1930) E Marcus and Burch 1965 K *Thorunna elitonata (Bergh, 1905) E *Thorunna decussata (Risbec , 1928) E,K *Thorunna norba (Marcus and Marcu s, 1970) E *Cadlinella ornatissima (Risbec, 1928) B,E,K *Dendrodoris coronata Kay and Youn g, 1969 E, K, R *Dendrodoris e/ongata Baba, 1936 B,E,K Dendrodoris erubescens (Bergh, 1905) E Marcus and Burch 1965 Dendrodoris nigra (Stimpson, 1855) E Marcus and Burch 1965; Young 1967 B,K,U *Dendrodoris tubercu/osa (Quoy and Gaimard, 1832) K *Gymnodoris cey/onica (Ke1aart , 1858) E 254 PACIFIC SCIENCE, Volume 37, July 1983

TABLEI (Cont.)

OPISTHOBRANC H SPECIES ATOLL SOURCE

Gymnodoris citrina (Bergh, 1877) E Marcus and Burch 1965; Young 1967 B,K *Gymnodoris okinawae Baba, 1936 E,K *Gymnodoris p/ebeia (Bergh, 1877) E "Gymnodoris striata (Eliot, 1908) E *Nem brotha kubaryana Bergh, 1877 K *P/ocamopherus cey/onicus (Kelaart, 1858) E *Aegires villosus Farran, 1905 E *Goniodoris joubin i Risbec, 1928 B,E,K " Goniodoridieila savignyi Pruvot-Fol, 1933 E Okadaia e/egans Baba, 1931 E Young 1967, 1969 Phy/lidia nobilis (Bergh, 1869) R Marcus 1965 *Phyllidia pustu/osa Cuvier, 1804 B, E,K Phy/lidia varicosa Lamarck, 1801 B Marcus 1965 E,K *Dermatobranchus fo rtunata (Bergh, 1888) E, K *Bornella adam sii Gray, 1850 E,K *Ma rianina rosea (Pruvot-Fol, 1930) B,E,K,U Flabellina alisonae Gosliner, 1980 E Gosliner 1980 B,K,U Phestilla /ugubris (Bergh, 1870) E Switzer 1971 K *Favorinusjaponicus Baba , 1949 E, K *Phyllodesmium hyalinum Ehrenber g, 1831 E,K Pteraeolidia ianthina (Angas, 1864) E Switzer 1971 K *Phidiana bourailli (Risbec, 1928) E, K *Phidiana indica (Bergh, 1896) E, K Herviella claror Burn, 1963 E Marcus and Burch 1965; Young 1967 Herviella mietta Marcus and Burch, 1965 E Marcus and Burch 1965; Young 1967 *G/aucus at /anticus Forster, 1777 E, K *Spurilla major (Eliot, 1903) E

NOTE: A = Arno, B = Bikini, E = Enewetak, K = Kwajalein, R = Rongelap, and U = Ujelang. A new record for the Marshall Islands is denoted by • .

Specimens and color slides of most of the MATERIAL: Empty shells are occasionally species discussed have been deposited in the found on Enewetak Atoll lagoon reefs and collections ofthe Bernice P. Bishop Museum, beaches. Honolulu, Hawaii , and the Mid Pacific DISTRIBUTION: Indo-west Pacific (Kay Research Laboratory, Enewetak, Marshall 1979). Islands.

Hydatina physis (Linnaeus, 1758) DESCRIPTION OF SPECIES REFER ENCE : Kay (1979); Bertsch and John ORDER CEPHA LASPID EA son (1981: 16). FAMILY APLUSTRIDAE MATERIAL: One specimen, 20 x 12mm, Enewetak, Bokandretok-Medren reef, in sand, Hydatina amplustre (Linnaeus, 1758) depth 8m, 20 Nov. 1981, P. Colin and L. Bell.

REFERENCE: Kay (1979); Bertsch and John DISTRIBUTION: Indo-west Pacific, parts of son (1981: 16). tropical Atlantic (Kay 1979). Marshall Islands Opisthobranchia-c-Jonxsox AND BOUCHER 255

Micromelo guamensis (Quoy and Gaimard, man 1970), Guam (Carlson and Hoff 1981), 1825) and the Caroline Islands (Marcus 1965). REFERENCE: Kay (1979); Bertsch and John son (1981: 17). Philinopsis gardineri (Eliot, 1903a)

MATERIAL: One specimen, 25 x 10 mm, REFERENCE: Bergh (1905: tafel 3, fig. 6, as Enewetak, lagoonside Jedrol Island, on sand, Chelidonura velutina). 13m, 21 August 1981,A. Thresher. One speci men, photographed at Kwajalein, D. John MATERIAL: One specimen, 30 mm, Enewe son. tak, lagoonside Mut Island, under dead coral, 5 m, 30 November 1978. One specimen, DESCRIPTION: Marshall Islands specimens photographed at Kwajalein, D. Johnson. differ from Hawaiian conspecifics in having a yellowish-white foot and cephalic shield DESCRIPTION: Body broad, rounded ante margins, which fade submarginally into a thin riorly, and notched posteriorly, with lateral line of bright green, then into a wide, almost parapodia held upright. Color jet black with irridescent band of bluish-purple. brilliant blue parapodial margins.

DISTRIBUTION: Pacific, including Hawaii, DISTRIBUTION: East Africa (Eliot 1903a), Guam, Japan, and Australia (Kay 1979). Indonesia (Bergh 1905), and Guam (Carlson and Hoff 1981).

FAMILY BULLIDAE REMARKS: Eliot's (1903a) description was ofa preserved specimen and does not mention the blue margins. Switzer (1971) may have Bulla vernicosa Gould, 1859 been referring to this species when she re . REFERENCE: Kay (1979); Johnson (1982: ported Chelidonura velutina Bergh, 1905, from 89). Enewetak. As Rudman (1973a) has pointed out, Bergh's species is actually a combination MATERIAL: Empty shells are common on of two previously described species, Philinop lagoon beaches at Enewetak. Occasional sis gardineri and Chelidonura varians Eliot, specimens crawl on sand or dead coral at 1903. night.

DISTRIBUTION: Pacific (Kay 1979). FAMILY GASTROPTERIDAE

FAMILY AGLAJIDAE Sagaminopteron psychedelicum Carlson and Hoff,1974

Chelidonura inornata Baba, 1949 REFERENCE: Carlson and Hoff(1974: pI. 10, REFERENCE: Baba (1949); Rudman (19:70); fig. 4). Johnson (1982: 88). MATERIAL: One specimen, photographed at MATERIAL: Six specimens, Enewetak, Me Kwajalein, D. Johnson. dren Pier pilings, 5 m, 6 July and 15 July 1981. DISTRIBUTION: Guam, northern Marianas One specimen, collected and photographed at Islands (Carlson and Hoff 1974). Kwajalein, D. Johnson.

DESCRIPTION: Marshall specimens possess two diffuse, light-blue patches on the anterior ORDER ANASPIDEA edge of the white band on the cephalic shield which are not mentioned in previous reports. FAMILY APLYSIIDAE DISTRIBUTION: Japan (Baba 1949), Austra lia (Burn 1966), the Solomon Islands (Rud- Aplysia parvula Guilding in March, 1863 256 PACIFIC SCIEN CE, Volume 37, July 1983

SYNONYMY: See Eales (1960). several deep in places, crawling across silty sand, was observed off Runit Island at a depth RE FERENCE: Bebbington (1977). of 24 m on 2 August 1981 (Colin and Kirby, MATERIAL: Specimens are common be pers. comm.). neath dead coral on the intertidal reefs of DEVELOPM ENT: Egg masses are frequently Enewetak and Kwajalein. found beneath rocks with groups ofSty lochei DESCRIPTION: Marshall Island s specimens Ius and consist of irregular tangles of bro wn are brownish, becoming yellow-brown toward spaghettilike strands, often very firmly at the foot. Body densely covered with small tached to the substrate. Three to five ova, each white spots and a few irregularly scattered measuring about 60 ,urn ,occur in each capsule white patches. Tail and parapodial margins, of 170,urn in diameter. Free swimming veliger the anterior edge of the foot , and rhinophore larvae begin hatching in 5 days. Larval shells and tentacle tips black . Black patches densely are tran sparent tan in color, with bro wn flecked with white surround the eyes, just columellae. anterior to the rhinophores. Approximately DI STRIB UTIO N: Circumtropical (Engle and one-third of the amber-yellow shell exposed Hummelinck 1936, Bebbington 1974, 1977). through a circular, black-rimmed opening in the mantle. Radula as figured by Bebbington (1977). ORDER SACOGL OSSA DI STRIBUTION: Circumtropical, including Indo-west Pacific (Bebbington 1974, 1977), the tropical western Atlantic (Marcus 1956), FAMILY HERM AEIDAE and England (Bebbington and Brown 1975). Stiliger smaragdinus Baba, 1949

REF ERENCE: Baba (1949: pl. 7, fig. 22).

Dolabella auricularia (Lightfoot, 1786) MATERI AL: One specimen, 22 x 5 mm, REFERENCE: Bebbington (1977); Kay Enewetak, Medren Pier pilings; on algae, 5 m, (1979). 6 July 1981.

MATERI AL: About 50 individuals observed DESCRIPTIO N: Body elongate, densely cov at Enewetak and Kwajalein, usually beneath ered with inflated, ceratalike pallial proce sses. rocks on lagoon reefs and pinn acles, often Base of each process translucent with light with grayish-green, spaghettilike egg masses. blue-green spots; distal three-fourths of each process green with opaque white patches and DISTRIBUTIO N: Circumtropical (Engle 1942, dense black freckling near the tip. Head green, Bebbington 1974, 1977). rhinophores long, smooth, green with white tips. Rhinophoral bases with several lines of white speckles extending posteriorly. Radul a S tylocheilus longicaudus (Quoy and Gaimard , as figured by Baba (1949). 1825) DI STRIB UTIO N: Japan (Baba 1949). REFERENCE: Bebbington (1977); Fielding (1979; 51); Kay (1979). FAMILY PLAKOBR ANCHIDAE MATERIAL: This species is one of the most frequentl y observed ofall the opisthobranchs at Enewetak. The animals are found abun Plakob ranchus ocellatus van Ha sselt, 1824 dantly with the alga Lyngbya, on and under REFERENCE: Kay (1949); Bertsch and John dead coral rocks and on sand y bottoms from son (1981: 20, 21). the intertidal to depths of at least 30 m. They often occur in large groups. A dense aggrega MATERIAL: Common in sand and rubble tion of man y thousands of individuals, piled areas on shallow lagoon reefs at Enewetak, Marshall Islands Opisthobranchia-i-Joanson AND BOUCHER 257

Bikini, Kwajalein, and Ujelang. Specimens touching only intermittently middorsally. frequently observed crawling exposed on the Color dark gray-black, the parapodial substrate. margins vivid orange, submarginally with a black band and a line of brilliant turquoise. DISTRIBUTION: Indo-west Pacific (Kay Base of each rhinophore with a patch of the 1979). same color sequence. Coloration within the parapodia as on the outer surface . Radula as Elysia bayeri Marcus, 1965 figured by Baba (1955) and Carlson and Hoff (1978). REFERENCE: Marcus (1965); Carlson and Hoff (1978). DISTRIBUTION: Japan (Baba 1949), Guam (Carlson and Hoff 1978). MATERIAL: More than 100 specimens ob served at Enewetak and Kwajalein, under and REMARKS: There may be some confusion on dead coral on lagoon reefs and pinnacles , as to the identity of this species. Our animals at depths of 3-10 m. differ in color from those originally described by Baba (1955) (that is, of bluish-black ani DEVELOPMENT: An egg mass was deposited mals with black, orange-red, and black bands by a 16mm individual on 16 December 198I. on the parapodial margins) but have a similar Ova orange, about 80/lm individually within radula. Abe's (1964) redescription of the ani capsules 120- I40 /lm in diameter. Ova depos mals ofE.livida is, however, identical with our ited in close-set whorls, coiled singly around specimens, as is the color description by Carl an orange extra capsular yolk string; about 29 son and Hoff (1978) of animals from Guam. ova per mm and 2500 in the entire mass. There is an eIysiid which occurs at Enewetak DISTRIBUTION: Bikini (Marcus 1965), and at Guam (Carlson and Hoff 1978) with Guam, and Palau (Carlson and Hoff 1978). coloration similar to that of Baba's (1955) original description of E. livida but which has a different radula. Elysia halimedae MacNae, 1954

REFERENCE: MacNae (1954); Carlson and Hoff (1978). Elysia marginata Pease, 1871

MATERIAL: One specimen, 12 x 2 mm, REFERENCE: Carlson and Hoff (1978). Enewetak, lagoonside Enewetak Island , on MATERIAL: Four specimens, the largest Halimeda, 3 m, I I October 1981. 45 mm in length, found at Rongelap and Ene DISTRIBUTION: Indo-west Pacific, including wetak, on or under dead coral at depths of South Africa (MacNae 1954), Japan (Baba 2-3m. 1957), Australia (Burn 1972), Guam, Ponape (Carlson and Hoff 1978), and Hawaii (Kay DISTRIBUTION: Indo-west Pacific (Carlson 1979). and Hoff 1978).

Elysia livida Baba, 1955 Elysia obtusa Baba, 1938 REFERENCE: Carlson and Hoff (1978); Abe Figure I (1964: pI. I I, fig. 39). REFERENCE: Carlson and Hoff(1978); Baba MATERIAL: Two specimens, 15 x 2 mm, (1949: pI. 9, figs. 28, 29). 14 x 2 mm, Enewetak, lagoonside Enewetak Island, on dead coral, 2-5m, 26 June 1981 MATERIAL: Three specimens, the largest 12 x 2.5 mm, found on lagoon reefs and pin and 26 August 1982. nacles of Enewetak and Bikini, at depths of DESCRIPTION: Body elongate, smooth. 2- 15m. One specimen observed crawling in Parapodia upright in slightly wavy folds, the open at night. 258 PACIFIC SCIENCE, Volume 37, July 1983

MATERIAL: More than 50 specimens, mea suring up to 16 x 2.5mm, recorded at Enewe tak and Kwaja1ein, usually on or under algal covered dead coral in I-10m of water, on both lagoon reefs and pinnacles.

DESCRIPTION: Body smooth; parapodia meet middorsally in a straight line extending to the tail. Aspects of the coloration vary betweenindividuals; usually light to dark gray, 40 with numerous black spots and fewer yellow spots; parapodia1 margins with a narrow to F IGURE I. Elysia obtusa Baba . Single radular tooth wide band of light yellow; rhinophores white, (scale bar in Jim). upper third orange-red. Radu1ar teeth as fig ured by Carlson and Hoff (1978).

DEVELOPMENT: An 11 mm specimen depos ited an egg mass 7 December 1981 and two DESCRIPTION: Radula of a 12mm specimen more on 12 December 1981. Ova bright with seven teeth in the ascending series and orange, arranged in six close-set whorls coiled seven in the descending. The teeth (Figure 1) flat on the substratum. Ova measure 55 differ from those figured by Baba (1949) in 65 pm individually within capsules 120pm in being slightly more humped dorsall y and lack diameter. Capsules piled in two layers around the minute serrations described by Carlson an orange extra capsular yolk string. Larvae and Hoff (1978). begin hatching in 5 days as free-swimming ve1igers with tran sparent shells. DISTRIBUTION: Japan (Baba 1938, 1949), Guam (Carlson and Hoff 1978). DI STR IBUTIO N: New Caledonia (Risbec 1928), Indonesia (Bergh 1905, as Elysia sp.), Guam, the northern Marianas, and Palau Elysia ratna Marcus, 1965 (Carlson and Hoff 1978). REFERE NCE: Marcus (1965); Carlson and REMARKS: Risbec (1928) figured this species Hoff (1978). as Elysia vatae and described it as Elysia MATERIAL: More than 100specimens, rang vataae. Since the figure precedes the descrip ing to 28 x 3 mm, observed on Enewetak tion, we follow Carlson and Hoff (1978) in and Kwaja1ein lagoon reefs and pinnacles, at using the former spelling. depths of2-20 m.

DEVELOPM ENT: Egg masses deposited by 14 FAMILY OXYNOEIDAE and 15mm individuals on 20 November 1981. Ova orange, measuring 80- 85pm individually within relatively large capsules 190pm in dia Oxynoe viridis (Pease, 1861 ) meter. Ova spiralled singly around an orange REFERENCE.: Hamatani (1980); Baba (1955: extra capsular yolk string. About 20 ova per pl. 2, figs. 3, 4, 5). mm of coil, and 2250 in the entire mass. MATERI AL: One specimen, 21 x 6 mm, DISTRIBUTION: Palau (Marcus 1965), Guam Rongelap, 1agoonside Maen Island, under (Carlson and Hoff 1978). dead coral, 1m, 12 September 1982.

DI STRIB UTIO N: Indo-west Pacific, including Elysia vatae Risbec, 1928 Tahiti (Pease 1861), Australia (Allan 1950, as RE FERENCE: Carlson and Hoff (1978); Ris O. olivacea Rafinesque), Japan (Baba 1952), bee (1928: pl. 12, fig. 7). and Zanzibar (Eliot 1906a). Marshall Islands Opisthobranchia-i-Joaxson AND BOUCHER 259

ORDER NOTASPIDEA FAMILY DORIDIDAE

FAMILY PLEUROBRANCHIDAE Platydoris cruenta (Quoy and Gaimard, 1832)

Berthellina citrina (Riippell and Leuckart, Figure 2 1828) REFERENCE: Risbec (1928); Bergh (1905: tafel 1, fig. 3). SYNONYMY: See Burn (1962); Thompson (1970). MATERIAL: One specimen, 70 x 46 mm (damaged), Enewetak, Mooring Buoy pin REFERENCE: Thompson (1970); Bertsch and Johnson (1981: 26, 27). nacle, under dead coral, 4 m, 8 August 1981.One specimen, 51 x 32mm, Kwajalein , MATERIAL: Relatively common at Enewe Gagan Quarry, under dead coral, 2 m, 15June tak and Kwaja1ein, under dead coral on 1982, L. Rousseau. One specimen, 95 x lagoon reefs and pinnacles at depths of 1 61mm, Kwajalein, Kwajalein intertidal reef, 10m. Individuals frequently observed preymg under dead coral, 21 June 1982, R. Alderson. on sponges and scleractinean corals in ledges One specimen, Ujelang, lagoonside Jereko and caves at night. Island, under dead coral, 1m, 28 July 1982. DESCRIPTION: Jaw elements of our speci One specimen, 100 x 53 mm, Kwajalein, mens differ slightly from those reported by Kwajalein intertidal reef, under dead coral, Thompson (1970) in having more regularly 14 September 1982, L. Bell. arranged and rounded, rather than pointed, denticulations. DESCRIPTION: Mantle rigid, granular, flat tened, and broadly oval with wide, flaring DISTRIBUTION: Indo-west Pacific, the Medi margins that cover the narrow foot. Colora terranean, the Atlantic coast of France, and tion varies among individuals: typical speci the southern part of Great Britain (Thompson mens cream, densely streaked with short, 1970). curved, black lines and splotched with large orange or red spots; others with black streaks missing, and replaced by patches of fine, ORDER NUDIBRANCHIA brown, sievelike reticulations. In the largest specimen from Kwajalein the reticulations FAMILY HEXABRANCHIDAE wereeasilyseen beneath the curved black lines. Elevated rhinophore sheaths white, edges Hexabranchus sanguineus (Riippell and scalloped; rhinophores white, each with about Leuckart, 1828) 40 lamellae varying from orange-red to gray brown. Branchial pocket convoluted with six REFERENCE: Thompson (1972a); Bertsch and Johnson (1981: cover). folds through which the six arborescent stalks protrude. Gillswithwhitebases,grayish-brown, MATERIAL: One specimen, 180mm, Enewe white-tipped branches. tak, washed ashore on the oceanside of Ene Radu1ar formulae of two specimens, both wetak Island, 30 December 1981 , R. Dubin. exceeding 80 mm in length, is 39 x 99.0.99 One specimen, 45mm, Kwajalein, oceanside and 42 x 170.0.170. Teeth long, curved, Ennubuj Island , under dead coral, 10m, 3 pointed rods with slightly thickened bases March 1982. (Figure 2). DISTRIBUTION: Indo-west Pacific (Thomp DISTRIBUTION: New Guinea (Quoy and son 1972a). Gaimard 1832), Indonesia (Bergh 1905), REMARKS: Hexabranchus sanguineus was Japan (Eliot 1913, Baba 1936, as Argus reported from Enewetak by Marcus (1965) as cruentus), New Caledonia (Risbec 1928), and H. marginatus (Quoy and Gaimard 1832). Vietnam (Risbec 1956). 260 PACIFIC SCIENCE, Volume 37, July 1983

250

F IGURE 2. Platydoris cruenta (Quoy and Gaimard). Right-half row of radular teeth (scale bar in 11m).

REMARKS: Our material indicates that Pla measuring about 80 llm individually within tydoris cruenta is variable both in coloration capsule s 10011m in diameter. Approximately and in radular formul a. Risbec (1928) and 210 ova per mm? and 152,250 in the entire Eliot (1910) havealso described color variation mass. Larvae hatched in 5 days as free in Platydoris spp. Considering this variation, swimming veligers. The type-one larval shells the validity of species similar to P. cruenta, were transparen t with purple- stained colu such as P. striata (Kelaart, 1858), P. sanguinea mellae. Bergh, 1905, P. flammulata Bergh, 1905, and DISTRIBUTION: Indo-west Pacific (Edmunds perhaps even P.formosa (Alder and Hancock, 1971). 1864) is suspect. REMARKS: We have observed Platydoris scabra in Hawaii, where it is rare around the Platydoris scabra (Cuvier, 1804) main islands but common within the lagoon REFERENCE: Edmunds (1971). of Kure. MAT ERIAL: Six specimens, the largest 34 x 20 mm, observed beneath dead coral and in Jorunna funebris (Kelaart, 1858a) caves at night on lagoon pinnacles of Kwaja Figure 3 lein and Enewetak, at depths of 10-15 m. REFERENCE: Marcus (1976). DESCRIPTION: Animals yellowish, densely speckled with tiny brown flecks, aggregated MATERIAL: About 20 specimens, ranging in irregularly as brown blotches. Scattered size to 30 x 12 mm, found at Enewetak , about the dorsum, but more or less in a longi mostly under dead coral on lagoon reefs and tudinal row on each side of the middorsal line, pinnacles. Seven specimens were feeding on a are small patches of opaque white. External, blue sponge , Haliclona sp. Two additional radular, and reproductive system morphology specimens, Kwajalein, J. and J. Wedge. as reported by Edmunds (1971). DESCRI PTIO N: Body broadly elongate oval, DEVELOPM ENT: A 30 mm individual depos soft, and finely papillose with tiny, pointed ited an egg mass in four whorls, loosely ere caryoph yllidia. Mantle margin wide, draping nulate on the free edge. Ova bright orange, about the foot. Background color white. Marshall Islands Opisthobranchia-i-Jonxson AND BOUCHER 261

specimens are definitely hooked, this dif ference seems insufficient to consider the two species separate and we retain the specific name funebris.

Discodorisfragilis (Alder and Hancock, 1864)

9 14 REFERENCE: Kay and Young (1969); Edmunds (1971); Bertsch and Johnson (1981: 40).

MATERIAL : More than 40 individuals , rang 150 ing to 85 x 50 mm, found at Enewetak, all FIGURE 3. Jorunna f unebris (Ke laart). R ight -half row beneath dead coral in 2- 4 m of water on the of radular teeth (sca le bar in JIm). lagoonside of Enewetak Island . DEVELOPMENT: An egg mass deposited by an 85 mm specimen on 28 August 1981 was Caryophyllidia appearing brown to black, 70 mm in diameter, and crenulated on the free but minutely, densely flecked with chocolate edge. Ova cream-white, measuring 80-90 f1m brown; some ofthe pigmentation spilling onto in diameter, individually encased in capsules adjacent dorsal surface. Pigmented patches 120zmi in diameter. Approximately 86 ova smaller around the margins. Foot white with per mm? and 3,160,500 in the mass. Free occasional brown spots.Rhinophores white swimming veligers begin to hatch in 4 days. with about 18 dark brown to black lamellae. The type-one larval shells were tran sparent Branchiae of 4-6 long, tripinnate stalks, white, tan with dark brown apertures. edged with dark brown or black. Radular formula of a 30 mm individual DISTRIBUTION: Indo-west Pacific (Edmunds 10 x 24.0.24. Teeth shaped like massive, 1971 ). smooth, cylindrical question marks (Figure 3). Tooth shape is similar to that ofa specimen Trippa intecta (Kelaart, 1858a) described by Marcus (1976), but app eared to be somewhat smoother and more regular. REFERENCE: Edmunds (1971); Baba (1949; pI. 24, fig. 89). DEVELOPMENT: An egg mass deposited by a 27 mm individual on 2 December 1981 con MATERIAL: Four specimens,' largest 40 x sisted of three loose, white whorls , attached 22 mm, Rongelap, lagoon side Rongelap by one edge to the substrate. Ova measure Island, beneath coral rubble, 2 m, 13 October approximately 170f1m within individual cap 1981. sules 240 f1m in diameter. About 75 ova per DISTRIBUTION: Indo-Pacifi c (Edmunds mm? and 21,900 in the mass. Larvae hatch in 5 days as free-swimming veligers with type 1971 ), including Hawaii. one shells. Larval shells pure white with purple stained columellae. Halgerda elegans Bergh, 1905 DISTRIB UTION: Indo- west Pacific (Marcus Figure 4 1976). REFERENCE: Bergh (1905: tafel 2, fig. 4a). REMARK S: The systematics of the known MATERIAL: One specimen, 12 x 3 mm, species as Jorunna were reviewed by Marcus Kwajalein, Ninni Island, under dead coral, (1976), who pointed out that the only reliable 8 m, 22 June 1981, K. DeGroff. difference between J. funebris and 1. gigas (Bergh, 1876) is the hook-shaped outermost DESCRIPTION: Body rigid, gelatinous, reti teeth in the latter. Although these teeth in our culated with low ridges that intersect at 262 PACIFIC SCIENCE, Volume 37, July 1983

F IGURE 4. Halgerda elegans Bergh. Right-half row of radular teeth (scale bar in jlm ).

pointed tubercles. Background color bright DEVELOPMENT: A 15mm specimen depos lemon-yellow, with numerous, irregular, criss ited an egg mass on 30 December 1981in three crossing white lines, several of which usually loose whorls att ached by one edge to the sub intersect at the tubercles. Margin wide, rather strate. Ova orange-yellow, 190,um individually thin, translucent white with numerous black within capsules 240 ,urn in diameter. Approxi spots and perpendicular streaks. Sides of foot mately 50 ova per mm? and 4000 in the entire and underside of the mantle whitish, spotted mass examined. Free-swimming veligers begin with black. Rhinophores with white peduncles, to hatch in 5 days. The type-one larval shells black clubs with white tips, and about 15 were tran sparent tan with brown columellae. lamellae each. Branchiae white, splashed with DISTRIBUTION: East Africa (Eliot 1904a, black, of six bipinnate stalks. Rudman 1978). Radular formula of the single specimen 29 x 28.0.28. Teeth mostly hamate (Figure 4), the outermost few straight and fringed with Sclerodoris paliensis Bertsch and Johnson, long, thin denticles. 1982

DISTRIBUTION: Indonesia (Bergh 1905). REFERENCE: Bertsch and Johnson (1981 : REMARKS: A nudibranch reported by Mar 45, as Sclerodoris sp.; 1982). cus (1965)as Halg erda eiegans from Rongelap MATERIAL: Three specimens, the largest 15 was "deep purple with white ring-like mark x 4 mm, found on lagoon pinnacles at Ene ings." This description bears no resemblance wetak, under dead coral or exposed on the to our specimen, and we question Marcus' substrate at night, at depths of 10-12 m. identification. DISTRIBUTION: Hawaii (Bertsch and John son 1982). Halgerda wasinensis Eliot, 1904a

REF EREN CE: Rudman (1978); Eliot (1904a: pI. 34, fig. 1). Aldisa pikokai Bertsch and Johnson, 1982 REFERENCE: Bertsch and Johnson (1981: MATERIAL: Four specimens, from 10 x 44, as Aldisa sp.; 1982). 5 mm to 17 x 7 mm, collected at Enewetak, all under dead coral on lagoon pinnacles at a MATERI AL: One specimen, photographed at depth of 10m. Kwaja1ein, D. Johnson. Marshall Islands Opisthobranchia-s-Joaxsox AND BOUCHER 263

1 5 16 23 27

B 50

FIGURE5. Casella atromarginata (Cuvier). A. Right-halfrow ofradular teeth; B. Anterior view ofinnermost lateral tooth (scale bar in Ilm) .

DISTRIBUTION: Hawaii (Bertsch and John main cusp (Figure 5A, B). Outer teeth hamate, son 1982). each with up to four denticles on the outer surface. Outermost laterals small, simply hamate, smooth. FAMILY CHROMODORIDIDAE DISTRIBUTION: Indo-west Pacific, including Casella atromarginata (Cuvier, 1804) Tahiti (Marcus and Marcus 1970), eastern Australia (Thompson I972a), east Africa and Figure 5A, B the Red Sea (Eliot 1904b, Gohar and Aboul REFERENCE: Thompson (1972b; 1976: pI. 5, Ela 1959), Japan (Baba 1949), Guam (Carlson fig. C). and Hoff 1981), and the small northwestern Hawaiian Islands of Nihoa, French Frigate MATERIAL: Three specimens, the largest 25 Shoals, Mara, and Laysan (pers. obs.). x 6 mm, collected on and under dead coral on Enewetak lagoon pinnacles. Additional specimens from Kwajalein, D. Johnson. Chromodoris albopunctatus (Garrett, 1879)

DESCRIPTION: Radular formula of a 25 mm SYNONYMY: Chromodoris imperialis Kay specimen 117 x 27.0.27. Innermost lateral and Young, 1969; Bertsch and Johnson, 1979a with 3-4 denticles on either side of a long, (non-Pease, 1860); C. sykesi Eliot, 1904. 264 PACIFIC SCIENCE, Volume 37, July 1983

REFERE NCE : Kay and Youn g (1969); albopunctatus were reported as C. imperialis Bertsch and Johnson (1981: 58). (Kay and Young 1969), a species that differs in coloration and external morphology, and MATERIAL: Approximately 20 specimens, from 8 x 5 mm to 75 x 33mm, found along may be synonymous with C. godeffroyana shallow lagoon reefs and pinnacles at Ene (Bergh, 1879). Marcus and Burch's (1965) wetak. Most were beneath dead coral, feeding description from Enewetak of C. briqua re on an unidentified orange-yellow encrusting sembles small specimens of C. albopunctatus sponge. in coloration, and with the exception of the outermost teeth, the radulae of the two species DESCRIPTION: Our specimens exhibit more are similar. variation in color than those described by Garrett (1879), Eliot (1904b), and Kay and Young (1969). Large specimens usually red dish purple with white rings. Individuals Chromodoris decora (Pease, 1860) approximately 30mm and smaller, orange to REFERENCE: Kay and Young (1969); red, with yellow, irregular rings (perhaps bet Bertsch and Johnson (1981: 55). ter described as yellow, densely spotted with MATERI AL: More than 75 specimens found orange or red). Margin tricolored with a wide at Enewetak, Kwajalein, and Bikini, mostly band of bluish-purple, becoming a thin , irreg beneath dead coral on lagoon reefs at depths ular darker purple-black line submarginally, ofl-5m. and a wide band of bright lemon-yellow. Bluish-purple marginal coloration also visible DESCRIPTION: Marshall Islands specimens beneath the mantle , but fades into maroon vary in two aspects of coloration: the purple submarginally. Underside , including the foot , spots on the dorsal white lines and marginal bright lemon-yellow. Lower half of each rhi orange band are not always present, and the nophore orange to light purple with red and orange margin is frequently studded with white spots, becoming orange , red brown, or . white granules. Radular teeth similar to those red-purple with white spots near the tip. figured by Kay and Young (1969), but differ in Branchiae colored as the rhinophores. Ani two minor respects: the mid-half row teeth mals in 70% ethanol become uniformly red have larger bases, and the innermost laterals brown with a yellowish-orange pigment possess two equal-sized inner denticles. leaching into the preservative. Radular tooth shape essentially as figured DEVELOPM ENT: A loosely coiled egg mass by Kay and Young (1969), although there are was deposited on 28 August 1981 by a 13mm individual. Ova cream-yellow, measuring 60 2 inner denticles on the innermost laterals - 80 pm individually within capsules 95 pm in in our specimens, and occasionally, a faint, diameter. A bright orange extra capsular yolk triangular central tooth. body, 60,um in diameter, situated like a cap DEVELOPM ENT: Egg masses deposited by external to the surface of each capsule. About these dorid s are unusual in having discrete 440 ova per mm? and 11 ,360 in the mass. extracapsular yolk bodies of irregular size and Veliger larvae free swimming, with type-one shape scattered throughout. For a discussion shells. of the ova and development, see Boucher (in DISTRIBUTIO N: Hawaii (Pease 1860; Kay press). and Young 1969), Guam (Carlson and Hoff DISTRIBUTION: Tahiti (Garrett 1979),East 1981), eastern Australia (Allan 1947), and Africa (Eliot 1904b), and Hawaii (Kay and Japan (Baba 1938, as Glossodoris setoensis, Young 1969). 1953, 1955).

REMARKS: Eliot's (1904b) Chromodoris REMARKS: Kay and Young (1969) note that sykesi agrees in all particulars with C. Doris lentiginosa Pease, 1871 from Tahiti may albopunctatus. The Hawaiian specimens of C. be a synonym. Marshall Islands Opisthobranchia-s-Joaxsox AND BOUCHER 265

Chromodoris elisabethina Bergh, 1877 white with gray-brown lamellae and distinct internal white cores. Branchial stalks 5-9, REFERENCE: Johnson (1982 : 92); Rudman gray-brown with internal beads of opaque (1982a: fig. IG). white. MATERIAL: More than 100 specimens ob DEVELOPM ENT: Egg masses, described in served on lagoon pinnacles and on the steep detail by Boucher (in press), possess extra slopes of the seaward reef of Enewetak and capsular yolk bodies similar to those of Kwajalein, at depths of 8-20 m. Most are Chromodoris decora. found -in ledges and caves in sheer cliffs, and feed on two unidentified sponges, Heteronema DISTRIBUTION: Sri Lanka (Kelaart 1858a), sp. and Dysidea sp. Indonesia (Bergh 1905), New Caledonia (Ris bee 1928), Japan (Baba 1953), Enewetak DESCRIPTION: Marshall Islands specimens (Marcus and Burch 1965), Guam (Carlson match those reported by Rudman (1982a). and Hoff 1981). DEVELOPMENT: Egg masses deposited on 22 REMARKS: Marcus and Burch (1965) dis August 1981 were cream-yellow and coiled cussed the synonymy of this species. They flat on the substrate rather than attached by followed Eliot (1909) and Pruvot-Fol (1951) one edge as is more typical of chromodorids. in regarding Chromodoris flammulata Bergh, Ova cream-yellow, irregularly oval, mea suring 1905, and C. lactea Bergh, 1905 as synonyms 75-105 Jim individually within capsules 105 of C. fidelis. We agree that C. fidelis and C. 125 Jim in diameter. Approximately 50 ova flammulata are synonymous, but have some per mm? and 5000 within mass. Free-swim doubts about C. lactea. The alcohol-preserved ming veliger larvae with transparent type-one C. lactea described by Bergh (1905) differs shells. in rhinophore and gill coloration from our DISTRIBUTION: West Pacific (reviewed by ethanol-preserved C. fi delis, and Bergh's Rudman 1982a) . radular figures of C.flammulata and C. lactea are far from identical. Chromodoris lactea should be regarded as a distinct species until Chromodoris fidelis (Kelaart, 1858a) living animals are found with preserved color REFERENCE : Eliot (1906b). ation and radular characteristics matching Bergh's description. MATERIAL: This is one ofthe most common nudibranchs at Enewetak. More than 200 individuals have been observed in a variety of Chromodoris geometrica Risbec, 1928 habitats, usually beneath loose coral rubble on inshore lagoon reefs and pinnacles. It is not Figure 6 unusual to find 10 or more in a single search REFERENCE: Risbec (1928) . hour. Specimens have also been found at Kwajalein and Bikini. These animals prey on MATERIAL: Over 60 specimens, ranging in at least two sponge species, including Aplysilla length to 35 mm, have been found on Ene violacea Lendenfeld, 1883. wetak lagoon reefs and pinnacles, under dead coral in 2-15 m of water. Several were photo DESCRIPTION: Typical coloration has been graphed by D. Johnson at Kwajalein. We have well figured in illustrations by Eliot (1906b) observed these chromodorids preying on at and as Chromodoris flammulata by Bergh least two species of sponges, including (1905) and Risbec (1928). Marshall Island Aplysilla violacea. specimens frequently exhibit color variation not mentioned in previous studies: the irregu DESCRIPTION: Marshall Islands specimens larly wide submarginal band that is usually dorsally light gray, with scattered compound maroon varies from yellow to dark reddish white pustules. A dark purple-black line forms purple, and the rhinophores are oftranslucent an irregular oval around the rhinophores and 266 PACIFIC SCIENCE, Volume 37, July 1983

1 4 16 25 26

FIGURE 6. Chromodoris geometrica Risbec. Right-half row of radular teeth (scale bar in JIm). gills, usually connected middorsally by a masses nearly surrounding the capsules. transverse line of the same color. Nine to 14 About 500 ova per mm? and 12,000 in a diffuse purple-black lines radiate from the typical mass. oval toward the margins. In larger specimens, DISTRIBUTION: New Caledonia (Risbec purple-black areas are studded with bright 1928), eastern Australia (Allan 1947), and white granules. Margin wide, white, thinly Enewetak (Young 1967). edged with purple anteriorly. Underside of mantle skirt translucent white, except ante REMARK S: Rudman (l973b) record s a riorly, where it is deep purple. Sides of foot Chromo doris cf. geome trica from East Africa with a longitudinal purple-black lineon a gray that differs considerably in coloration from ish background. Rhinophores and branchiae our specimens, and suggests that perhaps the yellow to green. In one of our specimens the coloration is quite variable. We feel that the gillswere inverted, projecting downward from differences in radular formulae , tooth shape, the underside of the posterior edge of the and external coloration are probably too mantle, as described for Chromodoridiella extreme to consider Rudman's specimens mirabilis (Eliot 1905a). conspecific. Radular formula of a 14mm specimen 32 x 26.1.26. Central tooth faint, present in Chromodoris inornata Pease, 1871 only a few rows. Innermost laterals with a large cusp and three small inner and five to six SYNONYM: Chrom odoris lilacina Young, outer denticles. Remain ing teeth hamate with 1967; Kay and Young, 1969 (non-Gould, outer denticles (Figure 6). 1852). REFERENCE: Rudman (1973b ); Bertsch and DEVELOPMENT: Two loosely coiled egg Johnson (1981: 54, as C. lilacina). masses deposited on 18 November 1981 were peach to orange in color. Ova white, measur MATERIAL: More than 75 specimens rang ing 90-105 flm individually within capsules ing in length to 35mm, observed on shallow 105-120 flm in diameter. Extra capsular yolk lagoon reefs of Enewetak , Kwajalein, and bodies orange, varying from small caps to Bikini. Marshall Islands Opisthobranchia-c-Jonxsox AND BOUCHER 267

1 5 25 36 42

100

FIGURE 7. Chromodoris marginat a (Pease). Right-half row of radular teeth (scale bar in /lm ).

DEVELOPMENT : On 12 December 1981, a REFERENCE: Kay and Young (1969 , as 10mm individual deposited an egg mass con Chromodoris trimarginata). sisting of six close set, spiral whorls laid flat MATERIAL: More than 60 specimens have on the substrate. Ova light yellow measuring been found at Enewetak, and one at Bikini, approximately 75 pm individually within cap mostly under dead coral rocks in 1-8 m of sules 90 J1.m in diameter. Free-swimming veli water along lagoon reefs. Individuals range to ger larvae with transparent, type-one shells. 34 mm in length. C. marginata feeds upon at DISTRIBUTION: West Pacific (reviewed by least three sponge species, including Aplysilla Rudman 1973b), Enewetak (Young 1967), violacea and A . sulphurea Schultz, 1878. and Hawaii (Kay and Young 1969). DESCRIPTION : Body soft, elongate oval, REMARKS : Specimens identical with these with a relatively wide mantle skirt draping Marshall Islands chromodorids were reported down around the foot. Mantle usually bright from Enewetak (Young 1967) and Hawaii white with maroon marginal and orange sub (Kay and Young 1969) as Chromodoris lila marginal bands. Larger specimens with irreg cina (Gould, 1852). However, Gould's Doris ular, translucent, almost grayish spots on a lilacina measured 85 mm in length , was lilac in slightly pustulose dorsum, and a bluish-white coloration with large, discrete darker spots , edging on the mantle margin. Rhinophores and possessed feathery, tripinnate branchiae with transparent peduncles, light reddish wider than the body. Young (1967) suggested clubs, each bearing 11-14 white lamellae. that Doris amabilis Kelaart, 1858 and Chro Seven to nine simply pinnate branchial stalks, modoris porcata Bergh, 1888 may be synony light reddish with rachides and pinnae edged mous with his " Chromodoris lilacina," Both with white. species are distinctly different in coloration Radular formula of a 16 mm specimen , 44 from C. inornata and may represent separate x 42.1.42. Central tooth small, triangular, species. and inconspicuous in small individuals. In nermost laterals with a large cusp and two equal inner, and four to five outer denticles Chromodoris marginata (Pease, 1860) (Figure 7). Most remaining teeth sharply Figure 7 hooked, each with a long cusp and three to 268 PACIFIC SCIENCE, Volume 37, July 1983

five denticles near the curved posterior edge. (1905) specimen was yellowish and grayish Several teeth near the outermost nearly bicus dorsally and had a red margin and light violet pid, the first denticle as large as the cusp and rhinophores and gills. Winckworth's (1946) larger than the remaining denticles. Outer C. trimarginata had a white outer margin and most teeth with-zero to three denticles. red dorsal spots. The radulae also vary among Buccal elements curved bifid rods approxi all these specimens. Additional research may mately 38 }J.m in length. Some appeared to be necessitate a reevaluation, but for the present inflated just behind the forked end, and had we follow other authors in considering all of thickened, sculptured bases. these records to be varieties of a single species.

DEVELOPMENT: An egg mass deposited by a 34mm specimen on 30 January 1982 con sisted of three loose whorls attached by one Chromodoris tryoni (Garrett, 1873) edge to the substrate. Ova bright yellow with Figure 8 orange, caplike extra capsular yolk bodies. REFERENCE: Young (1967, as Hypselodoris Ova measuring about 125 }J.m individually within capsules 160}J.m in diameter, arranged tryoni). in rows perpendicular to the long axis of the MATERIAL: More than 40 specimens vary ribbon. About 115ova per mm? and 13,800 in ing in size to 65 x 14mm observed at Ene the mass. Free-swimming veliger larvae with wetak on the pilings at Medren Pier in 1-5 m transparent, type-one shells. of water, usually feedingon the sponge Dysidea fragilis (Montagu, 1818). A few additional DISTRIBUTION: Hawaii (Pease 1860; Kay and Young 1969), Indonesia (Bergh 1905), individuals and pairs were found among dead coral rocks at Enewetak and Runit Islands New Caledonia (Risbec 1928), India (Winck worth 1946), Japan (Baba 1953), Vietnam and in a lagoon Halimeda patch near Ene wetak. One specimen collected and photo (Risbec 1956), and Australia (Thompson graphed at Kwajalein by J. Hammon. Adults 1972b). are usually paired and queue up when they REMARKS: We are tentatively assigning move, the following individual placing the Pease's name to this species, although our anterior edge of its foot on the tail of the specimens differ from those reported by Kay leader and following directly behind. and Young (1969) and Kay (1979) from the type locality in several respects: (1) the mar DESCRIPTION: Our specimens differ from ginal bands of color are maroon and orange, those of Young (1967) in certain aspects of instead of red-orange and yellow respectively; radular tooth morphology. Young's figure of (2) the rhinophores and gills, which are white the radula shows a smooth, hamate innermost lateral divided into two small cusps at the tip. in the Hawaiian specimens, are tinted with Although the teeth are difficult to distinguish red; (3) Enewetak specimens attain a larger because of crowding, none of the five speci size, becoming pustulose with grayish spots mens we examined possessed bicuspid inner rather than smooth and white; (4) a central most laterals. Instead, these teeth were hamate radular tooth is present; (5) the cusp is more with three to four denticles on either side of prominent and the outer teeth are smoother; and (6) some of the buccal elements have an elongate main cusp (Figure 8), similar to those figured by Bergh (1879)for Chromodoris inflated and sculptured portions. We consider tryoni . our animals to be Chromodoris marginata, in part because of the apparent variability of the DEVELOPMENT: An egg mass deposited by a species, both at Enewetak and throughout 55mm specimen on 3 January 1982 consisted the Indo-west Pacific. Specimens reported by of four loose whorls crenulated on the free Thompson (1972b) were white with a pepper edge. Ova rose-pink measuring 11O-120}J.m ing of gray on the dorsum and had a red individually within capsules 150}J.m in dia margin and gray rhinophores and gills. Bergh's meter. About 90 ova per mm? and 130,000 in Marshall Islands Opisthobranchia-s-Jorrxsox AND BOUCHER 269

19 40 60

FIGURE 8. Chromodoris tryoni (Garrett). Right-half row of radular teeth (scale bar in {tm).

FIGURE 9. Hypselodoris australis (Risbec). Right-half row of radular teeth (scale bar in jim) . the entire mass. The larvae are free-swimming morphology of the radular teeth traditionally veligers. determines the generic placement of chromo dorids. Our radular dissections show unicuspid DISTRIBUTION: Society Islands (Garrett teeth, and we have, at least for the present, 1973), Palmyra (Young 1967), and Guam retained the genus Chromodoris. (Carlson and Hoff 1981). Eliot (l904b) erroneously reported C. tryoni REMARKS: The buccal elements, reproduc from east Africa; his specimen had orange tive system, and external shape of Chromo spots and was probably C. aureopurpurea doris tryoni are clearly Hypselodoris-like Collingwood, 1881. Despite Eliot's (1904b) (Young 1967, Rudman 1977). However, the suggestion that the two are synonymous, C. 270 PACIFIC SCIENCE, Volume 37, July 1983 tryoni and C. aureopurpurea differ sufficiently SYNONYM: Chromodoris decorata Risbec, in shape and coloration that they cannot be 1928. considered conspecific. REFERENCE: Risbec (1928 : pI. 7, fig. 4).

MAT ERIAL : Over 60 individuals ranging to Hypselodoris australis (Risbec, 1928) 40 mm in length observed at Enewetak. They Figure 9 are usually found beneath dead coral rocks and in ledges along lagoon reefs and on pin SYNONYM: Chromodoris australis Risbec, nacles, as well as on the pier pilings at Medren 1928. Island. Specimensphotographed at Kwajalein, REFERENCE: Risbec (1928); Johnson (1982: D. Johnson. Enewetak specimens feed on at 92). least two species of sponges, including Dysidea fragilis. MATERIAL: Over 75 specimens of this com mon species, ranging in length to 18 mm, were DESCRIPTION: Our specimens exhibited observed beneath dead coral lagoon reefs and more color variation than indicated by Risbec on pinnacles. Individuals are often paired and (1928). Color cream-yellow middorsally, with are usually found feeding on an unidentified, a wide orange-red marginal band and a white encrusting, tan-colored sponge. Populations submarginal line that widens at intervals at Enewetak appear to fluctuate : specimens forming fingers of white extending into the are far more common in summer and fall orange-red margin . Two additional longitu months . Specimens photographed at Kwaja dinal white lines beginning between the rhino lein, D. Johnson and J. Wedge. phores and extending around the gills, some times connected by transverse white lines DESCRIPTIO N: Radular formula of a 9 mm specimen 38 x 22.0.22. Teeth long, narrow, middorsally. Notum cream-yellow, usually and difficult to distinguish. Innermost laterals with scattered carmine or purple spots, which bifid with one to three inner and two to are also present along the innermost edge of three outer denticles. Mid-half row teeth bifid the orange-red margin ; laterally, the orange or trifid with distinct outer denticulation red marginal bands are scattered with opaque (Figure 9). white granules in most, but not all, specimens. Sides of foot colored as the dorsum. Rhino DEVELOPM ENT: An egg mass, cream-white, phores cream-white, with three orange rings consisting of three loose whorls attached by and 14-15 lamellae. Seven simply pinnate gill one edge to the substrate, was deposited by a stalks, white with fine red lines and orange 16 mm individual on 12 December 1981. Ova rachides and tips. measure about 80 flm individually within cap Radular formula of a 25 mm specimen sules 90 zmi in diameter. About 325 ova per 52 x 45.0.45. Teeth strongly bicuspid along mm? and 35,000 in the mass. Free-swimming the entire radular row, warranting placement veligerlarvae with transparent type-one shells of this species in the genus Hypselodoris begin hatching in 5 days. (Figure lOA). Buccal elements of the same DISTRIBUTION: New Caledonia (Risbec individual are short, stout, thickened, and 1928), Guam (Carlson and Hoff 1981). ribbed near the top, each with single curved point at one end (Figure lOB). REMARK S: The narrow, elongate radular teeth are reminiscent of Babaina Odhner, 1968, DEVELOPM ENT : Each of the several egg but that genus is characterized by bifid teeth masses observed was deposited in one or two without denticulation (Bertsch 1977). loose whorls, attached by one edge to the substrate. Ova rose-pink, measurin g 80 90flm individually within capsules 125 flm Hypselodoris decorata (Risbec, 1928) in diameter. The largest egg mass measured Figure lOA , B 105mm in length and contained approxi- Marshall Islands Opisthobranchia-s-Jonxsox AND BOUCHER 271

1 7

FIGURE 10. Hypselodorisdecorata (Risbec) . A . Right-halfrow ofradular teeth; B. Buccal element (scale bars in /lm). mately 56,000 ova. Veliger larvae free-swim MATERIAL: Three specimens, the largest 35 ming with transparent type-one shells. x 7mm, found on Enewetak lagoon reefs at depths of 5-8 m. DISTRIBUTION: New Caledonia (Risbec 1928), Vietnam (Risbec 1956), and Guam DESCRIPTION: Enewetak specimens with (Carlson and Hoff 1981). three wide longitudinal bands of white, sep arated by bands of gray that join anterior to the rhinophores. Gray bands usually irregu Hypselodoris infucata (Riippell and Leuckart, larly splotched with yellow, and mantle with 1828) scattered purplish-black spots. Mantle mar REFERENCE: Edmunds (1971); Rudman gin edged with purple and submarginally with (1973b); Bertsch and Johnson (1981: 62, 63). somewhat regularly spaced purplish-black 272 PACIFIC SCIENCE, Volume 37, July 1983

1 7 30 40 43 44

FIGURE II . Hypselodoris mouaci (Risbec). A. Right-half row of radular teeth ; B, C. Buccal elements (scale bars in tim) .

spots surrounded by patches oflighter purple . REFERENCE : Risbec (1930: pI. 1, fig. 2). Rhinophores with white peduncles and red MATERIAL: Over 75 specimens observed at clubs. Eleven to 13 branchial stalks whitish, Enewetak, on pier pilings and dead coral on edged with red. Radula and buccal armature shallow lagoon reefs and pinnacles, to depths as figured by Edmunds (1971). of 20 m. Specimens photographed at Kwa DISTRIBUTION: Indo-west Pacific (Edmunds jalein, D. Johnson and J. Wedge. Enewetak 1971 , Rudman 1973b); Hawaii (Bertsch and specimens range in length to 40 mm and are John son 1979b). frequently observed feeding on the sponge Dysidea fragilis.

Hypselodoris mouaci (Risbec, 1930) DESCRIPTION: Radular formula of a 32 mm individual 67 x 54.0.54, that of a 34mm speci Figure l1A, B, C men, 62 x 44.0.44. Teeth strongly bicuspid SYNONYMY: Chromodoris mouaci Risbec, (Figure IIA), the inner ones with long, pointed 1930. cusps and thickened bases. Outermost teeth Hypselodoris hilaris Marcus rather squat, with small, rounded denticles on and Burch, 1965 (non Bergh, the posterior edge. Buccal elements short and 1890). thickened , with a single large point on one Marshall Islands Opisthobranchia-c-Jonxso» AND BOUCHER 273 end. Thickening on the shaft most commonly as in Figure lIB; occasionally, with one or two subpoints as in Figure II C, and as de scribed by Marcus and Burch (1965). DEVELOPMENT: An egg mass deposited by a 23 mm individual on 18 November 1981 con 3 9 14 sisted of a loose, irregular, brown coil, crenu lated on its free edge. Ova orange, measuring 95-120 Jim individually within capsules only 80 slightly larger. About 150 ova per mm? and 45,000 in the entire coil. Larvae are free swimming veligers with type-one shells.

DISTRIBUTION: New Caledonia (Risbec 1930), Enewetak (Marcus and Burch 1965). 8

REMARKS: Marcus and Burch (1965) erro neously identified this species as Hypselodoris hilaris (Bergh, 1890), a species that differs somewhat in external coloration. The speci mens Baba (1953) identified as Glossodoris hilaris from Japan may also represent H. FIGURE12. Thorunna clitonata (Bergh). A . Right-half mouaci. row ofradular teeth; B. Anteriorview ofinnermost lateral tooth (scale bar in pm) .

Thorunna clitonata (Bergh, 1905) posed ofeight simply pinnate, bladelike stalks Figure 12A, B set in a nearly complete circle; stalks pale SYNONYM: Chromodoris clitonata Bergh, orange at the base and purple at the tips. 1905. Radular formula of an 8 mm specimen 27 x 14.0.14. Innermost laterals extremely wide REFERENCE: Bergh (1905: tafel 5, fig. 16). with two or three denticles on either side of a MATERIAL : Four specimens, ranging from large cusp (Figure 12A, B). Remaining teeth 4 x 1mm to 12 x 2 mm, found on Enewetak hamate, becoming only slightly curved by the lagoon reefs, under dead coral at depths of outermost. 1-4m. DEVELOPMENT: An egg mass deposited by a DESCRIPTION: In motion, the body is elon 12mm specimen on 10July 1982consisted of gate and slender. Mantle margin bearing a a single white whorl attached by one edge to widewhite or light yellowband with a narrow, the substrate. Ova white, measuring 86 Jim irregular, submarginal band of purple, and a individually within capsules 100-115 Jim in wide middorsal white band from between the diameter. rhinophores to the gills. On either side of, and DISTRIBUTION: Indonesia (Bergh 1905), parallel to, the white band is a dark reddish Japan (Baba 1937a), eastern Australia purple line, followed by a relatively narrow (Thompson 1972b), and perhaps Guam band of peach or orange, then a line of light (Carlson and Hoff 1981, as Chromodoris cf. reddish-purple. Light purple background clitonata), New Caledonia (Risbec 1928), and nearly obscured by the dorsal color bands. Vietnam (Risbec 1956). Rhinophores with transparent peduncles and tricolored clubs: light purple at the tip, dark REMARKS: Bergh (1878) erected the chro purple in the middle, and orange basally. modoris genus Thorunna ( = Noumea Risbec, Branchiae situated far posterior, and com- 1928; see Bertsch 1977) for T. furtiva Bergh, 274 PACIFIC SCIENCE, Volume 37, July 1983

2 4 10 16

FIGURE 13 . Thorunna decussata (Risbec). Right-half row of radular teeth (scale bar in Jim).

1878, based on the wide innermost lateral teeth Thorunna decussata (Risbec, 1928) and lack ofbuccal armature. Buccal armature Figure 13 occurs in T. clitonata (see Bergh 1905) and in some other species with wide innermost lat SYNONYMY: N oumea decussata Risbec, erals, but is sufficiently small that, if present in 1928. T. furtiva, it may have been overlooked. We RE FERENCE: Risbec (1928: pI. 8, fig. 6). retain Thorunna because the width of these MATERIAL: This is the most abundant ofthe inner!ll0st teeth seems at least as significant as Enewetak chromodorids. As many as 20 indi the dIfferenc~ between unicuspid and bicuspid viduals were observed per search hour. Well teeth, the pnmary characteristic used to dif over 300 individuals have been found , mostly ferentiate Chromodoris and Hypselodoris. on the lagoonside of Enewetak Island at 2 Unfortunately, there is some confusion as 10m, but some on lagoon reefs ofother islands to the identity of this and similarly colored and on pinnacles. Their abundance fluctuates species. Our specimens resemble the original dramatically; almost none was observed dur description and figure by Bergh (1905), who ing the late summer and fall of1982. Thorunna states that the radula is typical of chromo decussata is found primarily in association dorids, but gives no figure. The color figure with the pink encrusting sponge Dendrilla sp. and radula of Risbec's (1928) Chrom odoris cf. D. cactus (Selenka , 1867), on which we clitonata are quite different from those of our have observed it feeding both in the field and animals. Adding to the confusion, Pru vot-Fol in aquaria. Less commonly, this nudibranch (1951) considered Glossodoris clitonata syn eats the yellow sponge Aplysilla sulphurea. onymous with G. scurra (Bergh, 1874), G. Specimens range in length to 20 mm, although luxuriosa (Bergh, 1875), G. variegata (Pease, they are typically less than 10mm, and are 1871), G. dorsalis (Gould, 1852), and G. lem usually ?bserve? clustered on their prey niscata (Quoy and Gaimard, 1832). Several of sponges ill multiples of 2 to 10 individuals these species differ in size and /or coloration frequently with eggs. Thorunna decussata has from our specimens. Clarification of the been photographed at Kwajalein, D. Johnson. systematics requires comparison of living spe~II~ens and radulae of all these species or DE.SCRIPTION: M.antle bulging conspicuous vaneties. ly midlaterally, bnght white to pink, usually Marshall Islands Opisthobranchia-i-Joaxsox AND BOUCHER 275 with a variable number of irregularly placed, Thorunna norba (Marcus and Marcus, 1970) small, diffuse spots of orange. Margin edged SYNONYM: Noumea norba Marcus and with a broken red line and a narrow, white Marcus, 1970. submarginal band. Rhinophore and gills tipped with varying amounts of orange-red. REFERENCE: Marcus and Marcus (1970). Combined radular formula of two 8 mm MATERIAL: About 15 specimens ranging to individuals 24-26.16.0.16. Innermost laterals 12 x 3mm in size observed at Enewetak, each with a main cusp with three small inner usually beneath dead coral on shallow lagoon and four larger outer denticles. Outermost reefs and pinnacles. laterals long and thin, each with two small denticles on the tip (Figure 13). DESCRIPTION: Mantle elongate oval, round ed in front and back; thin, somewhat flaring DEVELOPMENT: Ova from several egg but notespecially wide, and colored by a wide, masses varied in color from cream to bright faint yellow band; dorsum orange-brown, yellow, and measured 95-1051lm individually approaching peach in color, with two patches within capsules 125-140llm in diameter. of faint yellow to white, one roughly teardrop Approximately 130 ova per mm? and 8800 in shaped around the gills with the wider end the entire coil. Free-swimming veligers with posteriorly, and the other a band extending transparent type-one shells. from between the rhinophores to the middle DISTRIBUTION: New Caledonia (Risbec of the dorsum. In one specimen, these two 1928). faint yellow patches were connected by a thin line ofthe same color. Between orange-brown REMARKS: Our specimens are readily rec dorsum and yellowish margin a submarginal ognized by the distinctive midlateral bulges, row of alternating diffuse purple and white and are certainly conspecific with those de patches , becoming a narrow, submarginal, scribed by Risbec (1928)from New Caledonia. purple band around the anterior and posterior They are similar to those Kay and Young ends. Foot pink. Rhinophores with a translu (1969) identified from Hawaii as Hypselodoris cent orange peduncle and about 14 orange peasei (Bergh, 1880), a species originally lamellae. Branchiae pale orange with five described by Pease (1860) as Doris pieta (pre simply pinnate stalks. occupied) from the Hawaiian Islands. How Radular formula of an 8 mm specimen 31 ever, none of the more than 300 specimens x 14.0.14. Innermost lateral teeth each with examined during this study matches Pease's fivedenticles on either side of a main cusp, as (1860) color description of D. pieta or opposed to the six inner and seven to eight approaches it in size. outer denticles noted in the original descrip Chromodoris simplex Pease, 1871 , described tion. The posterior surface of the innermost from the Society Islands, may be an earlier laterals between the main cusp and the outer name for Thorunna deeussata. Pease's descrip denticles deeply grooved . tion of a three-fourths inch (19mm) animal, DEVELOPMENT: An egg mass deposited by a pale pinkish, white toward the margin, edged 10mm individual consisted of three close-set with a broken red line, and with one dorsal red whorls attached by one edge to the substrate. spot and red-tipped rhinophores and gills,falls Ova cream-white measuring approximately wellwithin the range of color variation ofour 95 11m individually within capsules 11011m in specimens. Pease (1871) did not mention the diameter. About 300 ova per mm" and 7500 in midlateral bulges, however, and until speci the mass. mens can be collected from the type locality for comparison, we retain the name T. REMARKS: Marcus and Marcus (1970)sum deeussata . marized the differencesin the innermost lateral The animal figured byAbe (1964)as Noumea teeth of the known species of Noumea, placing deeussata does not appear to be conspecific importance on the number of denticles on with our animals. either side of the main cusp. While the den- 276 PACIFIC SCIENCE, Volume 37, July 1983

ticulation in our specimens is not exactly the Approximately 165 ova per mm? and 39,600 same as that of Marcus and Marcus's N. in the ribbon. Over a period of 10 days the norba, the difference of a few denticles may 40mm specimen was observed to deposit eight reflect regional variation and is alone insuf eggmasses containing a total of approximately ficient to warrant a new species designation. 750,000 ova. Free-swimming veligers with Our record of this species is apparently the type-one shells begin to hatch in 6 days. first since its original description from a single DISTRIBUTION: Hawaii (Kay and Young preserved specimen collected in Fiji. 1969).

Cadline/la ornatissima (Risbec, 1928) Dendrodoris elongata Baba, 1936 REFERENCE: Baba (1949: pI. 22, fig. 79). REFERENCE: Baba (1949: pI. 28, fig. 104) MATERIAL: More than 100specimens, rang MATERIAL: About 30 individuals observed ing to 25 x 8 mm in size, observed on Ene at Enewetak, Bikini, and Kwajalein. They wetak lagoon patch reefs and pinnacles, range in length to 75mm and are usually found usually beneath dead coral in 3-20 m of water. beneath dead coral on lagoon reefs at depths These nudibranchs feed on the sponge Hali ofO-lOm. sarca metabola deLaubenfels, 1954, and on a thin, encrusting, light brown sponge species. DESCRIPTION: The Marshall Islands' speci Also collected at Bikini. Specimens photo mens possess the extremely elongate shape graphed at Kwajalein, D. Johnson. and slightly pustulose dorsum Baba (1936) described as characteristic of Dendrodoris DEVELOPMENT: Egg masses of this species elongata. Baba (1936) also listed the "peculiar contain discrete extra capsular yolk bodies of coloration" as one of the distinguishing fea 15-30 pm diameter scattered among the ova. tures of this species, although his description Details of ova and development are discussed was based on a single individual. Coloration by Boucher (in press). of Marshall Island specimens variable, from DISTRIBUTION: New Caledonia (Risbec translucent white to brown with numerous 1928), Japan (Baba 1949). diffuse, dark brown spots that consist of aggregations of tiny brown flecks scattered lessdensely over the entire dorsum; scattered, FAMILY DENDRODORIDAE white, star-shaped spots less evident on some specimens; low dorsal pustules tipped with white or dark gray. Foot and underside of the Dendrodoris coronata Kay and Young, 1969 mantle white with scattered brown spots. REFERENCE: Kay and Young (1969). DEVELOPMENT: An egg mass deposited by a MATERIAL: At least 25 specimens observed 45 mm individual on 17 December 1981 con at Enewetak, Rongelap, and Kwajalein, sisted of a wide, cream-yellow ribbon in six mostly beneath dead coral on lagoon reefs at loose whorls, slightly crenulate on the free depths of I-10m. One individual measured edge. Ova measure 160-170llm individually 40 x 26 mm, others ranged to 20 x 14mm. within capsules 240llm in diameter. There were 28 ova per mm? and about 35,000 in the DESCRIPTION: Background color of the Marshall Islands' specimens translucent egg mass. white to pale orange, with varying amounts of DISTRIBUTION: Japan (Baba 1936), New dorsal black and white pigmentation. Caledonia (Risbec 1953), Vietnam (Risbec 1956),and Hawaii (pers. obs.). DEVELOPMENT: A 12mm individual depos ited an egg mass 13mm in diameter on 17 REMARKS: Considering the color variation August 1981. Ova yellow, 50-60llm individ in other dendrodorids, such as Dendrodoris ually with capsules 75-80llm in diameter. nigra (see Edmunds 1971), D. rubra (see Col- Marshall Islands Opisthobranchia-c-Joaxsox AND BOUCHER 277

coral on the intertidal reefat low tide, in June and September 1982.

DISTRIBUTION: Indo-west Pacific, including Zanzibar (Eliot 1905b), Mauritius (Bergh 1889), Sri Lanka (Kelaart 1858a), India (Alder and Hancock 1864). Vietnam (Risbec 1956), Indonesia (Bergh 1905), Japan (Baba 1936, 1937a, 1949), and Hawaii (Pease 1860; Kay and Young 1969). 200

FIGURE 14. Gymnodoris ceylonica (Kelaart). Right FAMILY POLYCERIDAE half row of radular teeth (scale bar in 11m). Gymnodoris ceylonica (Kelaart, 1858b) lingwood 1881), and D. tuberculosa (see Figure 14 Bertsch and Johnson 1"981), it is not surprising SYNONYMY : see MacNae (1958). that D. elongata should show the variation in color that we have observed. REFER ENCE : Risbec (1928: pI. 7, fig. 11). MATERIAL: On 20 September 1981, seven Dendrodoris nigra (Stimpson, 1855) specimens were observed crawling up a sandy REFERENCE: Edmunds (1971); Bertsch and lagoon slope or drifting in a strong tidal cur Johnson (1981: 72). rent in 6-15 m of water near Medren Island, Enewetak. Three days later we observed more MATERIAL: Over 50 specimens found at than 80 specimens, ranging to 80 mm in length, Enewetak, Bikini, Ujelang, and Kwajalein, on lagoon sand flats near Enewetak Island. usually beneath dead coral rocks on intertidal These nudibranchs were actively crawling and shallow subtidal lagoon reefs to depths of obliquely shoreward (northeast) in a loose 3m. column at a depth of 25 m. Within one week, DEVELOPMENT: A 16 mm individual depos the gymnodorids had completely disappeared ited an egg mass on 16 August 1981. Ova light from the collection site. This migration may yellow, measuring 60-70,um individually with have been related to reproduction, as all the in capsules 70-85,um in diameter. Approxi specimens observed were ripe. Eliot (1904c) mately 360 ova per mm? and 7000 in the noted that Gymnodoris crocea (Bergh, 1889) ribbon. Free-swimming veliger larvae begin to "flocks to shallow water for the deposition of hatch in 4 days. Apex of larval shell dark spawn." On 28 February 1982, possibly fol brown and the rest translucent tan. lowing another shoreward migration, numer ous individuals of G. ceylonica were observed DISTRIBUTION: Indo-west Pacific (Edmunds in 5-6 m ofwater on the lagoonside ofMedren, 1971). mostly depositing egg masses on and under dead coral rocks. These opisthobranchs prey Dendrodoris tuberculosa (Quoy and Gaimard, on Stylocheilus longicaudus. 1832) DESCRIPTION: Radular formula ofa 40 mm SYNONYMY : Doris carbunculosa Kelaart, individual 19 x 23.0.23 . Large innermost 1858. D. rugosa Pease, 1860. lateral tooth with a large, clublike base. Teeth smaller laterally, the outermost as narrow, REFERENCE: Kay and Young (1969); pointed spikes (Figure 14). Although the ex Bertsch and Johnson (1981: 73). ternal morphology of this species is quite dis MATERIAL : Four specimens, the largest 180 tinctive, tooth shape is variable, especially in x 140mm, found at Kwajalein, under dead the shape ofthe base, among specimens from 278 PACIFIC SCIENCE, Volume 37, July 1983

DESCRIPTION: Our specimens differed from those of Young (1967)only in that the radular teeth are slightly straighter, with more distinc tivebases.The base of a midlateral tooth forms a basal cusp, present in varying degrees on all but the innermost tooth (Figure 15).

DEVELOPMENT: An egg mass deposited on 31 January 1982 by a 17mm individual con sisted of three whorls attached by one edge to the substrate. Ova yellow, measuring approxi 3 mately 175.um individually within capsules 215.um in diameter. About 50 ova per mm? FIGURE 15. Gymnodoris citrina (Bergh) . Two teeth and 6500 in the mass. from the right-h alf row. DISTRIBUTION: Palau (Bergh 1877), Japan (Baba 1930, as Gymnodoris japonica; 1949), different regions (see figures in Eliot 1904c, Tahiti (Marcus and Marcus 1970), Australia Bergh 1905,Risbec 1928,and MacNae 1958). (Burn 1975), Samoa (Eliot 1900), Enewetak (Marcus and Burch 1965, as G. bicolor; DEVELOPMENT: Specimens collected on Young 1967), the Solomon Islands (Miller both 20 and 23 September 1981 deposited egg 1969), and Guam (Carlson and Hoff 1981). masses in aquaria. Ova arranged in clusters of20-40 in a double spiral within a gelatinous string, the clusters 0.5-0.6 mm in greatest dia Gymnodoris okinawae Baba, 1936 meter. Ova yellow-orange, measuring approx imately 107.um individually within capsules Figure 16 170.urn in diameter. REFERENCE: Kay and Young (1969); DISTRIBUTION: Sri Lanka (Kelaart 1858b), Bertsch and Johnson (1981 : 79). Zanzibar (Eliot 1904c), South Africa (Mac MATERIAL: Twelve specimens, ranging to Nae 1958), Indonesia (Bergh 1905), New 25 x 3 mm from Enewetak, usually beneath Caledonia (Risbec 1928), Australia (Burn dead coral rocks on lagoon reefs and pin 1975), the Society Islands (Pease 1871), and nacles at depths of 4-10 m. Three additional Guam (Carlson and Hoff 1981). individualscollected at Kwajalein, K. DeGroff and D. Johnson. Unlike the Hawaiian speci Gymnodoris citrina (Bergh, 1877) mens of Gymnodoris okinawae, which eat elysiid sacoglossans (Kay and Young 1969), Figure 15 our specimens prey on small cephalaspideans. REFERENCE: Young (1967); Baba (1949: Several species of Elysia offered to G. oki pI. 11, fig. 37,38). nawae in aquaria were not accepted .

MATERIAL: More than 150specimens, rang DESCRIPTION: The Marshall Islands' speci ing in length to 30mm, found on lagoon reefs mens differ from previous records in some and pinnacles at Enewetak, Bikini, and Kwa aspects of coloration and radular morphol jalein, beneath dead coral rocks at depths of ogy. Body translucent, cream colored, scat 2-15 m. Specimens have been observed in the tered with low, white pustules, between which field and in aquaria feeding on other gymno are rather dense, irregular rows of bright dorids, including conspecifics, with which orange-red spots, forming a network around they copulate while, at the same time, con the pustules. Radular teeth of a 20 mm Ene suming them as prey. These opisthobranchs wetak individual smaller than those of larger also eat the egg masses of other species of specimensreported by Kay and'Young (1969). gymnodorid. Innermost laterals are proportionately larger Marshall Islands Opisthobranchia-e-Jonxsox AND BOUCHER 279

2 8 16

100

FIGURE 16. Gymnodoris okinawae Baba. Right-half row of radular teeth (scale bar in 11m). than the outer teeth, which do not increase in dead coral on lagoon reefs at depths of3-5 m. sizeto the outermost laterals; instead, the teeth The largest measured 16 x 3 mm, but most do are largest near the innermost, then become not exceed 10mm in length. slightly smaller outward (Figure 16). DISTRIBUTION: Pacific (Bergh 1877), Ha DISTRIBUTION: Japan (Baba 1936), Hawaii waii (Kay and Young 1969). (Kay and Young 1969).

REMARKS : The slight differences in colora Gymnodoris striata (Eliot, 1908) tion, radular morphology, and diet are prob Figure 17 ably insufficient to consider the Marshall Islands' specimens separate from Gymnodoris SYNONYMY: Trevelyana striata Eliot, 1908. okinawae. Analogium striatum (Eliot) Risbec, 1928. REFERENCE: Risbec (1928); Johnson (1982: Gymnodoris plebeia (Bergh, 1877) 93, as Analogium striatum). REFERENCE: Kay and Young (1969). MATERIAL : More than 50 specimens, rang MATERIAL: At least 40 specimens observed ing in size to 55 x 22 mm, observed at Ene at Enewetak and Kwajalein, usually beneath wetak, usually in sandy areas of lagoon reefs 280 PACIFIC SCIENCE, Volume 37, July 1983

longitudinally arranged lamellar pinnae; plates and pinnae edged with orange. Radular formula of a 50 mm specimen 16 x 19.0.19. Innermost laterals small and bicus pid; remaining teeth elongate and smooth, the inner two-thirds with sharp, outer, basal cusps 1 (Figure 17).

DEV ELOPM ENT: An egg mass deposited by a 40 mm individual on 10 December 1981 con sisted ofthree loose, cylindrical whorls, lightly 2 attached to the substrate. Ova bright yellow, measuring approximately 240.urn,individually encased in capsules 250.urn in diameter. About 14 ova per mm? and 2900 in the coil. 8 13 DISTRIBUTION: Red Sea (Eliot 1908), New Caledonia (Risbec 1928),Japan (Baba 1937b), Fiji (Marcus and Marcus 1970), and Australia (Burn 1975). 350 REMARKS: Risbec (1928) erected Analogium for species with the peculiar branchial arrange F IGURE 17. Gymnodoris striata (Eliot) . Right-halfrow ment of Gymnodoris striata. MacNae (1958) of radular teeth (scale bar in .urn). questioned the need for this genus, stating that the gills of one of his specimens of G. bicolor also "appear to be arranged in a transverse row and pinnacles at depths of 3-25 m. Gymno in the preserved specimen," a statement which doris striata preys voraciously upon Plako suggests that he misunderstood Risbec's (1928) branchus ocellatus, both in the field and in description of peculiar lamellate plates in living aquaria. specimens. The distinctive bicuspid inner teeth and unique arrangement of the gills may war DESCRIPTION: External descriptions pro rant the separate generic designation, but we vided by different authors vary. In our speci retain Gymnodoris until we complete studies of mens the body is high sided, soft in texture, several other, apparently undescribed species widest at midbody, rounded anteriorly, and from Enewetak with similar gill structure. tapers to a blunt point posteriorly. Mantle margin distinct, narrow, and thin. Background Nembrotha kubaryana Bergh, 1877 color translucent white, densely speckled with opaque white; foot and mantle margins bright REFERENCE: Bergh (1877). orange. Middorsal, elevated ridge originating MATERIAL: Specimens range to 60 mm in behind the rhinophores and extending pos length at Kwajalein and are most often in teriorly to the gillsbright orange; shorter ridges ledges and small caves on lagoon pinnacles on one or both sides of the middorsal ridge. and on the steep, seaward slope at depths of Posterior to the gills one or more broader 8-20m. ridges oforange extend about one-third ofthe way to the tail. Tail with an angular middorsal DESCRIPTION: Body soft, slender, elongate, keel, edged with orange. An additional ridge rounded anteriorly, highest and widest at mid of orange along each side of the foot ; bottom body, tapering to a blunt point posteriorly. of foot white. Rhinophores close set, with Background color black ; dorsum and sides transparent peduncles and orange clubs, each with large, scattered, irregular, green spots, bearing about 15 lamellae . Branchiae distinc some ofwhich combine forming a short mar tive, near midbody, and comprised ofa trans gin around the cephalic hood; foot margins verse series of five to nine parallel plates with and rhinophore sheaths brilliant red. Rhino-

• as@! n·bt¥M §# 'Sf!! ¥ii¢NttAU ,¢il#iI!!Jgi@mMe u fli¥ffi#¥!il!i mg;re.tftl!H1ilfiffif~ Marshall Islands Opisthobranchia-s-Joaxsox AND BOUCHER 281

0000

200

FIGURE 18. Plocamopherus cey lonicus (Kelaa rt). Entire right-ha lf row of radular teet h (scale bar in /lm) . phores rodlike , black, with 15- 20 bright red most laterals overlapping hooks, each with a lamellae. Branchiae located at midbody, of large denticle. Remaining teeth flat plates, five black, bipinnate stalks, edged with brilliant some with posterior keels (Figure 18). red. DISTRIB UTION: Sri Lanka (Kelaart 1858a, REMARKS: Our specimens match the origi as Poly cera cey lonica), eastern Australia nal description by Bergh (1877) from Palau, (Thompson 1975), and New Caledonia Bergh's (1905) color illustration of Nembrotha (Risbec 1928). kubaryana from Indonesia shows an animal REMARKS: Like Plocamopherus tilesii Bergh, more longitudinally lined with green rather 1877, the body has several patches that phos than spotted. phoresce when the animal is disturbed. These animal s are capable of swimming by laterall y Plocamopherus ceylonicus (Kelaart, 1858a) bending the body and using a flattened , pos Figure 18 terior keel as a fin.

REFERENCE: Thompson (1975); Risbec (1928: pI. 4, fig. 6). FAMILY AEGIRETIDAE

MATERIAL: Six specimens, ranging in size to 23 x 5 mm, at Enewetak , all on the lagoon Aegires villosus Farran, 1905 side of Enewetak Island, beneath dead coral, REFERENCE: Edmunds (1971); Baba (1955: at depths of 2-5 m in September and December pI. 4, fig. 13). 1981. MATERIAL: One specimen, 9 x 1mm,Ene DESCRIPTIO N: Radular formul a of a 14mm wetak, lagoon side Jinimi Island, under dead individual 12 x 7.3.0.3.7. The three inner- coral , 5 m, 24 August 1982. 282 PACIFIC SCIENCE, Volume 37, July 1983

DISTRIBUTION: Sri Lanka (Farran 1905), MATERIAL: More than 150specimens, rang East Africa (Edmunds 1971), New Caledonia ing in size to 30 x 9 mm, observed . These (Risbec 1928), and Japan (Baba 1955). nudibranchs crawl about in the open during the day. At Kwajalein, Enewetak, and Bikini they are found on lagoon pinnacles and the FAMILY GONIODORIDIDAE seaward slopes at depths of5-20 m, and occa sionally along interisland lagoon reefs.

Goniodoris joubini Risbec, 1928 DEVELOPMENT: A double spiral egg mass about 10mm in diameter contained ova mea REFERENCE: Risbec (1928: pI. 5, fig. 2). suring 170-200,um individually within cap MATERIAL: More than 50 specimens, rang sules 230-260,um in diameter. Approximately ing in length to 17mm, observed beneath dead 33 ova per mm? and 1850 in the mass. Free coral on shallow, inshore reefs of Enewetak, swimming veliger larvae with transparent Kwajalein, and Bikini. brown, type-one shells begin hatching in 5 days. DESCRIPTION: Body coloration rather var iable: specimens are typically cream-white, DISTRIBUTION: Indo-west Pacific (Baba and irregularly spotted with dark purplish-black, Hamatani 1975), Hawaii (Bertsch and Johnson white, and yellow; darker spotting varying in 1981). density, nearly obscuring the cream back ground in some individuals. REMARKS: Individuals of this species pro duce a grayish-white film ofmucus, which has DEVELOPMENT: An egg mass deposited by a a strong odor and kills other organisms placed 10mm specimen possessed white ova measur in the same closed container. The sponge ing 120-150,um individually within capsules prey, tentatively identified as a species of 180-210,um in diameter. About 30 ova per Hymeniacodon, produces the same odor. mm? and 350 in the entire mass. Free swimming veliger larvae with transparent type-one shells begin hatching in 6 days. Phyllidia varicosa Lamarck, 1801 REFERENCE: Edmunds (1971); Bertsch and DISTRIBUTION: New Caledonia (Risbec Johnson (1981: 74, 75). 1928), Hawaii (Kay and Young 1969, as Goniodoris sp. cf.joubini; pers. obs.). MATERIAL: More than 50 specimens of Phyllidia varicosa, ranging in size to 56 x 21 mm, observed at Enewetak and Kwajalein Goniodoridiella savignyi Pruvot-Fol, 1933 in a variety of habitats. These nudibranchs, REFERENCE: Baba (1960). like those ofP.pustulosa, are frequently found exposed on the substratum and secrete a MATERIAL: Two specimens, 4 x 1mm, grayish mucus toxic to other organisms (see Enewetak, lagoonside Enewetak Island, under Johannes 1963). dead coral, 3-5 m, 19 September and 22 November 1981. DISTRIBUTION: Indo-west Pacific (Edmunds 1971). DISTRIBUTION: Gulf of Suez (Pruvot-Fol 1933); Japan (Baba 1960).

FAMILY ARMINIDAE

FAMILY PHYLLIDIIDAE Dermatobranchus fortunata Bergh, 1888

Phyllidia pustulosa Cuvier, 1804 REFERENCE: Bergh (1888).

REFERENCE: Baba and Hamatani (1975); MATERIAL: Generally, 5-10 specimens Bertsch and Johnson (1981: 76). ranging in size to 16 x 4mm, can be found

.n sa;p & 1'11#1 W M@ft M#¥-#UW!lJii!ilfj '''Y iN,," . "4Z¥4i!i1&!I1&D Marshall Islands Opisthobranchia-i-Joaxso» AND BO UCHER 283

per search hour at Enewetak and Kwajalein, Marianina rosea (Pruvot-Fol, 1930) beneath dead coral on the exposed interisland REFER ENCE: Carlson and Hoff (1973, as reef flat at low tide. White egg masses were Aranucus bifidus Odhner, 1936). observed throughout the year. MATERIAL : More than 150specimens found DESCRIPTION: Body flattened , elongate, beneath dead coral rocks on shallow lagoon widest anteriorly, tapering to a blunt point reefs at depths of 1-5m, at Enewetak, Kwa posteriorly. Dorsum white, faintly pustulose, jalein , Bikini, and Ujelang. frequently with faint brown lines scattered between the pustules. Lateral margins with DISTRIBUTIO N: New Caledonia, Australia eight irregularly shaped , black blotches, con (Burn 1974), Gilbert Islands (Odhner 1936), taining small, white pustules. Anterior margin and Guam (Carlson and Hoff 1973). centrally indented, and , except for this inden tation, colored bright orange, as is the lateral margin on those sections adjacent to the black AEOLIDIACEA4 blotches. Rhinophore sheaths extend upward from just within the anterior mantle indenta Flabellina alisonae Gosliner, 1980 tion, white at the base, and crowded with vertical, short lines oforange, black, and white REFERENCE: Gosliner (1980); Bertsch and distally. Rhinophores narrow, orange with Johnson (1981: 88). white tips. Radula as figured by Bergh (1888). MAT ERIAL : More than 60 specimens ob DISTRIBUTIO N: Indonesia (Bergh 1888). served at Enewetak, Kwajalein, Bikini, and Ujelang. They and their egg masses are usually found beneath dead coral rocks on exposed reef flats at low tide. Rarel y, specimens are FAMILY TRITONIIDAE found subtidallyon lagoon reefs and pinnacles.

DESCRIPTION: The lateral teeth of the Mar Bornella adam sii Gray, 1850 shall Islands' specimens are more triangular in shape than those described by Gosliner SYNONYM: see Bertsch (1980). (1980). The denticles on the central teeth are REFERENCE: Bertsch and Johnson (1981 : more elongate and separated by grooves that 85). extend well into the base of each tooth.

MATERIAL: Over 25 specimens, ranging in DISTRIB UTIO N: Okinawa (Baba 1936, as size to 50 x 4 mm, observed at Enewetak and Coryph ella ornata Risbec, 1928), Japan (Baba Kwajalein . During the day they are found 1955, as Flabellina ornata), Hawaii and Ene beneath rocks on shallow lagoon reefs, and wetak (Gosliner 1980). nocturnally in ledges on the steep slopes of lagoon pinnacles and the seaward reef. Several individuals were observed feeding on hydroids Favorinusjaponicus Baba, 1949 of the genus Sertularia. REFER ENCE: Kay (1979); Rudman (1980); Bertsch and Johnson (1981: 96, 97). DESCRIPTION: The external characteristics of Bornella adamsii have been described fre MATERI AL : Eight specimens, the largest 12 quently (usually as B. digitata Adams and x 2mm, observed at Enewetak and Kwaja Reeve, 1850). Marshall Islands' specimens lein, all beneath dead coral on lagoon reefs vary from uniform orange-brown to pale and pinnacles, at depths of 1-8 m. These cream with a network of orange and orange brown subapical rings on the dorso-lateral processes. 4 Family names are not ut ilized in this group of nud i branchs becau se of apparent lack of agreement on the DISTRIBUTION: Indo-west Pacific boundaries of the families. 284 PACIFIC SCIENCE, Volume 37, July 1983

aeolids prey on the eggs of other opistho branchs, including Dendrodoris nigra and Hallaxa sp., and tend to pick up the color of their food source (see Kay 1979).

DESCRIPTION: Cerata vary in color from yellow to orange to pink; rhinophores with three annular swellings rather than the two reported for Japanese and Hawaiian speci mens (Baba 1949, Gosl iner 1980). Radular teeth and jaws as figured by Baba (1949). 90 DI STRIBUTION: Japan (Baba 1949), Hawaii (Kay 1979, Gosliner 1980), and East Africa F IGU RE 19. Phestilla lugubris (Bergh). Selected radu lar teeth from a single individual (scale bar in )lm). (Rudman 1980).

REMARKS: Gosliner (1980) stated that this species is similar to Favorinus go uaroi (Risbec, 1928), but differs in the smooth radular teeth; figured by Gosliner (1980), but vary consider Risbec's species has teeth with lateral den ably within a single individual (Figure 19). ticulation. However, as Rudman (1980) has Central cusp variable in size, and may be pointed out, lateral denticles appear to be a absent. Lateral denticles vary in height but variable characteristic in F.j aponicus, as is the are uniformly comb shaped, similar to those case in two closely related species, F. bran figured by Rudman (1981 a). chialis (Rathke, 1806) and F. auritulus Marcus, DISTRIB UTIO N: Indo- west Pacific from East 1955. Since two of our specimens ofF.japoni Africa to Hawaii (Rudman 1981a). cus were light yellow, as described for F. gouaroi by Risbec (1928), it seems likely that they are conspecific. A final determination of Phyllo desmium hyalinum Ehrenberg, 1831 synonymy should await the comparison of F. REFERENCE: Rudman (1981b); Baba japonicus with New Caledonian specimens (1937a: pI. 2, fig. 5). fitting the description of F. gouaro i. MATERI AL: Five specimens, rangin g to 20 x 2.5 mm in size (cerata not included), found Phestilla lugubris (Bergh, 1870) on lagoon pinnacles at Enewetak and Kwa Figure 19 jalein feeding on the soft coral Xenia sp.

REFER ENCE: Rudman (1981 a); Bertsch and DISTRIBUTIO N: Red Sea (Gohar and Aboul Johnson (1981: 90, 91, as P. sibogae). Ela 1957, as Phyllodesmium x eniae), East Africa (Eliot 1905b ; Rudman 1981b), and MAT ERIAL: More than 30 specimens of this Japan (Baba 1937a). cryptic species have been found in association with the scleractinean coral Porites at Enewe tak and Kwajalein. Typically, groups of two Pteraeolidia ianthina (Angas, 1864) or more individuals, frequently with cream SYNONY MY: see Go sliner (1980). colored egg masses, are found beneath coral heads. At Enewetak, Phestilla lugubris is REFER ENCE: Ka y (1979); Rudman (1982b); readily eaten by several species of wrasse Bertsch and Johnson (1981: 98, 99). (Labridae). MATERI AL : Solitary specimens of Pt eraeo DESCRIPTION: Radulae of 17 and 18mm lidia ianthina measuring up to 70mm in length specimens with 40 and 44 uniseriate teeth, are found infrequently in a variety ofhabitats respectively. Teeth ofMarshall Islands' speci at Enewetak and Kwajalein. We have observed mens basically similar to those of specimens them on and under dead coral , on pier pilings, Marshall Islands Opisthobranchia-c-Joaxsox AND BOUCHER 285

DISTRIBUTION: New Caledonia (Risbec 1928), East Africa (Rudman 1980).

Phidiana indica (Bergh, 1896)

REF ER ENCE: Rudman (1980); Bertsch and 100 Johnson (1981: 95, as Caloria militaris).

MATERIAL: About 20 specimens, ranging to 20 mm in length, observed at Enewetak and Kwajalein, usually beneath dead coral , on pier pilings, or on buoy lines at depths of 3-40m.

DISTRIBUTION: Indo-west Pacific (Baba FIGURE 20. Phidiana bourailli (Risbec) . Radular tooth 1969; Gosliner 1980; Rudman 1980). (scale bar in Jim). REMARKS: It is obvious from the literature that there is no clear consensus on the status and on the hull of a sunken ship at depths of of various genera within the Glaucidae or 3-20m. Facellinidae (see Edmunds 1969, Burn and Narayanan 1970, Miller 1974, Gosliner 1980, DEVELOPM ENT: A 70 mm individual depos ited an egg mass on II December 1981 . Ova Rudman 1980). We follow Rudman (1980) in white, spiraled into two loose whorls, measur retaining Phidiana for this species. ing about 140p.m individually within capsules 190p.m in diameter. About 97 ova per mm? Glaucus atlanticus Forster, 1777 and approximately 10,000 in the entire mass. REF ER ENCE: Thompson and McFarlane Free-swimming veliger larvae begin hatching (1967); Bertsch and Johnson (1981: 100). in 5 days. MAT ERIAL: Occasional specimens of Glau DISTRIBUTION: Indo-west Pacific (Kay cus atlanticus are washed ashore on windward 1979; Gosliner 1980), including Truk, eastern beaches of Enewetak. Specimens have also Caroline Islands, and the northwestern Ha been found at Kwajalein by D. Johnson. waiian Islands , especially Nihoa and Necker (pers. obs.). DISTRIBUTION: Circumtropical (Thompson and McFarlane 1967). Phidiana bourailli (Risbec, 1928) Spurilla major (Eliot, 1903b) Figure 20 REFERENCE: Kay (1979, as Berghia major); REFERENCE : Rudman (1980: fig. IB). Rudman (1982b); Bertsch and Johnson (1981: MAT ERIAL: At least 30 specimens, generally 89, as Berghia major). 0 less than 10mm in length, bserved under MATERIAL: One specimen, 23 x 7 mm, dead coral at depths of 1-5 m on lagoon reefs Enewetak, Bokandretok-Medren reef, under of Enewetak and Kwajalein . dead coral, 8 m, 14 July 1981. DESCRIPTION: Radula of a 9 mm specimen DISTRIBUTION: Indo-west Pacific, including uniseriate with 38 teeth. Number and shape of East Africa (Eliot 1903b) and Hawaii (Ka y denticles on the different teeth of a single 1979, Gosliner 1980). specimen vary; on either side of a main cusp are five to eight not necessarily symmetrical REMARKS: Rudman (1982b) believes the denticles. Posterior surface ofeach tooth with characteristics of the genus Berghia Trinchese, a central groove (Figure 20), and minor bumps 1877are adequately covered by the definition at the base ofeach denticle . of Spurilla Bergh, 1864. 286 PACIFIC SCIENCE, Volume 37, July 1983

DISCUSSION Alderson, Lori Bell, Patrick L. Colin, Dennis Devaney , Ruth Dubin, Karen DeGroff, After examining large numbers of living Jeanette Hammon, John and Buffy Harrison, opisthobranchs during this study and previous Dave Hayes, Regina Kawamoto, E. Alison studies in Hawaii (Johnson and Bertsch 1979; Kay, Terry Kerby, D. J. MacDonald, Lee Bertsch and Johnson 1979a, 1979b,1981, 1982; Rousseau, W. B. Rudman, Dan Self, Ann Johnson 1983), we conclude that external Thresher and Judy Wedge. Special thanks to morphology and coloration are generally re Dave Johnson and Jim Wedge of Kwajalein, liable species characteristics. While there is both for specimens and for allowing us access some intraspecific variation, the identity of a to their valuable collections ofopisthobranch specimen is seldom in question on~e familiarity photographs; to John and Mary Lavery, who with the species has been established. Many afforded us the opportunity to make collec species are similar in coloration or other char tions at Rongelap Atoll as guests aboard their acteristics, but a glance at living specimens yacht Si Ti Si III;and to Hans Bertsch, for his will usually readily differentiate among them substantial help with identification and taxo (see Rudman 1982a). nomic problems. This research was under It should be noted that intraspecific varia taken primarily at the Mid Pacific Research tion is also observed in radular characteristics, Laboratory, a facility sponsored by the U.S. both within and between geographic locations. Department of Energy through contract DE In many cases, conspecifics collected from AC08-76EV00703 with the University of Hawaii and Enewetak exhibit differences in Hawaii. details of radular morphology. Even within a single individual (e.g., Phestilla lugubris), tooth shape varies in teeth from different rows. Obviously , descriptions based solely on radu lar morphology are insufficient, parti~ularly LITERATURE CITED in genera within which the radulae of dlffere~t ABE, T. 1964. Opisthobranchia of Toyama species are similar, as they are, for example, III Bay and adjacent waters. Hokuro Ken, the genus Chromodoris. . Tokyo. 99 pp. A further problem is the oversynonynuza ALDER, J., and A. HANCOCK. 1864. Notice of tion of species, which can create problems in a collection of nudibranchiate Mollusca the determination of geographic distributions. made in India by Walter Elliot Esq., with For example, acceptance of Eliot's (1904b) descriptions of several new genera and statement that Chromodo ris tryoni and C. species. Trans. Zoo!' Soc. London 5: 113 aureopurpurea are conspecific would indic~te 147. that C. tryoni is distributed from East Afr!ca ALLAN, J. K .,1947. Nudibranchia from the to the Society Islands, instead of OCCUrrIng Clarence River Heads, North Coast, N. S. only in the Pacific, as we presently believe. W. Rec. Aust. Mus. 21 :432-463. Another example is Bergh's (1905) apparent - --. 1950. Australian Shells. Georgian synonymization of C. quadricolor and C. eli House, Melbourne. 470 pp. sabethina, which would make the former a BABA, K. 1930. Studies on Japanese nudi widespread Indo-west Pacific species rather branchs. 2. Venus 2 :43-50. than restricting it to East Africa and the Red --.1936. Opisthobranchia of the Ryukyu Sea as Rudman (1977) has done. (Okinawa) Islands. J. Dept. Agric. Kyushu Imper. Univ. 5(1): 1-50. ---. 1937a. Opisthobranchia of Japan. 2. J. Dept. Agric. Kyushu Imper. Univ . 5(7) : ACKNOWLEDGMENTS 289-344. We wish to thank the many people who ---. 1937b. Record of a nudibranch, have provided us with specimens , informa Gymnodoris striata (Eliot), from Amakusa, tion, and /or suggestions : Roger and Mary Japan. Zool. Mag. 49(6) :216-218. Marshall Islands Opisthobranchia-i-Jonnson AND BOUCHER 287

--- . 1938. Opisthobranchia ofKii, Middle der nudibranchiaten gastraopoden. ZooI. Japan. J. Dept. Agric. Kyushu Imper. Univ. Jahrb. Abt. Syst. 3(3): 348-364. 6(1) : 1-19. - --. 1889. Malacologische untersuchungen. ---. 1949. Opisthobranchia of Sagami In C. Semper, ed. Reisen im Archipel der Bay, collected by his majesty, the Emperor Philippinen 3(16) :815-872. ofJapan. Iwanami Shoten, Tokyo. 194 pp. --- . 1905. Die Opisthobranchia der Sibo ---.1952. Record ofan ascoglossan mol ga Expedition. Monographie 50 : 1-248. lusk, Oxynoe viridis (Pease) from Sagami BERTSCH, H. 1977. The Chromodoridinae Bay, Japan. Venus 17(2) :77-80. nudibranchs from the Pacific coast of --- .1953. Three new species and two new America. I. Investigative methods and records ofthe genus Glossodorisfrom Japan. supra-specific taxonomy. Veliger 20 : 107- . • PubI. Seto Mar. BioI. Lab. 3: 205-211. ll8. ---. 1955. Opisthobranchia of Sagami ---. 1980. A new species of Bornella from Bay, supplement. Iwanami Shoten, Tokyo. tropical west America (Mollusca, Opistho 59 pp. branchia). Spixiana 3(1): 33-42. - -- . 1957. The species of the genus Elysia BERTSCH, H., and S. JOHNSON. 1979a. Ha from Japan. PubI. Seto Mar. BioI. Lab. 6 : waiian chromodorids. West. Soc. Malac. 69-75. Ann. Rep. 12 : 12. - - - . 1960. The genera Okenia, Gonio --- . 1979b. Three new opisthobranch doridiella and Goniodoris from Japan records for the Hawaiian Islands. Veliger (Nudibranchia-Goniodorididae). PubI. Seto 22 :41-44. Mar. BioI. Lab. 8 :75-83. --- . 1981. Hawaiian Nudibranchs. Orien - --. 1969. Records of Learchis indica tal PubI. Co ., Honolulu. 112 pp. Bergh, 1896 from Japan and Hawaii. PubI. - -- . 1982. Three new species ofdorid nu Seto Mar. BioI. Lab. 16 : 399-403. dibranchs (Gastropoda: Opisthobranchia) BABA, K. , and l. HAMATANI. 1975. An illus from the Hawaiian Islands. Veliger 24 : trated list ofthe Phyllidiidae from Seto, Kii, 208-218. Middle Japan. Veliger 18 : 174-179. BOUCHER, L. M . In press. Extra-capsular yolk BEBBINGTON, A. 1974. Aplysiid species from bodies in the egg masses of some tropical east Africa with notes on the Indian Ocean Opisthobranchia. J. Molluscan Studies. Aplysiomorpha (Gastropoda, Opistho BROST, F. B., and R. D . COALE. 1971. A guide branchia). ZooI. J. Linn. Soc. 54: 63-99. to shell collecting in the Kwajalein Atoll. - --. 1977. Aplysiid species from eastern Charles E. Tuttle Co., Rutland, Vt. 157 pp. Australia with notes on the Pacific Ocean BURN, R . 1962. On the new pleurobranch Aplysiomorpha (Gastropoda, Opistho subfamily Berthellinae (Mollusca: Gastro branchia). Trans. Zool. Soc. London 34 : poda); a revision and new classification of 87-147. the species of New South Wales and Vic BEBBINGTON, A., and G . H . BROWN. 1975. toria. Mem. Natn. Mus. Melbourne 25 : Aplysia parvula Guilding in March, an 129-148. opisthobranch new to the British fauna. J. - --. 1966. Some Opisthobranchia from ConchoI. London 28 : 329-333. southern Queensland. J. Malac. Soc. Aus BERGH, R . 1877. Malacologische untersuch tralia 9 :96-109. ungen. In C. Semper, ed. Reisen im Archipel - -- . 1972. A guide to the Ascoglossa or der Philippinen 2(11) :429-494. sap sucking sea slugs of Australia. Aust. ---. 1878. Malacologische untersuchun Nat. Hist. 17(5): 174-178. gen. In C. Semper, ed. Reisen im Archipel - --. 1974. The taxonomy and distribution der Philippinen 2(13) :547-601. of Marianina rosea (Pruvot-Fol, 1930) and ---. 1879. Neue nacktschnecken der sud Thecacera pacifica (Bergh, 1883) comb. nov . see malacologische untersuchungen. 4. (Opisthobranchia: Dendronotacea and Journ. Mus. Godef. 5(14) : I-50. Doridacea). Veliger 16(3):305-306. --- . 1888. Die pleuroleuriden, eine familie - - -. 1975. A list ofthe dorid nudibranchs 288 PACIFIC SCIENCE, Volume 37, July 1983

ofAustralia (Gastropoda; Opisthobranchia). East Africa and Zanzibar. 3. Proc. Zoo!. J. Zoo!' London 176:514-517. Soc . London 1903(2): 354-385. BURN, R., andK. R. NARAYANAN. 1970. Tax --- . 1904b. On some nudibranchs from onomic notes on Eolis militaris Alder and East Africa and Zanzibar. 4. Proc. Zoo!' Hancock, 1864 (Opisthobranchia, Eoli Soc. London 1904(1): 380-406. dacea). J. Malac. Soc . Australia 12: 83-86. ---. 1904c. On some nudibranchs from CARLSON, C. H ., and P. J. HOFF. 1973. Exter East Africa and Zanzibar. 5. Proc. Zoo!. nal description of a living Aranucus bifidus Soc. London 1904(2) :83-105. (Odhner, 1936) (Opisthobranchia: Dendro - - -. 1905a. On some nudibranchs from notacea). Veliger 15:172-173. the Pacific, including a new genus, Chro ---. 1974. The Gastropteridae of Guam, modoridiella. Proc. Malac. Soc. London with descriptions offour new species (Opis 6(4) :229- 238. thobranchia: Cephalaspidea). Pub!. Seto - - -. 1905b. On some nudibranchs from Mar. Bio!. Lab. 11(5/6) :345-363. East Africa and Zanzibar. 6. Proc. Zoo!' ---. 1978. The identifiable Elysia from Soc. London 1904(2) :268-298. Guam (Elysiidae, Sacoglossa, Opistho - - - . 1906a. Nudibranchs and tectibranchs branchia). Micronesica 14:89-114. from the Indo-Pacific. 2. Notes on Lopho ---. 1981. Preliminary checklist of the cerus, Lobiger, Haminaea, and Newnesia. J. Opisthobranchia found on Guam. Univ. Concho!. London 11(10) :349-360. Guam Mar. Lab. Tech. Rep. 70:46-53. --- . 1906b. On the nudibranchs of south COLLINGWOOD, C. 1881. On some new species ern India and Ceylon, with special reference of nudibranchiate Mollusca from the east to the drawings by Kelaart and the col ern seas. Trans. Linn. Soc. London Zoo!' lections belonging to Alder and Hancock 2(2): 123-140. preserved in the Hancock Museum at New EALES, N . B. 1960. Revision of the world castle-on-Tyne. Proc. Zoo!. Soc. London species of Aplysia (Gastropoda, Opistho 1906 :636-691. branchia). Bull. British Mus. Nat. Hist. - -- . 1908. Reports on the marine biology (Zoo!.) 5 :269-404. of the Sudanese Red Sea. 11. Notes on a EDMUNDS, M. 1969. Opisthobranchiate Mol collection ofnudibranchs from the Red Sea. lusca from Tanzania. 2. Eolidacea (Cutho J. Linn. Soc. London Zoo!. 31:86-112. nidae, Piseinotecidae, and Facelinidae). --. 1909. Notes on a collection of nudi Proc. Malac. Soc. London 39 : 15-57. branchs from Ceylon. Spolia Zeylanica ---. 1971. Opisthobranchiate Mollusca 6(23): 79-95. from Tanzania (Suborder: Doridacea). ---. 1910. Nudibranchs collected by Mr. Zoo!' J. Linn. Soc. London 50 :339-396. Stanley Gardiner from the Indian Ocean in EKDALE, A. A. , S. J. HILLS, S. R. FISHER, and H. M. S. Sealark. Trans. Linn. Soc. London J. E. WARME. 1979. Quantitative ecology Zoo!' 13 :411 -438. and paleoecology of molluscan commu ---. 1913. Japanese nudibranchs. J. Col!. nities in reef associated, soft bottom Sci. Imper. Univ. T()kyo 35: 1-47. environments. Mid. Pac. Mar. Lab. Ann. Engel, H. 1942. The genus Dolabella. Zoo Rep. 1979: 28-33. Iogische Meded. 24; 197-239. ELIOT, C. N. E. 1900. Notes on tectibranchs ENGEL, H., and P. W. HUMMELINCK. 1936. and naked mollusks from Samoa. Proc. Ober westindische Aplysiidae und verwand Acad. Nat. Sci. Philadelphia 1899: 512-523. tenandere geberte. Capite. Zoo!' 8 : 1- 76. ---. 1903a. Notes on some new or little FARRAN, G. P. 1905. Report on the opistho known members of the family Doridiidae. branchiate Mollusca collected by Professor Proc. Malac. Soc. London 5 :331-337. Herdman, at Ceylon, in 1902. In W. A. --- . 1903b. On some nudibranchs from Herdman, Report to the Government of East Africa and Zanzibar. 2. Proc. Zoo!' Ceylon on the Pearl Oyster Fisheries of the Soc. London 1903(1): 250-257. Gulf of Manaar 3, supp!. report 21 :329 - --. 1904a. On some nudibranchs from 364.

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FIELDING, A. 1979. Hawaiian reefs and tide and little known species of Ceylon nudi pools. Oriental Publishing Co. , Honolulu. branchiate molluscs and zoophytes. J. Roy. 103 pp. Asiatic Soc. Ceylon Branch, Colombo 3 : GARRETT, A. G. 1873. Description of a new 84-139. species ofGoniodoris . Proc. Acad. Nat. Sci. ---. 1858b. Description of a new Cey Philadelphia 1873:232. lonese nudibranch. Ann. and Mag. Nat. - --. 1879. Description of a new species Hist. (3) 1(4):257-258. of Goniobranchus. Proc. Acad. Nat. Sci. MACNAE, W. 1954. On four sacoglossan mol Philadelphia 1879:31. lusks new to South Africa. Ann. Natal GOHAR, H. A. F., and I. A. ABOUL-ELA. 1957. Mus. 13:51 -64. On a new nudibranch "Phyllodesmium ---. 1958. The families Polyceridae and xeniae" from the Red Sea with a descrip Goniodorididae (Mollusca, Nudibranchia tion of its development. Publ. Mar. BioI. ta) in southern Africa. Trans. Roy. Soc. Sta . Al-Ghardaqa 9 : 131-144. South Africa 35 :341-372. - -. 1959. On the biology and develop MARCUS, E. 1956. Notes on Opisthobranchia. ment of three nudibranchs (from the Red BioI. Inst. Oceanografico Sao Paolo 7(1): Sea). Publ, Mar. BioI. Sta . Al-Ghardaqa 31-78. 10:41 -62. --- . 1965. Some Opisthobranchia from GOSLINER, T. M . 1980. The systematics of the Micronesia. Malacologia 3 :263-286. Aeolidacea (Nudibranchia: Mollusca) of MARCUS, E. D. B.-R. 1976. On Kentrodoris the Hawaiian Islands, with descriptions of and Jorunna (Gastropoda, Opistho two new species. Pac. Sci. 33 :37-77. branchia). Bolm. Zooi. Univ. Sao Paolo GOULD, A. A. 1852. Mollusca and shells. In I : 11-68. United States Exploring Expedition during MARCUS, E., and J. B. BURCH. 1965. Marine the Years 1839-1842, under the Command euthyneuran gastropods from Eniwetok of Charles Wilkes, U. S. N. Vol. 12. pp. Atoll, western Pacific. Malacologia 3: 235 1-510. -262. HAMATANI, I. 1980. On the species of the MARCUS, E., and E. D. B.-R. Marcus. 1970. genus Oxynoe Rafinesque, 1819 from Japan, Opisthobranch mollusks from the southern inclusive ofa new species (Opisthobranchia: tropical Pacific. Pac. Sci. 24 : 155-179. Ascoglossa). Publ. Seto Mar. BioI. Lab. MILLER, M. C. 1969. The habits and habitats 15(5/6) :349-360. ofthe opisthobranch molluscs ofthe British JOHANNES, R. E. 1963. A poison-secreting Solomon Islands. Phil. Trans. Roy. Soc. nudibranch (Mollusca: Opisthobranchia). Ser. B 255 :541-548. Veliger 5(3) : 104-105. ---. 1974. Aeolid nudibranchs (Gastro JOHNSON, S. 1982. Living seashells. Oriental poda: Opisthobranchia) ofthe family Glau Publishing Co ., Honolulu. 117 pp. cidae from New Zealand waters. Zool, J. ---. 1983. Distribution oftwo nudibranch Linn. Soc. 54 :31-61. species on a subtidal reef on the western ODHNER, N. 1936. Nudibranchia Dendrono shore of Oahu, Hawaii. Veliger 25 :356 tacea. A revision of the system. Melanges 364. Paul Pelseneer. Mem. Mus. Roy. d'Hist. JOHNSON, S., and H . BERTSCH. 1979. A popu Nat. Belg., ser II , Fasc. 3: 1057-1128. lation study ofthe nudibranch Chromodoris P EASE, W. H . 1860. Descriptions ofnew species godeffroyana. West. Soc. Malac. Ann. Rep. of Mollusca from the Sandwich Islands. II :8. Proc. Zool. Soc. London 1860: 18-36. KAY, E. A. 1979. Hawaiian marine shells. B. ---. 1861. Descriptions of new species of P. Bishop Mus. Spec . Publ, 64(4). 653 pp. Mollusca from the Pacific islands. Proc. KAY, E. A., and D. K. YOUNG. 1969. The Zoo!' Soc. London 1861 :242- 247. Doridacea (Opisthobranchia; Mollusca) of - --. 1871. Descriptions ofnudibranchiate the Hawaiian Islands. Pac. Sci. 23 : I72-231. Mollusca, inhabiting Polynesia. Am . J. KELAART, E. F. 1858a. Description of new Conchol. 6 :299-305. 290 PACIFIC SCIENCE, Volume 37, July 1983

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invertebrates. Medde!. fra Komm. for Dan branchia (Gastropoda: Opisthobranchia: marks Fiskeri- eg Havund., serie: Plankton Nudibranchia) from the central and west 4: 1-523. central Pacific with a description of a new WINCKWORTH, H. C. 1946. Glossodoris from species. Veliger 10: 159-173. Bombay. Proc. Malac. Soc. London 26(6) : - - -. 1969. Okadaia elegans, a tube-boring 155-160. mollusc from the central and west Pacific. YOUNG, D. K. 1967. New records of Nudi- Am . Zoo!' 9: 903-907.