Pacific Science (1979), vol. 33, no. I © 1980 by The University Press of . All rights reserved

The Systematics of the Aeolidacea (Nudibranchia: ) of the , with Descriptions of Two New !

TERRENCE M. GOSLINER 2

ABSTRACT: Nineteen species of aeolid nudibranchs are here recorded from the Hawaiian Islands. The natural history and distribution of each species is described. Morphological accounts of these taxa are provided and their sys­ tematic status is discussed. The status of an additional three species, previously recorded from the Hawaiian Islands but not encountered in this study, is reviewed. Based on the study of the Hawaiian aeolidacean fauna, the systematics of the are reviewed. An examination of the zoo­ geographical affinities of the Hawaiian aeolid fauna suggests that many taxa have a typical Indo-West Pacific distribution. However, a significant portion of the fauna is substantially more widely distributed and appears to be circum­ tropical.

THE FIRST SERIOUS ATTEMPTS to characterize the islands of and and span a the Hawaiian opisthobranch fauna are those period ofover 10 years. My own 0 bservations of Kay (1961, 1962a, 1962b, 1964a, 1964b) were made from September 1972 to October and Kay and Young (1969). Kay described 1973. During this period, monthly collections most of the anaspideans and several saco­ were made at four principal sites on Oahu: glossans, and Kay and Young dealt with the Kaneohe Bay, , dorid nudibranchs. Aeolid nudibranchs have Diamond Head Beach Park, and Kewalo only sporadically been described or recorded Basin. Collections were also made at other from Hawaii. Pease (1860) described two localities in the islands, including Fleming's species, Aeolis semidecora and A. parvula. Beach, , and on the Kona coast, Bergh (1900) described Samla annuligera Hawaii. from Laysan Island. Edmondson (1946) At least 22 species of aeolid nudibranchs recorded five species of aeolids, but identified can be attributed to the Hawaiian Islands. only Glaucus by name. Zahl (1959) figured Three species have been recorded that were semperi and Aeolidiella sp. from not encountered in this study: Samla annu­ Kaneohe Bay, Oahu. Harris (1968, 1970) ligera Bergh (known only from the holotype recorded melanobrachia and P. collected at Laysan); Embletonia gracile sibogae from the Hawaiian Islands. Rosin Risbec 1928 [here tentatively identified from (1969) recorded sp., and Baba a figure by Edmondson (1946)]; and Aeolis (1969) recorded indica. parvula Pease 1860 (which is not identifiable The purpose of this study is to revise the from Pease's brief description). At least five earlier work on the Hawaiian aeolids and to additional species of aeolids have been col­ report on previously unrecorded species. In lected by Kay but are not included in this this synthesis I include the collection records study because few specimens are available. of E. Alison Kay which are primarily from

I Manuscript accepted 15 May 1977. 2 University of Hawaii, Department of Zoology, , Hawaii 96822. Present address: University of New Hampshire, Department of Zoology, Durham, New Hampshire 03824. 38 PACIFIC SCIENCE, Volume 33, January 1979

Ghiselin (1965) have suggested that the gical variation and number of species. Two Aeolidacea are polyphyletic, but the evidence major groups exist, the facelinids with cuspi­ is not conclusive as the more primitive mem­ date radular teeth and the aeolidids with bers of the suborder, Notaeloidia and the pectinate radular teeth. The Coryphellidae, have been poorly studied. appear to be monophyletic, since members Odhner (1939) proposed the system of are similar in morphology and habits. Most classification whereby the aeolids are divided species are known to feed on sea anemones. into three tribes, the Pleuroprocta, Acleio­ The facelinids include a wide range of procta, and Cleioprocta, on the basis of anal morphological forms. They are the subject of position. This system is not entirely satisfac­ a great deal of taxonomic confusion and are tory, because there are numerous examples in need of revision. The controversy falls of genera that differ in anal position from into two schools of thought. One (Ernst other members of the family. has Marcus 1958, Miller 1974) contends that the the anus intermediate to the pleuroproct and major distinction is between the favorinids, acleioproct position, whereas the remainder with the grouped in arches, and the of the Coryphellidae are typically pleuro­ facelinids, with the cerata arranged in rows. proctic. Selva, a cuthonid, has the anus in the The other point of view is that of Edmunds cleioproct position, but others of the Cu­ (1970), who states that reproductive charac­ thonidae are acleioproctic (Edmunds 1964). ters, particularly the development of the One specimen of perea (Ernst receptaculum seminis and bursa copulatrix, Marcus, 1958) in this study was cleioproctic, should take taxonomic precedence over cera­ while seven other specimens were acleio­ tal arrangement, and that ceratal arrange­ proctic. Basing aeolid classification solely on ment should be considered polyphyletic.! anal position produces a polyphyletic, and agree with Edmund's contention that a serial hence artificial, classification system. receptaculum seminis is more primitive than The pleuroprocts are considered primitive, a semiserial receptaculum. and most members (except Roller, Miller (1974) contends that Babakina Rol­ 1973) have a triseriate radula. The families ler, 1973 is not closely allied to the Cory­ within the tribe are not easily distinguished, phellidae but with the glaucids and represents and various workers subdivide the tribe into a subfamily in the Glaucidae. His argument one to five families. is primarily based on the similarity of the Like the pleuroprocts, the more primitive uniseriate radula and oral glands. Miller acleioprocts, the Eubranchidae, and possibly suggests that the two bursae in Babakina are Cuthona (Ernst Marcus 1961), are distin­ not significant in their taxonomic placement guished by a triseriate radula. The more since some coryphellids and glaucids share advanced aeolids have a uniseriate radula, as this characteristic. do most cuthonids, Fiona, and the Piseino­ I contend that it is not so much the fact that tecidae. Most acleioprocts have a simply two bursae are present that is significant, but rounded head and anterior margin of the that their structure and form are distinctive. foot. The reproductive system is distinctive The fact that Antionetta and Dietata both within the Cuthonidae, where there is a distal have two bursae and are unquestionably receptaculum seminis near the gonopore and glaucids, is not supportive of Babakina's frequently a penial gland. The acleioprocts placement in the Glaucidae as stated by seemingly have polyphyletic origins within Miller (1974). While they have advanced the Aeolidacea (Edmunds 1970). This is glaucid characteristics of cleioproctic anus, particularly evident in the cuthonids and well-developed liver system, and general body Piseinotecidae, which are similar in external form of a glaucid, they also have a serial form but have markedly different radulae and receptaculum. It is unlikely that Babakina reproductive systems. would maintain its primitive coryphellid The cleioprocts represent the most diverse characters of pleuroproctic anus, primitive tribe of aeolids in both degree of morpholo- digestive system, prominent notal brim, Aeolidacea of the Hawaiian Islands-GosLINER 39

and club-shaped , and be more of his Facelininae in -but advanced reproductively, with a semiserial recognizing several genera within his Favor­ receptaculum, when Antionetta and Dictata ininae-is inconsistent, because characteris­ retain a serial receptaculum. It seems much tics of the penis are different in both groups. more reasonable that Babakina is allied to While the distinctions of genera are not the Coryphellidae. The are identical precise, the maintenance of genera seems to members of the Coryphellidae except for appropriate because it stresses differences their lack of lateral rows of teeth. Miller among the groups rather than combining maintains that the oral glands of Babakina them because the differences are difficult to are glaucid. Oral glands have not been studied discern. to any degree in aeolids and cannot be deemed Miller also stated that differences in what systematically significant at present. he considers Phidiana are based entirely on Miller (1974) did not make clear whether the structure of the penis. The statement is the rhinophores in Babakina caprinsulensis erroneous. Facelinella does not have an are joined in a common insertion. Roller ejaculatory vas deferens, while does. (1972) questioned the taxonomic level at Palisa differs in having papillate rhinophores which joined rhinophores can be considered and a serial receptaculum, and its significance significant, and this question remains largely in discerning the Facelinidae is well known unsettled. (Edmunds 1970). Other genera (i.e., Hermis­ Miller suggested that the genera Glaucus senda, Emarcusia, etc.) differ significantly in and Pteraeolidia are closely allied to the radular and external form. Phidiana is unique facelinids. I feel that, while they are obviously in that members of the genus lack tentacular allied on the basis ofreproductive and radular foot corners. On the above bases, I suggest form, they are significantly aberrant in their that the genera placed in synonomy under external form and ecology as to warrant Phidiana by Miller be retained as distinct familial status and that the Facelinidae, genera until more accurate relationships can Glaucidae, and Pteraeolididae, should be be assessed. maintained as familial taxa. Edmunds (1970) argued that the Favor­ inidae and Facelinidae are polyphyletic and SYSTEMATIC ACCOUNT that separation of the two families on the basis of branching of the digestive system Family Coryphellidae does not accurately reflect evolution within Flabellina annuligera (Bergh, 1900) the groups. Miller's (1974) placement of the Samla annuligera Bergh, 1900; Flabellina Favorininae, Crateninae, Facelininae, and annuligera (Marcus & Marcus 1967) Herviellinae as subfamilies within the Glau­ cidae does not solve the problem. They are This species was described from a single still polyphyletic units established on the specimen from Laysan Atoll (Bergh 1900), basis of convergence in ceratal arrangement. and has not since been recognized. Bergh Establishment of phyletic units on the basis separated Samla from Flabellina on the basis of reproductive system seems more accurate of its rounded anterior margin of the foot and decreases the number of polyphyletic compared to the tentacular foot corners of taxa. I suggest that the subfamilies proposed Flabellina. Marcus and Marcus (1967) re­ by Miller be discarded and that the species ferred to this species as Flabellina annuligera be retained in the single large family, the without stating any reasons. However, the Facelinidae. generic separation of Samla and Flabellina Miller (1974) combined all the genera of solely on the basis of pedal termination his Facelininae, except Antionetta, in the should not be maintained. genus Phidiana. While the generic distinctions Abbott (1974) misinterpreted Marcus and within the family are not entirely clear, Marcus (1967) and erroneously reported this Miller's method of placing all but a single species from Puget Sound, Washington. 40 PACIFIC SCIENCE, Volume 33, January 1979

DESCRIPTION: slender (Figure I), 7 to 15 mm in length. Body color purple or white. Rhinophores cream-yellow distally, red-brown at base, bulbous in form, and perfoliate. Cerata white with an orange subapical ring. Oral tentacles elongate, more than half the body length. Foot corners short, sharply recurved, tentacular. Cerata pedunculate in groups of two or three, with four to eight pairs of peduncles. Gonopore directly ventral to anteriormost group of cerata. Anus intermediate between pleuro­ proctic and acleioproctic position, dorsola­ teral, immediately anterior to second ceratal group. Radular formula 14-18 x 1.1.1 in three specimens. Rachidian teeth (Figure 2A) 55 }lm in length, sharply arched with seven or eight denticles on each side of central }lm FIGURE I. Flabel/ina alisonae. Length, 7 mm. cusp. Laterals (Figure 2B) triangular, 28 in length with four to six denticles. Jaws (Figure 2C) thin, 0.42 to 0.46 mm long, masti­ Flabellina alisonae spec. nov. catory border denticulate with four rows of ornata (Baba 1936); Flabel­ denticles. Thirty-five denticles in outermost lina ornata (Baba 1955, Abe 1964, row of cutting edge. Penis unarmed. Repro­ Marcus and Marcus 1967, Schmekel ductive system (Figure 2D) with saccate 1970) ampulla, proximal receptaculum seminis, distal bursa copulatrix, and nodular pro­ TYPE LOCALITY: Diamond Head Beach state. Park, Oahu, Hawaii TYPE MATERIAL: The holotype is deposited DISCUSSION: Baba (1936) identified an in the Bernice P. , Honolulu, aeolid from Okinawa as Coryphella ornata Hawaii (BPBM 218720). A paratype radula has Risbec, 1928, stating that his specimen had been deposited in the invertebrate zoology several rows of denticles on the jaws while collection of the California Academy of Risbec's contained only one. Subsequently, Sciences, San Francisco, California (CASIZ Baba (1955) transferred the species to Flabel­ paratype 00714 and type slide series 532). lina because ofthe pedunculate cerata present in his material from Japan and Okinawa. DISTRIBUTION: Southern Japan (Abe 1964, Marcus and Marcus (1967), in a discussion of Baba 1955), Okinawa (Baba 1936) the generic distinctions between Coryphella HABITS AND OCCURRENCE: Flabellina ali­ and Flabellina, noted that the cerata in Baba's sonae was collected in the vicinity of various material are clearly pedunculate while in hydroids, but no direct relationship can be Risbec's description the cerata do not appear suggested. One egg mass was found on an to be pedunculate. unidentified ectoproct: Eleven specimens In a review of the genus Flabellina, have been found in tidepools at Poipu, Kauai; Schmekel (1970) noted the differences in on patch reefs in Kaneohe Bay, Oahu; at ceratal arrangement between the Risbec Diamond Head, Oahu, in less than 1 m of animals and those ofBaba and suggested that water, and from a depth of 2 m, Honaunau, there was a high probability they represent Hawaii. This species has also been found at two species. Schmekel also noted differences Enewetak, Marshall Islands (Kay, personal in dentition of the lateral teeth. communication). In comparing the shape of the radular

= i adl;;; Wi::: - Aeolidacea of the Hawaiian Islands-GOSLINER 41

A B

I .1mm I D m.g.

O.t. b.c.

me.g. o.g.

.4mm

FIGURE 2. Flabellina alisonae; A, rachidian tooth; B, lateral tooth; C, jaw; D, reproductive system (a.g., albumen gland; am, ampulla; b.c., bursa copulatrix; m.g., mucous gland; me.g., membrane gland; o.t., ovotestis; pr, prostate; r.s., receptaculum seminis).

teeth of Baba's and Risbec's specimens, I be examined. Within the description of C. note that the rachidian of the New Caledon­ ornata, Risbec indicated the existence of ian examples has a prominent central cusp another individual described as C. ornata var. while the specimens from Japan and Okinawa violacea. The radular teeth and coloration have a central cusp that is not significantly differ markedly from the typical form. In my larger than the adjacent denticles. The later­ opinion, these differences warrant specific als also differ significantly. In Coryphella separation, and the name Coryphella violacea ornata Risbec, the lateral is quadrate with comb. nov. is suggested. denticles only on the upper half of the tooth; The specimens described in this study are in Baba's material, the teeth are triangular virtually identical to Baba's animals from with denticles only along the basal two-thirds Okinawa and Japan. The coloration in all ofthe inner margin. There are two clusters of cases is white, but with a purple tinge in some cerata anterior to the anus in the New of the Hawaiian specimens. The cerata are Caledonian animals while in the Japanese and uniformly spindle-shaped with a subapical Okinawan animals there is a single preanal orange band. In the Hawaiian material, as in ceratal peduncle. There seems to be little Baba's specimens, the cerata are arranged in doubt that the animals described by Baba peduncles of one, two, or three cerata with represent a species distinct from C. ornata one preanal peduncle per side. The extremely Risbec. Because there is no evidence to elongate oral tentacles are characteristic of suggest that C. ornata Risbec has pedunculate all the material. The shape of the jaws from cerata, it is preferable to maintain the species the Okinawan animal is close to that of the within Coryphella until further material can three Hawaiian specimens examined. The TABLE I COMPARATIVE DATA ON Flabellina AND Two SPECIES OF Coryphel/a

NUMBER OF DENTICLES RIGHT ON EACH ANTERIOR NUMBER OF SIDE OF NUMBER OF DIGESTIVE RADULAR RACHIDIAN DENTICLES MASTICATORY SYSTEM REPRODUCTIVE SPECIES LOCALE COLOR ROWS CUSP ON LATERAL BORDER CERATA SYSTEM

Flabellina affinis Mediterranean Lilac 34 6-7 6 - Compound Receptaculum (Gmelin, 1791) peduncles only F. alisonae sp. Hawaii, Japan, Purple-white 14-20 7-12 4-7 Short Simple, Receptaculum nov. Okinawa cerata with I preanal and bursa orange dng row F. annuligera Laysan Atoll Blue, white 20 8-10 5-7 Elongate Simple, Receptaculum (Bergh, 1900) cerata with 1 preanal only orange ring row F. babai Naples Blue-white 18-24 5-10 5-8 Short Simple, Receptaculum Schmekel, 1970 cerata with 2 preanal only orange ring rows F. engeli Cura9ao Pink with 20 7-11 5-10 Unknown Simple, Receptaculum Marcus & brown cerata I preanal only Marcus, 1968 with white row F. macassarana Macassar Pink-yellow 17 5 4-5 Unknown Simple, Unknown Bergh, 1905 2 preanal rows(?) F. stohleri Gulf of Orange with 14 6-8 9 Unknown Simple, Unknown Bertsch & California white 3 preanal Ferreira, 1974 rows F. telja . Gulf of Blue cerata; 28 8-11 6-8 Elongate Simple, Receptaculum Marcus & California orange with 4 preanal and bursa Marcus, 1967 white tip rows Coryphel/a ornata New Caledonia White cerata 20 7 8 Elongate Compound, Unknown Risbec, 1928 with orange 2 preanal groups C. violacea comb. New Caledonia Rose-violet - 5 5 Unknown Unknown Unknown nov. 3 longitudinal red lines Aeolidacea of the Hawaiian Islands-GosLINER 43 jaws are broad and ovoid with a short Of the species that have only a receptacu­ masticatory border containing several rows lum seminis, (Omelin, 179 I) of denticles. The radulae bear a strong can be distinguished on the basis of its num­ resemblance, with 16 or 17 rows of teeth in erous radular rows (34 as compared to 14-20 the Japanese and Okinawan animals and 14 in F. alisonae), compound ceratal peduncles to 18 rows in theHawaiian material. Schmekel with two preanal peduncles, and coloration. (1970), apparently based on a specimen from Flabellina babai Schmekel, 1970 possesses Japan, described a radula with 20 rows. In two preanal ceratal peduncles and propor­ Baba's and Schmekel's animals, the rachidian tionately shorter oral tentacles than does F. tooth contains nine or ten denticles on each alisonae. Flabellina annuligera (Bergh, 1900) side of the central cusp compared to seven and F. engeli Marcus and Marcus, 1968 have or eight in the Hawaiian aeolids, while the a simply rounded anterior foot margin and laterals in their specimens have six to nine only a single row of denticles on an elongate denticles compared to four to seven in the masticatory border. present study. These differences are not con­ The present species is named in honor of sidered significant, particularly in view of the E. Alison Kay of the University of Hawaii, similarity in the shape of the teeth in all Honolulu, Hawaii. citations. Therefore, I consider the specimens described by Baba (1936) as Coryphella ornata and Flabellina ornata (Abe 1964, Baba Family Cuthonidae 1955, Schmekel, 1970) to be conspecific with Cuthona perea (Ernst Marcus, 1958) Flabellina alisonae spec. nov. Catriona perea Ernst Marcus, 1958 The only description of the reproductive (Edmunds 1964); Trinehesia perea Flabellina alisonae system of is that of Marcus & Hughes 1974; Cuthona reflexa Schmekel (1970). She noted that the penis is Miller, 1977 unarmed, as in Baba's (1955) and the present material, and that a distal bursa copulatrix DISTRIBUTION: Brazil (Ernst Marcus 1958); and proximal receptaculum seminis are both Jamaica (Edmunds 1964); Florida (Eveline present. Marcus 1972) ; Barbados (Marcus and Several characteristics of the described Hughes 1974); New Zealand (Miller 1977) species of Flabellina and those of Coryphella HABITS AD OCCURRENCE: Three specimens ornata and C. violaeea are summarized in were found associated with the sea anemone, Table I. Of the six species of Flabellina for Aiptaisia sp., on the floating docks at Coconut which the reproductive system has been Island, Kaneohe Bay, Oahu, in October 1972; described, only F. alisonae and F. telja Marcus four specimens in the thermal pollution & Marcus, 1967 are known to have both a tables at Coconut Island in July 1973; and proximal receptaculum and distal bursa. single specimens in August and September F. Flabellina telja is clearly separable from 1973 from the same locality. Several speci­ alisonae because it has more numerous rows mens have been found from St. Petersburg, of radular teeth (28 as compared to 14-20 Florida (Joseph L. Simon, University of in F. alisonae), and has four preanal ceratal South Florida, personal communication, rows compared to a single row in F. alisonae. 1975). The coloration is also strikingly different. Flabellina alisonae differs from F. stohleri DESCRIPTION: Animal (Figure 3A) 5 to 15 Bertsch & Ferreira, 1974 in its coloration and mm in length. Color translucent white with the fact that F. stohleri has three preanal scatterings of opaque white and dark olive ceratal rows. Flabellina maeassarana Bergh, green; orange tinge sometimes present on the 1905 differs from F. alisonae in its pink­ head and dorsal surface; olive green pigment yellow body color, more numerous cerata in ceratal core. Cerata often unusually elon­ (six cerata each within the first two ceratal gate. Animals typically cuthonid, with round­ rows, which are preanal), and rachidian ed head and smooth rhinophores. Cephalic teeth with a prominent central cusp. tentacles shorter than rhinophores and terete. 44 PACIFIC SCIENCE, Volume 33, January 1979

B

D

I 5mm

FIGURE 3. Cuthona perea: A, dorsal view of living animal; B, posterior radular tooth; C, anterior radular tooth; D, jaw; E, penial stylet.

Anterior margin of foot simply rounded. cus and Hughes 1974); and from New Zea­ Anterior digestive system of two rows of land (Miller 1977). The specimens from cerata: Left digestive system containing five Hawaii and St. Petersburg, Florida, agree to ten pairs of rows. Gonopore between first with the described variation of the species. two rows of cerata. Anus acleioproctic, The dark-green to black color in the cerata, cleioproctic in one specimen. Strongly taper­ with dark coloration on the head anterior to ing radula (Figure 3B, C) of 35 rows of teeth the rhinophores, is distinctive. All the Hawai­ in a 12-mm specimen. Posterior teeth smal­ ian animals had two subapical bands of lest, 38 pm wide, with four denticles on each opaque white on the cerata, but several side of central cusp. Anterior teeth 2.5 times specimens lacked orange pigment on the broader, up to 87 pm wide, with 11 denticles head and notum. The cerata of the larger on each side of central cusp. Jaws thin and Hawaiian animals were significantly more elongate (Figure 3D), 0.78 mm long. Mastica­ elongate than previously described material tory border with a row of 18 denticles in one from the Atlantic. The radular configuration specimen. Penis (Figure 3£) with accessory is similar to the range described (Edmunds gland and terminal stylet, 75 pm long. 1964, Ernst Marcus 1958, Miller 1977): 35 teeth in the present material compared to DISCUSSION: Cut/wna perea (Ernst Marcus, 16 to 33. The unusual tapering of the radula 1958) is known from the western Atlantic with marked differences in the width of the from Cananeia, Brazil (Ernst Marcus 1958); teeth and number of denticles from anterior Jamaica (Edmunds 1964); Miami, Florida to posterior is characteristic of this species in (Eveline Marcus 1972); the Barbados (Mar- all accounts. This tapering is especially Aeolidacea of the Hawaiian Islands-GosLiNER 45 evident in members ofthe Aeolidiidae, which A single egg mass was also found on the feed largely on sea anemones. Cuthona perea same hydroid colony. feeds exclusively on the sea anemone, Aipta­ DESCRIPTION: Single individual (Figure sia sp. in Hawaii. It is suggested that this radular shape represents a convergent adap­ 4A) 7 mm in length. Color translucent white tation in C. perea and the Aeolidiidae with opaque white pigment throughout body associated with feeding on sea anemones. surface. Rhinophores (Figure 4B) with seven The configuration of the digestive system to eight semicircular annulations. Length of rhinophores and oral tentacles equal. Foot in the Hawaiian specimens agrees with that described by Edmunds (1964), but at least simply rounded anteriorly. Cerata in seven rows, three rows in anterior right digestive two more rows were present in the posterior digestive system of the larger Hawaiian system. Cerata (Figure 4C) with two bulbous animals than in those described from Jamai­ swellings in distal portion. Opaque white ca. The presence of a penial stylet and penial pigment present at swellings. Gonopore accessory gland in the Hawaiian material between first two ceratal rows. Anus sub­ further substantiates the conspecific nature of dorsal, acleioproctic, anterior to fourth cera­ the organisms. Penial armature cannot be tal row. Radula uniseriate (Figure 4D), elongate, of 41 rows of teeth 45 Jim wide. used to separate Trinehesia from Cuthona, Central cusp elongate with five or six den­ and the species must be regarded as belonging ticles on each side. Jaws elongate (Figure 4£), to Cuthona (Williams and Gosliner, in press). 0.70 mm long. Masticatory border a single The coloration of Cuthona reflexa Miller, row of 28 denticles. Penis bulbous (Figure 1977 is similar to that described for C. perea 4F) with apical stylet at apex and penial in that it also possesses dark hepatic tissue in gland. the ducts of the digestive gland. The radular morphology, number ofteeth, and denticula­ DISCUSSION: In 1949 Baba described Cra­ tion of Cuthona reflexa lie within the range of tena pinnifera from Sagami Bay. Burn (1964) variation described for C. perea. Both taxa applied the name Trinehesia pinnifera in a have an armed penis with an accessory discussion of the rhinophores in Toorna, penial gland and a prostatic vas deferens. stating that T. pinnifera is the only Trinehesia The only morphological character which with annulate rhinophores. Again, penial differs is the number of ceratal rows in the armature cannot be used to separate Trin­ right anterior digestive system; there are two ehesia from Cuthona, and the species must be rows in C. perea and three to four rows in C. regarded as belonging to Cuthona (Williams reflexa. Owing to their extreme morpholo­ and Gosliner, in press). gical similarity, Cuthona reflexa Miller, 1977 The single specimen found in this study is is here regarded as a junior synonym of C. similar to those described by Baba from perea (Marcus, 1958). Sagami Bay, Japan, and is considered con­ specific with the Japanese animals. Baba's animals ranged in length from 7 to 10 mm, as Cuthona pinnifera (Baba, 1949) compared to 7 mm in the Hawaiian animal. pinnifera Baba, 1949; Trinehesia In Baba's animals, the cerata were arranged pinnifera (Burn 1964) in eight to nine rows, as compared to seven in the specimen from Kaneohe Bay. Both the DISTRIBUTION: Sagami Bay, Japan (Baba Japanese and the Hawaiian animals had 1949) three ceratal rows in the anterior right diges­ HABITS AND OCCURRENCE: A single speci­ tive system. In the Japanese specimens, the men was found under a dead coral head on gonopore was located ventral to the second a largely dead reef in Kaneohe Bay, Oahu, ceratal row; in the Hawaiian animal, it is in August 1973. It was associated with the between the first and second rows. The anus plumularid hydroid, Halopteris diaphana in animals from both localities is located (Heller, 1868), on which it presumably feeds. anteriorly, adjacent to the dorsal end of the 46 PACIFIC SCIENCE, Volume 33, January 1979

.lmm I--l

1mm 1 sl.

v.d. E

.5mm ~

.lmm I---l

FIGURE 4. Cuthona pinnif'era: A, dorsal view of living animal; B, ; C, ceras; D, radular tooth; E, jaw; F, penis (p.g., penial gland; st., stylet; v.d., vas deferens). fourth ceratal group, in a typical acleioproct ring below the opaque white apex. The position. In animals from both localities, the Hawaiian specimen lacks yellow pigment. foot corners are simply rounded. Baba described five to eight semicircular The coloration of Baba's animals was a annulations on the rhinophores as compared uniform white, but some specimens had a to seven or eight in the Hawaiian animal. yellow ring around the middle of the cephalic Baba did not mention the two bulbous tentacles and cerata with a subapical yellow swellings seen in the cerata of the Hawaiian

II _ .,milS;;;;W: us i JAi k Z1iii$l Aeolidacea of the Hawaiian Islands-GosLINER 47

individual, nor did he figure or describe the long with large expanded masticatory border general shape of the jaws. The Japanese bearing 18 denticles along its length. Penis animals had 20 to 22 denticles on the mastica­ armed with a curved stylet (Figure 5E) tory border of the jaws as compared to 28 in 110 ,um long, with penial gland. Receptacu­ the animal from Kaneohe Bay. lum seminis distal, opening near female The radula in the material from Japan gonopore. consisted of a uniseriate radula of 45 rows of teeth with six to seven denticles on each side DISCUSSION: Cuthona diminutiva reaches a of the central cusp. This compares with a maximum length of 5 mm. Animals main­ radula of 41 rows with five denticles on each tained in the laboratory and fed on corynid side of the central cusp in the animal from hydroids never exceeded this size. The specific Hawaii. epithet diminutiva was chosen because of the small size of the animals. Cuthona diminutiva resembles Trinchesia Cuthona diminutiva spec. nov. boma Edmunds, 1970 and Trinchesia sp. TYPE LOCALITY: The seawater tables of the (Edmunds 1970) in several aspects of mor­ thermal pollution study at the Hawaii Insti­ phology, but there are significant differences tute of Marine Biology (Coconut Island), that warrant specific separation. Trinchesia Oahu, Hawaii, serve as the type locality. is considered a junior synonym of Cuthona (Williams and Gosliner, in press). Cuthona TYPE MATERIAL: The holotype is deposited boma, C. diminutiva, and Cuthona sp. are all in the Bernice P. Bishop Museum, Honolulu, sexually mature when only 3 mm in length Hawaii, where it bears the number BPBM and have few cerata (three to five rows, with 218719. Paratype material resides in the one to four cerata per row and two preanal author's personal collection. rows). Each of the three species possesses a HABITS AND OCCURRENCE: Cuthona diminu­ penial stylet; all three are similar in repro­ tiva was abundant in the thermal pollution ductive morphology. The radular teeth are study tables at Coconut Island from July to similar in number of rows and shape. In C. August 1973, where it apparently feeds on diminutiva and C. boma the cerata are elon­ an unidentified corynid hydroid and is fre­ gate, but in C. diminutiva there is a prom­ quently found with its egg masses. inent bulbous swelling bear the middle of the ceras. In C. boma the cerata begin near DESCRIPTION: Animals small (Figure 5A), the base of the rhinophores, whereas in C. 3 to 5 mm in length. Animals sexually mature, diminutiva the cerata begin in the posterior laying small, semicircular egg masses 3 to 5 two-thirds of the animal's length. Cuthona mm long. Coloration generally white with boma has seven to nine denticles on each encrustations of white and brown pigment. side of the central cusp of the rachidian, Eyes readily visible. Rhinophores smooth while C. diminutiva possesses five or six. The and terete. Oral tentacles approximately penial stylet of C. boma is straight, while it equal to rhinophores in length and diameter. is curved in C. diminutiva. Cuthona boma and Anterior foot corners simply rounded. Cer'l­ C. diminutiva also differ in coloration: there tal formula (after Edmunds 1970) 4, 2; 1, I is orange pigment on the dorsum and rhino­ and 2, 3; I, I in two specimens. Cerata phores of C. boma that is absent in C. (Figure 5B) one to four per row with bulbous diminutiva, and C. diminutiva has uniformly inflation about midway of their length. distributed brown flecks that are absent in Gonopore ventral to most anterior ceratal C. boma. row. Anus acleioproctic in posterior half of Cuthona sp. (Edmunds 1970) differs from body. Radula (Figure 5C) consisting of 24 to C. diminutiva in having fewer, shorter cerata, 34 teeth 35 ,um wide with five or six denticles a straight penial stylet, and orange pigment on each side ofthe slightly prominent central on the dorsum. cusp. Jaws elongate (Figure 5D), 0.48 mm The following species also have two preanal 48 PACIFIC SCIENCE, Volume 33, January 1979

1mm 1 5mm 1· I

D

,1OO~m I 40~m

FIGURE 5. Cuthona diminutiva: A, dorsal view of living animal; B, ceras; C, radular tooth; D,jaw; E, penial stylet. ceratal rows: Cuthona bylgia (Bergh, 1870); diminutiva. Ernst Marcus (1958) listed most C. colmani (Burn, 1961); C. cuculatta (Bergh, species that can currently be considered as 1905); C. ornata (Baba, 1937); C. pupillae members of the genus Cuthona. Schmekel (Baba, 1961); C. rubrata (Edmunds, 1970); (1968) summarized the species described after and C. taita (Edmunds, 1970). All these have Marcus' work. Cuthona diminutiva differs significantly more numerous cerata than C. from all other species listed. Only C. pusilla Aeolidacea of the Hawaiian Islands-GosLiNER 49

(Bergh, 1898) is similar to C. diminutiva, in seventh ceratal row. Reproductive system that the anus is located in the posterior half (Figure 6D) typically cuthonid. Penis with of the body. short, 40 flm long stylet (Figure 6C) and penial gland. PhestilIa sibogae Bergh, 1905 DISCUSSION: PhestilIa sibogae was de­ PhestilIa sibogae Bergh, 1905, (Bonar scribed from Roti Island in the Timor Sea and Hadfield 1974, Hadfield and Karlson (Bergh 1905). The animals were described 1969, Harris 1970, Kawaguti 1943); from preserved material; the color in the Aeolidiella edmondsoni Ostergaard, 1955 living state was not described. The radula was variable, consisting of 24 to 44 teeth DISTRIBUTION: Roti Island, Timor Sea (Bergh 1905); Palau (Kawaguti 1943); Ha­ with six to seven denticles on each side of the waii (Bonar and Hadfield 1974, Hadfield and central cusp. Karlson 1969, Harris 1970, 1975, Ostergaard Kawaguti (1943) recorded PhestilIa sibogae from Palau and cited its association with the 1955). coral genus Porites. In 1955, Ostergaard HABITS AND OCCURRENCE: PhestilIa sibo­ described Aeolidiella edmondsoni from Ha­ gae is apparently quite uncommon in the field. waii on the basis of external characteristics. Only one specimen was seen during the course Hadfield and Karlson (1969) reported that of this study. Kay (personal communication) the larvae in the laboratory are induced to reports only two specimens in more than 10 metamorphose in the presence ofmucus from years of field collection. The single specimen living Porites. Bonar and Hadfield (1974) in this study was found under a dead chunk described the development of the species. of the coral Porites compressa in Kaneohe Harris (1970) stated that Ostergaard's Bay, Oahu. A single egg mass was found on species is a junior synonym of P. sibogae but the underside ofa living Porites lobata colony gave no specific basis for this contention. in . Under laboratory condi­ However, it is obvious from Ostergaard's tions, where a head of Porites is in a water plate of the animal that they are conspecific. table and unfiltered seawater is allowed to pass over it, the species flourishes (Hadfield and Karlson 1969). In these colonies, Phes­ Phestilla melanobrachia Bergh, 1874 tilIa feeds voraciously upon the polyps, and DISTRIBUTION: (Bergh 1874, egg masses are deposited abundantly on the Harris 1968), Sagami Bay, Japan; Okinawa, undersurface of the coral. The coloration of Singapore, Hawaii (all Harris 1968) the cerata in Phestilla is largely dependent upon the color ofthe Porites colony on which HABITS AND OCCURRENCE: The occurrence it has been feeding. and feeding habits of Phestilla melanobrachia are described by Harris (1968). During the DESCRIPTION: Color variable, green, yel­ present study two specimens were collected low-green, or purple, with opaque white on from the ahermatypic coral, Tubastrea aurea, head and cerata. Head and anterior margin from a depth of 2 to 5 m in Kaneohe Bay, of foot rounded. Rhinophores smooth and Oahu, in December 1972. conical. Cerata nodular with numerous swel­ lings, arranged in pedunculate rows, six in DESCRIPTION: Length 10 to 16 mm. Color anterior right digestive system, eight rows in orange, with areas of translucent white in posterior digestive system. Radula (Figure oral region. Rhinophores smooth, conical. 6A) uniseriate, 21 to 31 teeth, 100 p.ffi wide, Oral tentacles short. Anterior margin of foot with five to nine denticles on each side of rounded. Cerata thin, elongate, in peduncu­ prominent central cusp. Jaws (Figure 6B) late rows: six in anterior right system, six in strong, 2.2 to 4.0 mm long, with smooth posterior system. Gonopore ventral to third masticatory border. Gonopore ventral to ceratal row. Anus acleioproctic anteriodorsal third ceratal row. Anus anteriodorsal to to seventh ceratal row. Uniseriate radula 50 PACIFIC SCIENCE, Volume 33, January 1979

c

bOp m --1 r-

v.d.

FIGURE 6. Phestilla sibogae: A, radular tooth; B, jaw; C, penial stylet; D, reproductive system (a.g., albumen gland; am, ampulla; m.g., mucous gland; me.g., membrane gland; o.t., ovotestis; p.g., penial gland; LS., recepta­ culum seminis; v.d., vas deferens). (Figure 7A) consisting of 71 teeth, 160 .urn brachia and P. sibogae in the present study wide with eight to ten denticles on each side both have an armed penis alters the generic of prominent central cusp. Jaws (Figure 7B) diagnosis, since Bergh (1874, 1905) described elongate, 4.0 mm long. Penis (Figure 7C) the penis as unarmed. with penial gland and armed with short cuti­ cular stylet. Reproductive system similar Embletonia gracile Risbec, 1928 to P. sibogae. Embletonia gracile was described from DISCUSSION: Harris (1968, 1970, 1975) two specimens collected in Canala Bay, New observed many of the biological aspects of Caledonia. The animals were white, the this species in several Indo-Pacific localities. cerata arranged in seven rows, and digitate He noted that variation in color of Phestilla at their apices. Baba (1959) recorded E. melanobrachia is dependent upon its food gracile from Sagami Bay, noting that the source. Most recently he described the life terminal branchial papillae (cerata) distin­ history ofthe species. The fact that P. melano- guish E. gracile from the type of the genus, Aeolidacea of the Hawaiian Islands-GosLiNER 51

A

100 pm lmm

c

I 40pm

FIGURE 7. Phestilla melanobrachia: A, radular tooth; B,jaw; C, penial stylet.

E. pulchra (Alder & Hancock, 1844) from specimen of this species off Beach England, E.faurei Labbe, 1923 from France, at a depth of25 m. and E. evelinae Marcus, 1957 from Brazil. A subspecies, E. gracile paucipapillata, is distinguished from the typical form in having Family Facelinidae four or five rows ofcerata per side, with vivid Facelinella semidecora (Pease, 1860) orange-yellow digestive diverticula, as op­ Aeolis semidecora Pease, 1860 (Pruvot­ posed to seven or eight ceratal rows with Fo11947); Coryphella semidecora (Bergh pale-yellow diverticula. This subspecies was 1890); Facelinella anulifera Baba, 1949 described by Baba and Hamatani (1963). (1965) The Hawaiian record is based on a short description and a figure by Edmondson DISTRIBUTION: Hawaii (Pease 1860) and Sagami Bay, Japan (Baba 1949, 1965) (1946, fig. 98b). The presence ofapical ceratal papillae and the distinctive body shape in HABITS AND OCCURRENCE: This aeolid is Embletonia strongly suggest that the animals commonly found on the undersurface of figured by Edmondson are conspecific with rocks in the intertidal to depths not exceeding E. gracile. Dr. Hans Bertsch (personal com­ 5 m. It is common throughout the year. The munication) has recently found a 15 mm species is widely distributed within the Ha- 52 PACIFIC SCIENCE, Volume 33, January 1979

~ 30jJm 1200 /Jm

c

m.g.

r. s. I 20", m

~ 2mm

FIGURE 8. Facelinella semidecora: A, radular tooth; B, jaw; C, reproductive system (a.g., albumen gland; am, ampulla; m.g., mucous gland; o.t., ovotestis; r.s., receptaculum seminis); D, penial spines.

USiJiJ ibil Ii ilia - Api

Aeolidacea of the Hawaiian Islands-GosLiNER 53

waiian Islands, found at Poipu, Kauai; Ala 1860). The color pattern and exter~al. mor­ Moana Beach Park, , Diamond phology are within the r~~ge of va.natlOn of Head Beach Park, and Kaneohe Bay, all t>n the material from Hawall. The rhmophores Oahu; Fleming's Beach, Maui; and Honau­ were annulated with four or five rings, and nau, Hawaii. Egg masses have been observed the cerata were arranged in eight to twelve throughout the year. rows with five to six rows in the right anterior digestive system. DESCRIPTION: Animals 6 to 12 mm in tra~slu­ The radula of Baba's specimens and the length. Body thin, elongate: Color ~awai­ cent white, with opaque white maculatlOns; present material is also similar. The oral tentacles banded with opaque white and ian specimens have 17 to 20 teeth With five to eight denticles on each side of the central red, with single band of each near center ~nd fi~e a long band of white at apex; apical portIOn cusp; there are 16 rows with three to of rhinophores with large band of opaque denticles on each side of the central cusp m sh~pe. white; varying amounts of red-orange often the Japanese animals. The general of the teeth is the same, despite the vanatlOn present around head and medi~lly;.cerata brown internally with opaque white pigment in the number of denticles. Baba (1965) men­ concentrated at apices. Rhinophores conical tioned a row of spines on the peculiarly with two to five slight annulations. Oral shaped, nonejaculatory penis, and stated tentacles sharply tapered, one-third length that the nonejaculatory vas deferens sepa­ rates Facelinella from Facelina. of rhinophores. Anterior foot corners tenta­ The similarity in reproductive morphology cular. Cerata short, five rows in anterior of the present material to that of F. quadri­ digestive system and seven rows in posterior lineata (Baba, 1930) and F. anulifera furt~er system. Ceratal rows of posterior digestive suggests the affinity between the matenal system in two distinct groups. Gonop<;>re from Japan and Hawaii (Baba 1965). ventral to posterior portion of right antenor digestive system. Anus anterior to second ceratal row of posterior digestive system. militaris (Alder & Hancock, 1864) Radula (Figure 8A) consisting of 17 t~ 20 Eolis militaris Alder & Hancock, 1864 teeth, 60 to 70 ,urn wide, with five to eight (Eliot 1906); Hervia militaris Farran denticles on each side of central cusp. Jaws 1905 (Narayanan 1969, O'Donoghue (Figure 8B) 0.92 mm long, with masticatory 1932, Satyamurthi 1952); Lew·chis indica border of 15 denticles. Penis (Figure 8C) Bergh, 1896 (Baba 1969); I-!ervia da~­ paddle-shaped, lacking an ejaculatory vas geri Risbec, 1953; Learchls howensls deferens. Lateral edge of penis with single Burn, 1966; Caloria militaris (Burn & row of hook-shaped spines (Figure 8D). Narayanan 1970); ~lon!Sate Ampulla saccate, entering <;>viduct. (Miller 1974) Receptaculum seminis semi senal, With duct entering oviduct near junction with albumen DISTRIBUTION: (Alder and Hancock gland. Female gland mass large. 1864, Burn and Narayanan 1970, Eliot 1906, Narayanan 1969, O'Donoghue 1932, Satya­ DISCUSSION: Aeolis semidecora was de­ murthi 1952); Ceylon (Farran 1905); Aus­ scribed from the Hawaiian Islands (Pease Naraya~an 1860). Pease's description closely matches tralia (Burn 1966, Burn and the specimens found in the present study, 1970); middle Japan (Baba 1969); Amboma (Bergh 1896); New Caledonia (Risbec 1953); and there is little doubt that the present Funafuti (Burn and Narayanan 1970); Ha­ material is conspecific. waii (Baba 1969, Burn and Narayanan 1970); Facelinella anulifera was described from and New Zealand (Miller 1974) Sagami Bay, Japan (Baba 1949). Similarity in color pattern, radular structure, and. re­ HABITS AND OCCURRENCE: Caloria militaris productive morphology lead me to consider feeds on colonies of the hydroid Halocordyle it a junior synonym of F. semidecora (Pease (= Pennaria) tiarella (McCrady, 1857). This 54 PACIFIC SCIENCE, Volume 33, January 1979

I- 60 IJ m

I 300 IJm

FIGURE 9. Cataria militaris: A, radular tooth; B, jaw. species is commonly encountered in Kaneohe followed by five to seven rows in posterior Bay, Oahu, on patch reefs, and also on the system. Gonopore ventral to first and second floating docks of Coconut Island, through­ ceratal rows. Anus posterior to first row of out the year. Halacardyle colonies on the posterior digestive system. Radula (Figure leeward coast of Oahu apparently support 9A) uniseriate, 20 rows ofteeth, 100 f..I.m wide, smaller populations of Calaria. Single speci­ with five to twelve denticles on each side of mens have been seen at Ala Moana Beach the more elongate central cusp. Jaws rounded Park and from 12 m off Pokai Bay (Kay, (Figure 9B), thickened anteriorly, 1.4 mm personal communication). long. Masticatory border a single row of 20 denticles. Reproductive system as described DESCRIPTION: Length 7 to 35 mm (from by Baba (1969). 50 animals). Body long and slender. Vividly and variably colored: body color translucent DISCUSSION: Burn and Narayanan (1970) yellow-white, head bright red-orange with presented an extensive synonomy under opaque white lines, oral tentacles opaque Calaria militaris, contending that this taxon white for distal three-fourths, rhinophores cannot be included in Facetina (=Hervia) translucent white with medial band ofopaque because of the presence of an unarmed penis white and apical band of red, cerata red and smooth rhinophores, and that it should from base to middle, followed by band of be placed in Learchis. They stated that their slate blue, terminated by white apex. Rhino­ specimens are identical with the junior syn­ phores conical, smooth. Oral tentacles long, onym, Learchis indica Bergh, 1896, but gave sharply acute. Foot corners tentacular, elon­ no basis for this statement. The type of Lear­ gate. Cerata arranged in seven to nine rows chis, L. indica, was distinguished from Calaria with two rows in anterior digestive system maculata, the type of Calaria, only on the Aeolidacea of the Hawaiian Islands-GosLINER 55

basis of jaw shape. Burn and Narayanan tables of the thermal pollution study at considered these two genera synonymous, Coconut Island, Kaneohe Bay, Oahu. with Caloria having priority. Miller (1974) contended that Caloria is a synonym of Phi­ DESCRIPTION: Length 7 to 35 mm. Body diana. My reasoning for separating Phidiana color translucent white with varying amounts and Caloria was explained earlier (see dis­ of opaque white and orange-brown, as in cussion of Facelinidae.) anterior half of oral tentacles; cerata trans­ In their synonomy Burn and Narayanan parent white with opaque white flecks and (1970) studied specimens from Lord Howe apices, often with orange and blue pigment; Island; Funafuti; Kaneohe Bay, Oahu; and ceratal core chocolate brown. Rhinophores the Gulf of Kutch, India. They indicated no smooth, translucent with opaque white and variation. Miller (1974) compared material orange. Body typically facelinid, with elon­ from New Zealand with that from other gate oral tentacles more than twice the length localities. The general body shape is the same of rhinophores. Foot corners tentacular, in each description. Burn (1966) described sharply recurved. Cerata elongate, thin, Learchis howensis as having six to eight an­ arranged in four to eight horseshoe-shaped nulae on the rhinophores but did not mention arches. Cerata become erect when disturbed the fact when he characterized Caloria mili­ and are readily detachable. Gonopore ventral taris as having smooth rhinophores. Annulate to anterior portion of first horseshoe. Anus rhinophores must, therefore, be within the within second horseshoe and nephroproct variation of the species but appear to be anterior to first arch of posterior digestive uncommon. Miller (1974) mentioned papil­ system. Radula (Figure lOA, B) consisting lations on the head in a few specimens from of 24, 26, 27 teeth in three specimens, 65 to New Zealand. 130 ,um wide with three or four large denticles The coloration of Caloria militaris is pre­ on each side of central cusp. Jaws (Figure dictable. Most workers cite red-orange pig­ 10C) 1.8 to 2.5 mm in length with 20 denticles ment arranged in a V shape on the head or along masticatory border. Penis (Figure lOD) other orange coloration over a translucent armed with curved spine 230 ,um long. Am­ cream to yellow body. The cerata are brick pulla narrow, elongate, convoluted. Vas red to brown with blue and white pigment. deferens thick, prostatic throughout. Recep­ There may be as many as nine pairs of rows taculum seminis semiserial. of cerata. DISCUSSION: Barnard (1927) described The jaws are always described as strong Hervia quadricolor from False Bay (near with a denticulate masticatory border con­ Cape of Good Hope), South Africa. The taining 9 to 25 denticles on the margin. The radula and jaws were described but not illus­ radula has 14 to 25 teeth with five to eight trated. The anatomy of the reproductive denticles on each side of the prominent system was not discussed. Barnard was quite central cusp. Both the radula and jaws are specific as to the pattern of coloration on consistent in shape. the cerata, describing it as orange, ultra­ The penis is unarmed in all citations. marine, dark brown, and sulfur yellow. MacNae (1954) described and de­ signated Hervia quadricolor as the type spe­ (Barnard, 1927) cies. He noted that the coloration was highly Hervia quadricolor Barnard, 1927; Aeo­ variable and that one or more of the bands lidiella sp. (Zahl 1959); Godiva quadri­ ofcolor may be reduced or wanting. MacNae color (MacNae 1954) illustrated the radula, jaws, elaboration of the digestive diverticula, and the reproductive DISTRIBUTION: South Africa (Barnard system, thus providing a detailed account 1927, MacNae 1954) and Hawaii (ZahI1959) ofthe morphology ofthe species. HABITS AND OCCURRENCE: This species was The material from Hawaii closely ap­ found throughout the year in the seawater proaches that from South Africa in color and 56 PACIFIC SCIENCE, Volume 33, January 1979

100 IJm

0.1.

1500pm I

FIGURE 10. Godiva quadricolor: A, radular tooth, dorsal view; E, radular tooth, lateral view; C,jaw; D, repro­ ductive system (am, ampulla; fgm, female gland mass; 0.1., ovotestis; ph, penial hook; r.s., receptaculum seminis). morphology. The specimen photographed contained 24 to 27 teeth as compared to 26 by Zahl (1959) especially resembles the Afri­ to 30 teeth in the African animals. The teeth can specimens in coloration. Other Hawaiian in all accounts have three or four large den­ specimens vary in color. tides on each side of the central cusp. The The radula of the Hawaiian specimens masticatory border of the jaw had slightly Aeolidacea of the Hawaiian Islands-GosLiNER 57

fewer denticles in the Hawaiian animals than with dense encrustations of opaque white; previously described for the species, but cerata light brown, pink, white, or green with this variation is considered within the range opaque white apices. Rhinophores conical of intraspecific variation. with two prominent bulbous swellings, one The reproductive system, as described by near the middle, the other just below the MacNae (1954), is virtually identical with apex. Oral tentacles elongate and sharply that of the present material. Specimens from pointed. Anterior foot corners elongate and both localities have an elongate ampulla; tentacular. Cerata in five pairs of horseshoe­ short, thick, prostatic vas deferens; and shaped arches. Single arch in anterior diges­ terminal penial spine. The only difference tive system and four arches in posterior noted was in the receptaculum seminis, where system. Cerata papillate and slightly recurved it was directed anteriorly lying on the top of toward central body axis. Gonopore ventral the prostate in the Hawaiian animal and to cerata of first horseshoe. Anus within was directed posteriorly and rested on the second horseshoe. Radula (Figure 11 A) uni­ mucous gland in the African specimens. seriate, consisting of 22 teeth, 140 pm long. MacNae (1954) included Rizzolia australis Teeth with single cusp, lacking lateral den­ Bergh, 1884 in Godiva, and subsequently, ticles. Jaw (Figure liB) elongate, 1.1 mm Edmunds (1964) described G. rubrolineata long, with three to five rows of denticles on from Jamaica. Both species lack a terminal masticatory border. penial spine and have a simple, nonglandular penis. They more closely approach Setoeolis DISCUSSION: japonicus was de­ Baba 1965 and are here regarded as members scribed by Baba (1949) based on three speci­ of this genus. mens from Sagami Bay, Japan. Baba and Hamatani (1964) redescribed the species in greater detail and tentatively placed it in Favorinus japonicus Baba, 1949 synonomy with Eolis nodulosa Keelart, 1859 Favorinus japonicus Baba, 1949 (Abe and Favorinus gouaroi (Risbec, 1928). Both 1964, Baba and Hamatani 1964, Carlson ofthese species seem distinct from the present and Hoff 1973) species. Eolis nodulosa is figured (Eliot 1906) DISTRIBUTION: Middle and southern Japan with smooth cerata and bulbous portions on (Abe 1964, Baba 1949, Baba and Hamatani the rhinophores. Because several species of 1964), Guam (Carlson and Hoff 1973) Favorinus have bulbous swellings on the rhinophores [F. branchialis (Rathke, 1806), HABITS AND OCCURRENCE: Favorinus japo­ F. auritulus Marcus, 1955, and F. ghanensis nicus has been found at Ala Moana Beach Edmunds, 1968], the presence of rhinophoral Park, along the seawall at Kewalo Basin, in swellings cannot be regarded as sufficient the seawater tables of the Pacific Biomedical evidence to imply that F. japonicus and E. Research Center at Kewalo Basin, all on nodulosa are conspecific. While F. gouaroi Oahu; and at Poipu, Kauai. The animals (Risbec, 1928) resembles F. japonicus in occur throughout the year as do their egg external features, there are six denticles on masses. each side of the central cusp of the radula These aeolids feed on the egg masses of teeth. Also, F. gouaroi lacks swellings on the various opisthobranchs, such as Aplysia rhinophores (Risbec 1928, pI. 12, fig. 13). juliana, A. dactylomela, Dolabrifera dolabri­ Favorinus branchialis, F. auritulus, and F. fera, M elibe pilosa, and Pleurobranchus ghanensis all have denticles adjacent to the semperi. median cusp ofthe radular teeth. DESCRIPTION: Length 8 to 17 mm. Color The Japanese specimens of F. japonicus, translucent white to green; oral tentacles those from Guam (Carlson and Hoff 1973), translucent white from middle to apex; foot and those from Hawaii are similar in form and rhinophores translucent white; remain­ and coloration. Baba (1949) described F. der of body, except cerata, translucent white japonicus as yellow with opaque white encru- 58 PACIFIC SCIENCE, Volume 33, January 1979

I AO ~m AOO~m

FIGURE 11. Favorinusjaponicus: A, radular tooth; B, jaw. stations and red cerata. The specimens from feeds on opisthobranch eggs as well as Guam were described as transparent with hydroids. white masking, but Carlson and Hoff (1973) stated that the color varies with feeding Herviella mietta Marcus & Burch, 1965 habits. Baba (1949) described the cerata as Herviella mietta Marcus & Burch, 1965; arranged in rows, whereas the Hawaiian Herviella sp. nov. Baba, Rosin 1969 animals had ceratal horseshoes. However, the cerata often appear as double rows rather DISTRIBUTION: Enewetak Atoll, Marshall than arches. The position of the gonopore Islands (Marcus and Burch 1965); Hawaii and anus are identical in animals from Japan (Rosin 1969) and Hawaii. HABITS AND OCCURRENCE: Two specimens Internally, Baba's animals and the present were found in this study, one from the north material are even more similar. The uni­ side of Coconut Island and a second from seriate radula from the Japanese specimens Portlock, Koko Head, Oahu. Kay (personal consisted of 17 to 20 teeth without any lateral communication) reports specimens from denticles, as compared to 22 teeth in a Ha­ Black Point and Kewalo Basin, Oahu. Rosin waiian specimen. The jaws in the Japanese (1969) reported specimens from Kewalo specimens contained five or six rows of den­ Basin, Oahu, where they reportedly feed ticles as compared to three to five in the upon Anthopleura nigrescens. Kay (personal Hawaiian material. communication) has also found this species Neither Baba (1949) nor Carlson and Hoff at Fanning Atoll, Line Islands. (1973) mentioned the feeding habits in their animals, but Haefelfinger (1969) stated that DESCRIPTION: Length 7 to 40 mm. Body F. branchialis [as F. alba, synonymized by colors translucent white and dark green to Lemche (1964)] feeds on the egg masses of black; dark pigment most pronounced on opisthobranchs and that its color is depen­ head; area around eyes and rhinophores dent on the eggs upon which it feeds. Thomp­ lacking dark pigment and appearing as two son and Brown (1976) noted that F. blianus translucent white patches; foot translucent Aeolidacea of the Hawaiian Islands-GosLiNER 59

I- lmm

lOOl.lm

FIGURE 12. Herviella miella: A, radular tooth; B,jaw. white. Rhinophores approximating oral ten­ cusp (Marcus and Burch 1965). In specimens tacles in length, smooth and conical. Both from Hawaii, there are 22 to 37 rows with six short relative to rest of body. Anterior por­ to eight denticles on each side of the central tion of foot simply rounded. Cerata long and cusp. The denticles in the Hawaiian animals thin, arranged in five or six rows, with a single are all approximately equal in length, while row in right anterior digestive system. Gono­ in the animals from Enewetak, the outer pore ventral to first row of cerata. Anus denticles are more elongate than the inner posterior to second ceratal row. Uniseriate and central denticles. radula (Figure 12A) consisting of 23 to 37 The jaws in specimens from both localities teeth, 150 to 200 fl-m wide with six to eight are similar in shape, but are somewhat denticles on each side of much thickened broader in specimens from Enewetak. The central cusp. Jaws (Figure 12B) elongate, 2.0 specimens from Enewetak had 18 denticles to 4.0 mm long, with ten irregularly spaced on the masticatory border as compared to denticulations. 10 in a Hawaiian animal. The type specimen had a penial stylet DISCUSSION: Herviella mietta was described (Marcus and Burch 1965). from Enewetak, Marshall Islands (Marcus and Burch 1965). The prey-predator rela­ tionship between Herviella sp. and the sea Noumeaella sp. cf. rehderi Ernst Marcus, anemone Anthopleura nigrescens was de­ 1965 scribed by Rosin (1969). The similar external HABITS AND OCCURRENCE: A single speci­ description between the specimens found by men of Noumeaella sp. was found in 2 m of Rosin and those found by Marcus and Burch, water at Keauhou Bay, Hawaii, on 3 Sep­ and the identical habitat between Rosin's tember 1973. and the present material, suggest that they are all conspecific. DESCRIPTION: Body slender and elongate, The radulae of the type specimens were 11 mm. Color translucent white with dense uniseriate, consisting of 18 teeth with eight patches of encrusting opaque white pigment or nine denticles on each side of the central arranged randomly. Rhinophores short and 60 PACIFIC SCIENCE, Volume 33, January 1979

I 20 jJm

200jJm I

FIGURE 13. Noumeaella sp. cf. rehderi: A, radular tooth; E, jaw. thick, anteriorly smooth, and densely papil­ denticles on the masticatory border as com­ late posteriorly. Oral tentacles approximating pared to 28 in the Hawaiian animal. rhinophores in length and tapering to acute The radulae demonstrate more substantial apex. Foot corners tentacular. Five pairs of similarity. The animal from Palau had 18 ceratal groupings arranged in horseshoe­ rows ofteeth with six or seven lateral denticles shaped arches, inserted commonly at base. on each side of the central cusp as compared Single arch in anterior digestive system. to 19 teeth with nine or ten denticles on each Cerata long, fusiform, and recurved. Gono­ side. The overall shape of the teeth is similar pore within first ceratal arch, anus within in the two animals. second. Uniseriate radula (Figure 13A) 40,um While it appears that the specimen de­ wide, consisting of 19 rows with nine or ten scribed by Ernst Marcus (1965) bears sub­ denticles on each side of the prominent me­ stantial similarity to the specimen from dian cusp. Jaws (Figure 13B) generally round, Hawaii, the lack ofmaterial in both instances 0.70 mm long. Masticatory border with 28 makes more positive identification difficult. denticles. DISCUSSION: The description of Noumea­ ella rehderi was based on a single specimen Family Pteraeolididae collected in Palau (Ernst Marcus 1965). The specimen was 3.5 mm long and was described (Angas, 1864) as colorless (apparently meaning translucent Flabellina Ianthina Angas, 1964; F. sem­ white rather than transparent and clear). peri Bergh, 1870; F. scalopendrella Ris­ The animal from Hawaii is similar in many bec, 1928; Pteraeolidia semperi Bergh, respects to N. rehderi. Both are translucent 1875 (Edmunds 1970, Eliot 1903, 1904a, white, although the Hawaiian animal has Ernst Marcus 1965, Ernst and Eveline opaque white pigment as well. The jaws are Marcus 1960, Risbec 1953, 1956); P. similar in shape in both specimens; however, Ianthina (Burn, 1965, Ernst and Eveline the jaws in Marcus' animal contained 40 Marcus 1970) Aeolidacea of the Hawaiian IsIands-GosLINER 61

1.75 mm

FIGURE 14. Pteraeolidia ianthina: A, radular tooth; B, jaw.

DISTRIBUTION: Red Sea (Ernst and Eveline form, as the generic name implies. Gonopore Marcus 1960); Zanzibar (Eliot 1903, 19040); below posterior portion of single ceratal Viet Nam (Risbec 1956); (Angas group. Anus within first arch of posterior 1864, Burn 1965); New Caledonia (Risbec digestive system. Uniseriate radula (Figure 1928, 1953); middle Japan (Baba 1949); 14A) consisting of 36 rows, 190 /lm wide Philippines (Bergh 1870); Micronesia (Ernst with five to seven denticles on each side of Marcus 1965); and Hawaii (Zahl 1959) the central cusp. Central cusp may have one to three small denticles. Jaws (Figure 14B) HABITS AND OCCURRENCE: A single speci­ 3.0 to 4.2 mm long, with many dense rows men was collected from each of three locali­ of small papillate denticles on masticatory ties: 10 to 20 m from Kaneohe Bay, Oahu, border. in February 1973; 20 m off Waikiki, Oahu, in March 1973; and from 10m, off Molokini, DISCUSSION: This species was described off the shore of Maui in November 1972. as Flabellina ianthina from Australia (Angas Specimens from the Kona coast of Hawaii 1864). Bergh (1870) described Flabel/ina found in August 1971 (Kay, personal commu­ semperi from the Philippines and in 1875 nication) were associated with the octocoral named it as the type species of Pteraeolidia. Sarcothelia edmondsoni. Many other workers have recorded this species from various localities. Burn (1965) DESCRIPTION: Animals large, reaching and Ernst and Eveline Marcus (1970) placed 80 mm. Color exceedingly variable: body Angas' F. ianthina in Pteraeolidia and as­ color generally green; green cephalic tentacles serted that P. semperi is a junior synonym banded with opaque white and black pig­ of P. ianthina, but they provided no basis ments; dorsal surface with gold reticulations for their decisions. or opaque white patches; cerata green or Baba (1949) recorded P. semperi from purple, sometimes with opaque white pig­ Japan, and described yellowish animals with ment, with a yellow or gold apex. Rhino­ blue cerata. The radula consisted of 20 rows phores perfoliate throughout. Oral tentacles of teeth with 10 to 20 denticulations on each smooth with acute apex. Anterior foot cor­ side of the central cusp, as compared to 35 ners tentacular. Body long and slender with rows with five to seven denticles in a Hawaiian 5 to 18 pairs of ceratal groups, winglike in specimen. Risbec (1953) stated that the num- 62 PACIFIC SCIENCE, Volume 33, January 1979

500IJm

50IJ m

FIGURE 15. G/aucus at/anticus: A, radular tooth; B, jaw. ber of rows of teeth and denticles per tooth colors dark blue-purple and silver. Rhino­ in material from New Caledonia is exceed­ phores and oral tentacles reduced. Anterior ingly variable. The jaws in Baba's .and Ris­ margin of foot simply rounded. Cerata in bec's accounts of P. semperi resemble those two to three groups on both sides of the of the Hawaiian specimens in shape and body, most anterior group largest. In a 30 mm denticulation of the masticatory border. specimen, 16 cerata in first group, II in second, and 6 in third. In a 7 mm specimen, uniseriate radula (Figure 15A) consisting of Family Glaucidae 16 teeth 62 J.Lm wide with five denticles on Gfaucus atfanticus Forster, 1777 each side of central cusp. Jaws (Figure 15B) Hirudinia marina Breyn, 1705; Gfaucus 1.0 mm long, sharply angular, with straight, atfanticus Forster, 1777 (Miller 1974, elongate masticatory edge bearing 130 den­ Thompson and McFarlane 1967); ticles. Penis armed with dark-brown penial radiata Gmelin, 1790; Scyffaea marga­ spine. ritacea Bosc, 1802; G. hexapterygius DISCUSSION: Glaucus atlanticus and its Cuvier, 1808; G. octopterygius Cuvier, many synonyms are known from throughout 1808; Laniogeras blanvilli Goldfuss, tropical seas. The genus Glaucus Forster, 1820; L. effortii Blainville, 1825; G. 1777 is separated from Gfauciffa Bergh, 1868 forsteri Lamarck, 1836; G. !inectus by the presence of a penial spine (Thompson Bergh, 1868; G. gracilis Bergh, 1868; and McFarlane 1967). Thompson and Mc­ G. fongicirrus Bergh, 1868 (Edmondson Farlane (1967) emphasize the high degree of 1946); G. marinus (Baba 1949) variability found in specimens of G. atlan­ DISTRIBUTION: Circumtropical ticus. The coloration is especially variable, but is basically silvery-white and blue. HABITS AND OCCURRENCE: Pelagic, feeds Thompson and McFarlane (1967) state on siphonophores and chondrophores. Two that the radula contained 18 to 20 teeth, specimens were collected off , with two to ten denticles on each side of the Oahu, in plankton hauls in December 1972. median cusp. The specimen from Hawaii, DESCRIPTION: Length, 7 to 30 mm. Body in which the radula was observed, contained Aeolidacea of the Hawaiian Islands-GosLiNER 63

60 IJm

lmm

FIGURE 16. Aeolidiella takanosimiensis: A, radular tooth; B,jaw.

16 teeth with five denticles on each side of brown; generally with a subapical white the median cusp. Neither Thompson and band on cerata; oral tentacles and rhino­ McFarlane nor Baba (1949) reported the phores translucent white or orange; speci­ number of denticulations on the masticatory mens from thermal pollution study tables border. lack orange. Rhinophores smooth and coni­ cal. Oral tentacles elongate. Anterior foot Family Aeolidiidae corners short and tentacular. Foot long and tapered posteriorly. Cerata with four to eight Aeolidiella takanosimiensis Baba, 1930 rows in right anterior digestive system in Aeolidiella takanosimiensis Baba, 1930, single group, with II to 16 in posterior system 1949 (Sphon 1971) in groups of one to four rows. Gonopore ventral to anteriormost ceratal group. Anus DISTRIBUTION: Middle Japan (Baba 1930, in first ceratal group of posterior digestive 1949); southern California (Sphon 1971) system. Uniseriate radula (Figure 16A) con­ HABITS AND OCCURRENCE: Four specimens sisting of 15 to 17 teeth, 150 to 200 J1.m wide, were collected from Kewalo Basin seawall, with 16 to 19 denticles on each side ofcentral Oahu, in April 1973; three specimens from cusp. Jaws (Figure 16B) rounded, 1.1 to the thermal pollution study tables, Coconut 2.3 mm long, with smooth masticatory Island, Kaneohe Bay, Oahu, in August 1973; border. a single specimen from Portlock, Koko DISCUSSION: Aeolidiella takanosimiensis Head, Oahu, in June 1973; and three speci­ was described from middle Japan (Baba mens from Poipu, Kauai, in January 1965 1930). The Japanese animals are identical (Kay, personal communication). Eggs were in coloration with the specimens found at found throughout the year. Kay (personal Kewalo Basin and at Portlock, Oahu (Baba communication) has found this species at 1949, pI. 49, fig. 167). Baba does not mention Fanning Atoll, Line Islands. a wide range in color, as was noted in the DESCRIPTION: Length 9 to 21 mm. Animals Hawaiian animals. translucent white with varying amounts of The radulae of the Japanese animals con­ orange in head region, often with opaque sisted of 16 teeth with 24 to 34 denticles on white spots posterior to rhinophores, encir­ each side of the median cusp as compared cled by red-orange; cerata brick red to dark to 15 to 17 teeth with 16 to 19 denticles in a 64 PACIFIC SCIENCE, Volume 33, January 1979

Hawaiian animal. The teeth are much more translucent white with dark gray-brown liver rounded in the Japanese animals. The jaws diverticula; remainder of animal translucent of the specimens from both localities are white. Rhinophores perfoliate with many virtually identical but slightly more rounded lamellae. Tips of oral tentacles expanded in the Hawaiian animals. as minute bulbs. Anterior foot corners short, Sphon (l971) recorded Aeolidiella takano­ tentacular appendages. Gonopore within simiensis from Los Angeles and San Diego, first ceratal arch. Anus within first arch of California. While Sphon gave no detailed posterior digestive system. Radula (Figure comparison of his animals, personal com­ 17A) uniseriate, with 29 broadly arched, munication with Sphon and my own obser­ pectinate teeth, 500 to 700 J.lm wide. Den­ vations of animals from San Diego confirm ticles: 37 to 53 on each side of central their identification. Some specimens from identation ofteeth. Jaws (Figure 17B) strong, California have a stouter body and lighter ovoid in shape, 4.5 mm long, with elongate, orange coloration than Japanese and Ha­ smooth masticatory border. waiian animals. DISCUSSION: is a well­ known aeolid, which is extremely variable. Spurilla neapolitana (Delle Chiaje, 1823) The large number of synonyms attests to Eolis neapolitana Delle Chiaje, 1823 ; its variability. Eolidina neapolitana Delle Chiaje, 1844; Spurilla neapolitana ranges in color from Eolis alderiana Deshayes, 1865; Flabel­ pink to orange, with brown and gray liver lina neapolitana A. Costa, 1866; F. inor­ diverticula. There mayor may not be opaque nata A. Costa, 1866; Spurilla neapolitana white pigment. The single specimen found (Bergh 1864, 1876, Edmunds 1964, Hae­ in Hawaii is within the described range of felfinger 1969, Eveline and Ernst Marcus color seen in this species. Haefelfinger (1969) 1959,1960,1962,1967,Pruvot-FoI1953, attributed the range of color in most nudi­ 1954, Trinchese 1878, Vayssiere 1888); branchs to variable feeding habits and cited S. sargassicola Bergh, 1861, 1877; S. S. neapolitana as an example. brasilana MacFarland, 1909; S. neapo­ litana var. brazilana Marcus, 1955, 1957; The cerata in all Spurilla neapolitana are S. mogribiana Pruvot-Fol, 1953; S. daka­ reportedly recurved at the apex, as in the Hawaiian specimen. The cerata are arranged rensis Pruvot-Fol, 1953 with a single arch in the anterior digestive DISTRIBUTION: Mediterranean (Bergh system followed by a variable number of 1864,1876,1877, A. Costa 1866, Delle Chiaje arches and rows in the posterior system. The 1823, 1844, Deshayes 1865, Haefelfinger single specimen from this study had a single 1969, Trinchese 1878, Vayssiere 1888); arch followed by three arches and two rows. Atlantic coast of France (Pruvot-Fol 1954); As in all material ofthe species, the gonopore Morocco, Senegal, Cape Verde Islands (all is located in the first arch of cerata, and Pruvot-Fol 1953); Sargasso Sea (Bergh the anus is in the first arch of the posterior 1861); Florida (Eveline and Ernst Marcus system. 1960, 1962, 1967); Texas (Eveline and Ernst In Italy (Trinchese 1878), S. neapolitana Marcus 1959); Jamaica (Edmunds 1964); Sao reaches a length of 70 mm, while in Brazil Paulo, Brazil (Ernst Marcus 1955, 1957) (Ernst Marcus 1955), it varies from 2 to 35 mm. HABITS AND OCCURRENCE: A single speci­ men of Spurilla neapolitana was found on The radular teeth vary in number and the undersurface of a rock at the seawall, denticulation. Previous workers report 11 to 28 teeth with 15 to 63 denticles on each side Kewalo Basin, Oahu, in November 1972. of the median cusp. The Hawaiian specimen DESCRIPTION: Length 22 mm. Head, has 29 teeth with 37 to 55 denticles on each rhinophores, oral tentacles, and pericardial side of the median notch. region with orange reticulate pigment; cerata The jaws in Spurilia neapolitana are vari- Aeolidacea of the Hawaiian Islands-GoSLlNER 65

B

200prn

FIGURE 17. Spuril/a neapolitana: A, radular tooth; B, jaw. able. Bergh (1877, 1882) described a denti­ Rhinophores short with enlarged bulbous culate masticatory border, while all other portion around middle. Head broad, with workers have described a smooth mastica­ red or orange pigment. Oral tentacles tory edge. The specimen from Hawaii has a prominently constricted at middle. Anterior smooth masticatory border. foot corners rounded. Cerata in 9 to II The occurrence of Spurilla neapolitana in groups, all curved toward central body axis. Hawaii represents the first record of the Gonopore within first group of cerata. Anus species in the Indo-Pacific and implies that in first group of cerata in posterior digestive the species is circumtropical in distribution. system. Radu1a (Figure 18A) uniseriate, consisting of 23 teeth, 210 to 230 Jim wide with 22 to 28 denticles on each side ofcentral Spurilla japonica (Eliot, 1913) cusp. Jaws (Figure 18B) thick, 1.3 to 2.3 mm Aeolidiella japonica Eliot, 1913 (Baba long, with smooth masticatory border. 1949); Spurilla japonica (Edmunds 1969) DISCUSSION: Burn (1969) differentiates DISTRIBUTION: Middle Japan (Baba 1949, Spurilla Bergh, 1864 from Aeolidiella Bergh, Eliot 1913) 1867 on the following bases: a more anterior anus in Aeolidiella, smooth rhinophores in Aeolidiella compared to perfoliate or annu­ HABITS AND OCCURRENCE: Collections of this species were made from Poipu, Kauai, late rhinophores in Spurilla, and difference June 1972; Kewa10 Basin, Oahu, April in jaw shape. Based on these differences, 1962 and January 1965; Kahuku, Oahu, Edmunds (1969) transferred Aeolidiella alba December 1962; Kaimalino, Oahu, August Risbec, 1928 and A. japonica Eliot, 1913 to 1963 and January 1966 (Kay, personal the genus Spurilla. communication). Spurillajaponica was described from Japan (Eliot 1913). Baba (1949) described living DESCRIPTION: Length 7 to 30 mm. Color material from Sagami Bay, Japan. The ex­ translucent white with opaque white on ternal body form in Baba's animals is similar rhinophores and as maculations on cerata, to the material from Hawaii, but the colora­ with solid white apex; cerata olive green. tion differs substantially. The Hawaiian 66 PACIFIC SCIENCE, Volume 33, January 1979

100 IJm

lmm

FIGURE 18. Spuri/lajaponica: A, radular tooth; B,jaw.

specimens lack the dark, reticulate pigment DISTRIBUTION: Mediterranean (Haefel- on the head evident in Baba's specimens. finger arid Stamm 1959); Bonaire (Eveline Eliot described the radula of Spurilla and Ernst Marcus 1970); Baja California japonica as having 18 teeth with as many as (Bertsch 1972) 65 denticles on each side of the central cusp. Baba described 16 to 30 teeth with 30 to 60 HABITS AND OCCURRENCE: A single speci­ denticles on each side of the central cusp. men was found on the surface of sand flats The material from Hawaii has 23 teeth with in Kaneohe Bay, Oahu, in I m of water on 22 to 28 denticles on each side of the central 15 September 1973. cusp. DESCRIPTION: Single specimen (Figure The jaws of specimens described by Eliot 19A, B) 26 mm in length. Body color brown, and Baba and those from Hawaii all have reticulated with translucent white; oral ten­ smooth masticatory borders. tacles, rhinophores, and foot brown with Edmunds (1969) described the affinities white maculations; dorsal surface of body between Spurilla alba and S. japonica, but with series of 14 ocellae of blue-green, en­ stated that they differ in color, size, and circled by yellow and with three to nine large number of digestive gland ramifications. The spots of opaque white; cerata yellow-brown bulbous swellings of S. japonica also appear reticulated with opaque white, and with a to be more pronounced than in Edmunds' subapical band of white and translucent figure of S. alba (1969, fig. 9A). white apex. Rhinophores densely papillate on posterior side; apex sharply acute. Oral Baeolidia nodosa (Haefelfinger & Stamm, tentacles elongate. Anterior foot corners 1959) tentacular, with deep groove across width. Limenandra nodosa Haefelfinger & Anterior digestive system of two rows and a Stamm, 1959 (Bertsch 1972, Eveline and reduced row, consisting of only two cerata; Ernst Marcus 1970) posterior digestive system of 16 rows of

a;gm· Ii;;;;$! Ii ;z Mil II • Aeolidacea of the Hawaiian Isiands-GOSLINER 67

E

E .,.,E

2mm

F

lmm

penis

pr

FIGURE 19. Baeolidia nodosa: A, ventral view of a living animal; B, lateral view of anterior portion of preserved animal; C-E, cerata showing variability in shape and orn,!lmentation; F, radular tooth; G, jaw; H, reproductive system (a.g., albumen gland; am., ampulla; m.g., mucous gland; me.g., membrane gland; o.t., ovotestis; pr, prostate; LS., receptaculum seminis). 68 PACIFIC SCIENCE, Volume 33, January 1979 cerata on each side. Cerata (Figure 19C-E) new genus and species, Limenandra, with irregular in shape, dorsoventrally flattened, L. nodosa as the type, and included Baeolidiq some papillate; apex of one to three cnido­ fusiformis Baba, 1949 in Limenandra. They sacs within single ceras; tip sharply recurved differentiated Limenandra, noting that aH or moderately arched. Gonopore between cerata are arranged in rows rather than in anterior and posterior digestive systems. horseshoe-shaped arches, with fewer row's Anus between first and second rows of pos­ than in Baeolidia, and the cerata are round terior digestive system. Radula (Figure 19F) rather than flattened as in Baeolidia. Th~ uniseriate, consisting of 21 pectinate teeth, shape of the cerata in Limenandra nodosa, 100 to 250 J1.m wide with 30 to 70 denticles. can be dorsoventrally flattened (Bertsch Jaws (Figure 19G) weak, 2.3 mm long, with 1972). Eveline and Ernst Marcus (1970) smooth masticatory border. Reproductive suggested that the shape of the cerata (flat~ system (Figure 19H) with elongate ovotestis, tened in Baeolidia, fusiform in Berghia) was convoluted ampulla. Receptaculum seminis the only acceptable difference and place4 large, bilobed, with elongate duct. Vas Baeolidia fusiformis in Berghia while main1 deferens elongate, convoluted, prostatic taining Limenandra nodosa. Basing the throughout most of its length. Penis paddle­ generic separation on the ceratal shape cre~ shaped, unarmed. ates more problems than it solves, becausje this placement unites species that have arche~ DISCUSSION: The generic distinctions for in the digestive system with species that have Berghia Trinchese, 1877, Baeolidia Bergh, linear rows. Also, species such as Limenandrq 1888, and Limenandra Haefelfinger & nodosa have both round and flattened cerata. Stamm, 1959 are confusing. All are members It seems that a more reasonable approach of the Aeolidiidae and have papillate rhino­ would be to place those species with the phores. Bergh (1890, 1892) contended that digestive system arranged in arches in Ber~ deeply incised radular teeth in Berghi(1 and ghia, as in the type, Berghia coeru/escens, and evenly curved teeth in Baeolidia served as an those with cerata in linear rows in Baeolidia; essential generic distinction. Ernst Marcus as in the type, Baeolidia moebii. Limenandrq (1958) argued that this radular distinction would then be considered as a junior syn~ is invalid on a generic level. Drawings by onym of Baeolidia since a slightly more Baba (1955, fig. 56) illustrate Baeolidia major anterior anal position in B. fusiformis and B. amakusana with both indented and evenly nodosa does not warrant generic separatioq, curved radular teeth. When one compares the reproductive Marcus (1958) attempted to settle this systems of the species for which the gener~ confusion by comparing the type species of have been described, some differences the two genera. The type of Baeolidia, B. emerge. In both Baeolidia benteva Marcus; moebii Bergh, 1888, has linear rows of cerata 1958 and Baeolidia nodosa (present paper), in the right anterior digestive system, while the receptaculum seminis is bilobed. In Berghia coerulescens (Laurillard, 1830) has Berghia coerulescens (Tardy 1962), B. verru~ cerata arranged in horseshoe-shaped arches. cicornis (Tardy 1962), B. erutzberghi (Eveline Marcus contended that the difference is and Ernst Marcus 1970), and B. major (pre­ sufficient to distinguish the genera. Marcus sent paper), the receptaculum is simple. This (1958) confirmed that Berghia coerulescens is a potentially valuable characteristic for has the anus in the first ceratal arch of the generic separation. ' posterior digestive system while Baeolidia Baeolidia nodosa has been found spora­ moebii has the anus located posterior to this dically at a variety of localities, which position, and utilized this feature to distin­ suggests a wide distribution. Haefelfinger guish the genera. Tardy (1962) considered and Stamm (1959) described the species frorn this a questionable generic character because 25 specimens from a single locale in Ville~ the position of the anus in Baeolidia moebii franche, France. Eveline and Ernst Marcus is not known. (1970) reported a single specimen from Haefelfinger and Stamm (1959) erected a Bonaire in the southern Caribbean. Bertsch ': Aeolidacea of the Hawaiian Islands-GosLINER 69

(1972) described a single specimen from the 1955; Berghia amakusana (Marcus, Gulf of California. The presence of this 1958, Farmer 1966); Berghia major species in Hawaii further substantiates the (Edmunds, 1970) circumtropical distribution of the species. The morphology of Baeolidia nodosa, with DISTRIBUTION: Tanzania (Edmunds 1970, the exception of the reproductive system, Eliot 1903); Japan (Baba 1933, 1937, 1955); was included in the original description. Australia (Allan 1947); New Zealand (Mor­ Eveline and Ernst Marcus (1970) stated only ton and Miller 1968); and Mexico (Farmer tpat the external morphology was virtually 1966). iclentical to Haefelfinger and Stamm's mate­ HABITS AND OCCURRENCE: More than fifty ria,l. Bertsch (1972) stated that his animal specimens have been found on the under" was somewhat intermediate between B. surface ofrocks at the Kewalo Basin seawall, nodosa and B. fusiformis. He noted that the from the floating docks and thermal pollu­ radular rows and coloration are similar tion study tables at Coconut Island, and between the two species, and the cerata from from patch reefs in Kaneohe Bay, Oahu, in his animal resemble those of Haefelfinger all months ofthe year. Egg masses also occur apq Stamm's material despite their statement throughout the year, usually in close prox­ that B. nodosa has round cerata. Bertsch imity to one or more specimens. Berghia stated that the yellow-pink colored circles major feeds on the sea anemones Boloceroides from his specimen were intermediate be­ sp. and Aiptasia sp. It is always in close prox­ tWt:en the white, yellow, white, red circles imity to one of these species of anemones, of B. nodosa from France and the yellow more commonly Aiptasia. However, there spots of B. fusiformis from Japan. The does not appear to be any feeding preference y¢llow, blue-green circles of the specimen between the two species. from Hawaii demonstrate that the essential dffference between B. nodosa and B. fusi­ DESCRIPTION: Length 10 to 100 mm. Gen­ formis is not in the color similarities but the eral body color variable: translucent white, distinct arrangement ofcoloration in circular golden brown, dark gray-green; many varia­ ~rtits on the dorsal surface of B. nodosa. tions in pattern of pigmentation; head region Bertsch also said that the rhinophores of his may have spot of dark pigment or may lack specimen more closely resemble those of dark pigment; cerata, especially in specimens 13. fusiformis, but described papillae begin­ exceeding 30 mm in length, may be orna­ niIlg halfway up the length ofthe rhinophores mented with opaque white spots and sub­ with very few on the anterior surface. This apical blue rings. Rhinophores most fre­ a,ccurately describes the figure of the rhino­ quently densely papillate, but occasionally phores of B. nodosa (Haefelfinger and Stamm may virtually lack papillae. Papillations 1959, fig. IB) and can also apply to all strikingly more dense on posterior side, members of Berghia and Baeolidia. The pre­ although variable. Oral tentacles elongate sence of papillate cerata, distinct circles of and used extensively in feeding where they coloration, and an elongate masticatory establish contact with prey anemone. When margin ensure that the specimen from contact is made with the anemone, an initial Mexico is indeed B. nodosa. The specimen period of I to 3 sec of contact followed by from Hawaii fits the original description of quick withdrawal is observed. The behavior B. nodosa in color pattern, external morpho­ pattern is a rapid sequence in which the logy, radula, and jaws, and eliminates any tentacles firmly grasp the anemone while doubts that it is conspecific and that B. engulfing it. Foot corners elongate and ten­ fusiformis represents a distinct taxon. tacular. Anterior border of foot with sharp notch. Cerata numerous, always dorsoven­ Berghia major (Eliot, 1903) trally flattened, although degree of flattening Baeolidia major Eliot, 1903 (Allan 1947, is variable; smaller animals of 10 to 20 mm Baba 1933, Morton and Miller 1968); with only slightly flattened cerata, while Baeolidia major amakusana Baba, 1937, cerata almost paddle-shaped in animals 20 70 PACIFIC SCIENCE, Volume 33, January 1979

I 200 JIm

lmm

E

m.g. r.S.

I 2mm

FIGURE 20. Berghia major: A, B, radular teeth; C, D, jaws; E, reproductive system (a.g., albumen gland; am, ampulla; m.g., mucous gland; me.g., membrane gland; o.t., ovotestis; r.s., receptaculum seminis). to 100 mm; cerata of anterior right digestive more or less entire at apex. Teeth progres­ system in a single horseshoe-shaped arch; sively smaller in posterior region of radula, first one to four ceratal groups of posterior shape of entire radula tapered toward pos­ system in this arrangement as well; cerata terior. Teeth in 16 to 28 rows, with 25 to 65 posterior to second to fifth arches in linear denticles on anterior margin. Denticles rows. Gonopore ventral to anterior portion somewhat recurved inwardly in more lateral of anterior ceratal group. Anus within first portion, becoming progressively straighter horseshoe of posterior right digestive system. toward center. Jaws (Figure 20C, D) 1.6 to Radular teeth (Figure 20A, B) 100 to 650 J.l.m 40 mm long, also variable, although general wide, uniform in shape within single speci­ shape rather constant. Most individuals with men. Teeth indented at central region in highly denticulate masticatory border of 50 some specimens. Most individuals with teeth to 110 denticles, but some specimens with Aeolidacea of the Hawaiian Islands-GosLiNER 71

TABLE 2 MORPHOLOGICAL VARIABILITY IN Berghia major (ELIOT, 1903)

MASTICATORY SOURCE LENGTH (mm) COLOR CERATA RADULA EDGE

Edmunds 1970 25 Pale gray with Flattened 21 teeth Smooth Tanzania brown and blue 4 arches 36-84 denticles 7 rows Baba 1955 60 Yellow-brown Flattened, 32 teeth Smooth Japan leaflike 140 denticles 7 arches 8 rows Farmer 1966 20 Yellow-brown 22 teeth Mexico Allan 1947 20-25 Light gray-brown Leaflike 32 teeth Australia with bluish cerata Eliot 1903 40 Grayish white Flattened 32 teeth Smooth Zanzibar 17 groups 150 denticles Morton and Miller 32 Brown, orange, with 1968 pale blue and New Zealand crimson Present paper 10-100 Translucent white, Flattened 16-28 teeth Smooth or Hawaii and dark gray-green, dorsoventrally 25-65 denticles denticulate Mexico or golden brown 2-5 arches with opaque white 3-6 rows and blue smooth striated border. Reproductive system noted that Eliot made no distinction of (Figure 20£) of two extremely large organs ceratal distribution, but contended that and several smaller ones. Ovotestis generally Baeolidia major should probably remain in yellow and comprising entire posterior half Baeolidia becaude the anus in Eliot's animals of animal. Female gland mass consists of was situated behind the second ceratal group albumen, membrane, and mucous glands, and maintained the separation of the two and comprises bulk of anterior region of species (Ernst and Eveline Marcus 1970). animal; it consists of several lobes and is Edmunds (1970) found specimens from Tan­ transparent in the living animal. Penis elon­ zania attributable to Baeolidia major Eliot, gate and unarmed. Receptaculum seminis 1903. He stated that these animals were small and inconspicuous. Ampulla wide conspecific with Eliot's, but did not give and convoluted. reasons. Based on a single horseshoe of cerata in the right anterior digestive system, DISCUSSION: Berghia major was described he placed Baeolidia major in Berghia. Ed­ from Zanzibar as Baeolidia (Eliot 1903). munds also synonymized Berghia amakusana Baba (1933) attributed animals from Japan as a junior synonym of B. major without to Baeolidia major but later (Baba 1937) giving a basis for the synonymy. From this distinguished B. major amakusana as a sub­ study it is readily evident that Baeolidia species, without giving a basis for distinction major and Berghia amakusana are conspe­ between his specimens and those described cific. Several characteristics of Berghia major by Eliot. are summarized in Table 2. Ernst Marcus (1958) transferred Baeolidia In Hawaii, Berghia major feeds on the sea major amakusana to Berghia, while simul­ anemones Aiptasia sp. and Boloceroides sp. taneously raising it to specific status as Specimens from New Zealand feed on the Berghia amakusana on the basis of ceratal anemone Cricoplorus nutrix (Morton and arrangement and anal position. Marcus Miller 1968). Body color is variable. The 72 PACIFIC SCIENCE, Volume 33, January 1919 specimens from Hawaii exhibit most of the added significant information. The opistho­ color range seen in the species known from branch fauna of Japan has been exceedingly other collections throughout the Indo­ weli studied (Baba 1930, 1949, 1955, Eliot Pacific. A specimen collected by Farmer in 1913). the Gulf of California shows little difference The opisthobranch fauna of the Hawaiiah from one specimen collected from Kaneohe Islands has only recently been well studied. Bay (Keen 1971, pI. 22, fig. 5), and, indeed, The zoogeographical affinities of the the variation within the Kaneohe Bay popu­ Hawaiian marine mollusks have been studied lation is more substantial than between these by Kay (1967), who estimated that approxi­ two individuals. mately 20 percent of the Hawaiian marine The Hawaiian specimens range from 10 mollusks are endemic to the archipelago. to 100 mm in length, which exceeds previous There is not enough information about the records for the species. The cerata of all four aeolids known only from the Hawaiian B. major recorded are dorsoventrally flat­ Islands (Flabellina annuligera, Cuthona dirni­ tened to varying degrees. nutiva, Noumeaella sp., Aeolis parvula) to The radula in the Hawaiian specimens imply that any species are, in fact, endemiC. consists of 16 to 28 rows of teeth, with 25 to Eight species, Flabellina alisonae, Cuth0'1a 65 denticles. Specimens have been recorded pinnij'era, Embletonia gracile, Facelinella that have 32 rows of teeth (Allan 1947, Baba semidecora, Favorinus japonicus, Hervielta 1955, Eliot 1903) with 140 denticles (Baba mietta, Aeolidiella takanosimiensis, arid 1955) and 150 denticles (Eliot 1903). Spurillajaponica, are reported from localities Probably the most unusual variation with­ within the Pacific basin. Five species, in the Hawaiian specimens occurs in the Phestilla sibogae, P. melanobrachia, Calaria jaws. All other citations of Berghia major militaris, Pteraeolidia ianthina, and Berghia report a smooth masticatory border. Several major, are known to occur in both Indian specimens from Hawaii have a smooth and Pacific oceans. Five species, Cuthd,ui masticatory. border, but most specimens perea, Godiva quadricolor, Glaucus atlanticus, from Oahu have from 50 to 110 denticles Spurilla neapolitana, and Baeolidia nodosa; on the cutting edge. are known to occur in the Atlantic and The reproductive system of Berghia major Pacific and are considered circumtropical. is previously unrecorded. It is very similar This latter group represents a significant to that of B. coerulescens (Laurillard, 1830) percentage of aeolids that apparently are (Tardy 1962) and B. crutzberghi Marcus & widespread in their distributions. Marcus, 1970. It is interesting that 15 of the 22 species described in this paper have been previously reported from Japan. Baba (1949) stated that the Japanese fauna is "rich in Indo­ ZOOGEOGRAPHICAL DISCUSSION Pacific elements." The zoogeographical affinities of the Members of the family Aeolidiidae appear opisthobranchs of the Indo-Pacific region to be particularly widespread.' Baeolidia are poorly understood because vast areas nodosa and Spurilla neapolitana are circutn­ have not been explored. Other areas have tropical, while Berghia major is known from had only cursory examination. In the Indian the Indo-Pacific from Tanzania to Mexico Ocean, only Tanzania (Edmunds 1969, 1970, (Edmunds 1970). Aeolidiella takanosimiensis Eliot 1902, 1903, 1904a, 1904b) has been is known from Japan to California, and examined to any degree. In the Pacific, Spurilla japonica is known from Japan arid restricted areas of Australia have been well Hawaii. Aeolidiapapillosa is boreal-temperate collected (Burn 1963, 1964, 1966). New in its distribution (Ernst Marcus 1961) as it Caledonia (Risbec 1928, 1953) has been has been found from Europe, both coasts fairly well studied. The recent work of of North America, northern Japan, arid Carlson and Hoff (1973) from Guam has southern Chile. Members of the family feed Aeolidacea of the Hawaiian Islands-GosLiNER 73 on sea anemones and seem to have a high ---. 1936. Opisthobranchia ofthe Ryukyu degree ofplasticity in the number ofavailable Islands. J. Dept. Agr. Kyushu Imp. Univ. prey species. 5: 1-50. ---. 1937. Opisthobranchia of Japan. 2. J. Dept. Agr. Kyushu Imp. Univ. ACKNOWLEDGMENTS 5(7): 289-344. ---. 1949. Opisthobranchia of Sagami I wish to extend my gratitude to E. Alison Bay. Iwanami Shoten, Tokyo. 194 pp. kay for her guidance, cooperation, and ---. 1955. Opisthobranchia of Sagami ~riticism, and for her willingness to assist Bay. Suppl. Iwanami Shoten, Tokyo. in my endeavors. I wish also to thank other 59 pp. triends and colleagues who assisted in the ---. 1959. A new record of an interesting collection of specimens and other aspects species, Embletonia gracile, Risbec, from 9f this research: Becky Abildskov Houck, Japan (Nudibranchia-Eolidacea). Publ. Frederick J. Lighter, and Steven Syrop. I Seto Mar. BioI. 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