On the World-wide Dispersal of a Hawaiian Barnacle, Balanus amphitrite hawaiiensis Broch'

HUZIO UTINOMI 2

IN RECENT YEARS a great deal of attention has forms (or subspecies); and to present the al­ been given by many workers, especially by leged distribution of these related forms, as far marine ecologists and oyster planters, to the in­ as can be determined from the records of their troduction of an Australian barnacle, Elmin ius occurrence. modestus Darwin, into British waters and to its subsequent rapid spread along the continental ACKNOWLEDGMENTS European coasts from Brittany to the mouth of I am especially indebted to Mr . William A. the Elbe River. The most recent extensive re­ Newman, Pacific Islands Central School, Truk, views of the occurrence of this barnacle on those East Caroline Islands, for data of occurrence in coasts have been given by Den Hartog ( 1953) America, and to Mr. Kasio Ora, Zoological In­ and Kiihl ( 1954). Similarly, another instance stitute, Agriculture Faculty, Hokkaido Univer­ of the introduction of an adventive barnacle into sity, for information on habitats. Thanks are British coastal waters has also focused more at­ also due to Dr. Masaru Kato, Zoological Insti­ tention on the possibility of other importations tute, Kyoto University, for translating a Russian of many harmful foreign animals, as well as the paper. extension of their natural geographical distribu­ tion. These studies strongly suggest that trans­ IDENTITY OF SUBSPECIES denticulata WITH portation on ships' bottoms or as stowaways SUBSPECIES hawaiiensis among oysters is the chief means of migration (Allen, 1953; Coe, 1956). Balanus amphitrite Darwin ( 1854) com­ The second example of a barnacle imported prises many local forms , occasionally treated as into European waters has long been considered either "subspecies" or "varieties," which are to be Balanus amphitrite var. denticulata Broch, diversified in the number of teeth on the labrum. which was originally described by Broch (1927 ) Most of the subspecies have only three teeth on from the Suez Canal. Since the first record of its each side of a median notch. The other sub­ occurrence on the southern coasts of Britain by species almost always have four teeth, or a larger Bishop (1950) , most of the European workers, number (Hiro, 1938). Even in the subspecies especially field ecologists who are not acquainted having three or four teeth, the actual number is with the Pacific species of barnacles, have over­ apt to vary among specimens, but in such cases looked the fact that it is identical with Balanus the innermost one is either small or only rudi­ amphitrite forma hawaiiensis Broch (1922) , mentary. which is thought to be native to the Hawaiian The two subspecies, hawaiiensis and denticu­ Islands. lata, have such a multidenticulate labrum. The The object of this paper is to state the reasons same is true of Balanus eburneus Gould and of for considering both of these forms as identical, B. improvisus Darwin, which are found as mi­ at least with each other and possibly with other grants on ships sailing between America and Europe. The number of teeth is also variable, 1 Contributions from the Seta Marine Biological ranging between 8 and 20; in most cases 10 to Laboratory, No. 335. Manuscript received January 2, 13 larger teeth are found on each side, but never 1958. fewer than 8. 2 Seta Marine Biological Laboratory, Sirahama, Wakayama-ken, Japan. The most outstanding features in external ap-

43 44 PACIFIC SCIENCE, Vol. XIV, January 1%0

pearance which prove the identity of both forms DIFFERENCES BElWEEN SUBSPECIES hawaiiensis are the colored strip es on the wall and the shape AND SUBSPECIES communis of the opercular valves. The shell is glossy white, Notwithstanding the peculiarities mentioned smooth, and furnished with dark violet longi­ above, sometimes there has been confusion tudinal strip es not crossed by any transverse among cirripede systematists between forma stripes, though growth lines are faintly defined. hawaiiensis (or denticulata) and Darwin's vari­ Usually a broad white area is present in the ety communis (see Table 3). middle of the larger compartments (carina, For example, N ilsson-Cantell (1938: 37) laterals, and rostrum ), and a similar white area recognized both subspecies as distinct, but men­ is present along both sides, the colored stripes tioned that "I do not consider it impossible that being indistinct or quite absent there (see Table they (denticulata ) represent a special race from 1) . The pari etal tubes (the number of which the canal zone, but I do not think that numerous corresponds with the number of the colored teeth on the labrum are a special characteristic stripes, both distinct and obsolete) are narrow of the subspecies denticu lata." He emphasized and numerous; the longitudinal septa on the that a great variability in the number of teeth inner lamina are strongly developed and run up on the labrum is not always of systematic im­ to the sheath as high ridges which are very finely portance, remarking that "the typical communis denticulate on both sides. The radii are very also has numerous teeth on the labrum." How­ broad, with summits almost parallel to the base ever, I most emphatically dissent from his (Fig. 1). opinion in this respect. As he says,it is true that In describing the new variety denticulata the balanids are highly variable as far as a par­ from the Suez Canal, Broch (1927) did not ticular character is concerned, but I hold that mention its close affinity with his earlier form there are certainly some admittable limitations hawaiiensis described in 1922. A comparison of in the range of variation. A similar unreliability his descriptions and figures of both forms with in identifications based only on the labrum was the Japanese specimens of hawaiiensis clearly pointed out by Pilsbry (1916: 88 ), in the case shows that there is no difference important of Balanus improvises. enough to separate them as different subspecies. Some specimens of subspecies communis re­ Therefore, I believe that denticulata must be the semble subspecies hawaiiensis superficially, be­ same as hawaiiensis. ing often found together in the same place, but

TABLE 1 THE N UMB ER OF LONGITUDIN AL COLORED STRIPES IN ALL COMP ARTMENTS OF Balanus amp hitrite hawaiiensis AND Balanus amphitrite communis"

LEFT RIGHT CARINO- CARINO-LEFT RIGHT SIZ E OF LATERAL LATERAL LAT ERAL LATERAL SPECIMENS CARINA COM- COM - COMPART- COM PART- ROSTRUM PART- PART- MENT MENT MENT MENT B. a. hawaiiensis c.r.d. 15.4 mm. (0) 5 (1 ) 5 (1 ) (1)2(1) ( 0)2(2) (1)4(2)4(1) (0) 5(1)4(2) (2)5(2)5(2 ) c.r.d. 18.0 mm. ( 1)3(2) 4 (1) (1 )2(1) ( 1) 2 ( 2 ) ( 2 ) 4 (2 ) 4 ( 1) ( 1) 5(2)4(2 ) (2 ) 5(3 )6(1 ) c.r.d. 18.5 mm. (0) 5(2) 4 ( 0) (0)3 (1) (1 )2(1) (2 )5(1) 3 (1) (1)4 ( 1)4(0) (2 ) 4(2) 4(3 ) c.r.d. 19.6 mm. (2 )4(1 ) 3(1) (1) 2(1) ( 1)2(1) (1 ) 3 (3 ) 4 ( 2 ) (2)2 ( 2) 2(0) (1 ) 5 (2) 4 (2) B. a. communis c.r.d. 14.0 mm. 10 3 2 10 9 9 c.r.d, 18.2 mm. 8 2 2 9 9 9 c.r.d, 20.0 mm. 9 4 2 10 10 10 . . • c.r.d. =carino -rostral diameter. Numbers with out parentheses represent distin ct stripes, Numbers rn parentheses represent obsolete or reduced stripes. T he total num ber corr esponds with the number of parietal tubes in each of the comp artments. Hawaiian Barnacle-UTINOMI 45

FIG . 1. Balanus amphitrite hau/aiiensis Broch (a, b and e) and B. amphitrite communis Darwin (d): a, Profile of left lateral compa rtment, showing the arrangement of longitudinal colored stripes, either distinct or indistinct (dotted in the figure ); b, c, basal view of parietal tub es; d, e, labrum. the two differ in several important details of Pearl Harbor, Hawaii (Broch, 1922 ) . structure, as summarized in Table 2. Honolulu Harbor, Hawaii (Pilsbry, 1928). Consideration of these characteristics will Sasebo, Kure , Maizuru, Seto, and Misaki, prove that the two subspecies are separable from Japan (Hiro, 1937, 1938 ). each other when large series of specimens are Persian Gulf (Nilsson-Cantell, 1938) . examined. Suo, Kiirun, Tansui , Takao, and Mako, DISTRIBUTION AND SYNONYMY Formosa (Hiro, 1939) . Aio, Seto Inland Sea, Japan (Hudinaga and Herewith is presented a summary of accounts Kasahara, 1942) . of the corresponding subspecies of Balanus am­ Pearl Harbor and Kaneohe Bay, Hawaii phitrite Darwin given by previous workers and (Edmondson and Ingram, 1939; the supposed geographical distribution of B. Edmondson, 1946) . amphitrite hawaiiensis Broch (Fig. 3). The evi­ Misaki, Japan (Hirano and Okushi, 1952) . dence is far from complete and some records of occurrence are not conclusive, since most of the 2. Recorded as B. a. denticulata: subspecies or forms of Balanus amphitrite, espe­ Suez Canal (Broch, 1927; Ciurea et al., 1933; cially the most familiar communis, have been Monod, 1937). insufficiently described (often without detailed Southern coasts of Britain (Bishop, 1950; figures), or are recorded only by a subspecific Norris et al., 1951; Crisp and Molesworth, name. Nevertheless, those I have listed may help 1951). furore workers towards the solution of the prob­ Atlantic coasts of France (Bishop et al., lems presented herein. 1957). 1. Recorded as B. a. hawaiiensis: Knysha Estuary, South Africa (Millard, Kaladis Point, Mindanao (Broch, 1922). 1950) . 46 PACIFIC SCIENCE, Vol. XIV, January 1960

Queensland and Torres Strait, Australia 6. Recorded as B. amphitrite, subspecific name (Allen, 1953). not given : Durban Bay, South Africa (Day and Morgan, Atlantic coasts of France (Prenant, 1929; 1956) . Fischer, 1929; Fischer-Pierre, 1932, erc.). 3. Recorded as B. a. communis (in part) : ? Biscayne Bay, Florida (Weiss, 1948). Ismailia, Timsah Sea, Suez Canal (N ilsson­ Oakland Estuary, California (Graham and Cantell, 1921). Gay, 1945 ) . ? Billiton, Sunda Islands (N ilsson-Canrell, 7. Unpublished records of occurrence: 1921) . San Diego, Salton Sea, and San Francisco Bay, ? Messina port, Italy (Nilsson-Carnell, 1921). California; and the Hawaiian Islands (New­ Istanbul harbor, Turk ey (Neu, 1939). man, 1955, 1956: personal communica­ Middle Harbor, Port Jackson, Australia tion ) . (Pope, 1945 ) . Dakar, West Africa (from I.F.A.N. Adriatic Sea (Kolosvary, 1947). Collection) . Black Sea (Zevina and Tarasov, 1957) . Seto Inland Sea and Ise Bay,Japan (Ora, 1955: personal communication) . 4. Recorded as B. a. venustus: A detailed discussion of the synonymies listed Krusadai, Gulf of Manaar, Ceylon (Raj, above is omitted, for the sake of saving space, 1927) . but the preceding illustrations and descriptions 5. Recorded as B. a. franciscanus and B. a. herzi: • I am not certain whether another new subspecies, San Francisco Bay, California (Rogers, B. a. saltonensis Rogers, from the Salton Sea, is the 1949) .3 same as the others or is distin ct from them.

" a c . ... . '\ ,: "\

d

FIG. 2. Balanus amphitrite hawaiiensis Broch (a and b) and B. amphitrite communis Da rwin (c and d): a and c, Inside of tergum; band d, insid e of scutum. Hawaiian Barnacle-UTINOMI 47

TABLE 2 DIFFERENCES IN DETAILS OF ST RUCTURE B ETW EEN Balanus amp hitrite hawaiiensis AND B. amp hitrite communis

DETAILS OF STRUCTURE B. amphitrite hawaiiensis B. amphitrite communis

Ground color of shell.,; Glossy white Rather reddish white, darkened with bluish tint in upper part Outside of pa rietes.c.c.... Onl y dark violet longitudinal stripes Pale or dark violet longitud inal stripes sharply defined; those in middle and crossed by the same colored or red­ along both sides indistinct or reduced dish trans verse stripes on the whole (Fig . Ia) area Inside of parietes__._._ ____ Strongly ribb ed up to the sheath; ribs Slightly ribb ed near base; ribs low, with flat on roof and finely denticulate on slight denticula tion at base which both sides but not bifurcate distally may extend as bifurcate secondary (Fig. Ib) ribs; a few shorter riblets often in­ terseptally (Fig. l c) Parietal tub es .. Numerous (a bout 16 in rostrum) , nar- Few (a bout 9 in rostrum ), large and row and square at base (Fig. 1b) rectangu lar at base ( Fig. l c) Radii...... __ ...... Very broad, flat, with almost level sum - N arrow, slightly sunken, with very ob­ mits, forming an angle of about lique summits, forming an angle of 100 0 _ 110 0 with the interlocking about 140 0 with the interlocking margin (Fig. l a) margin Rostrum ...... __ .. .. Apex usually incurved Apex often turned outwards when iso­ lated Scutum ..__ Depression below adductor ridge deep, Depr ession below adductor ridge shal­ oblongly circumscribed ( Fig. 2b) low, lengthened along adductor ridge (Fig. 2d ) Tergum...... __ ..... Spur very broad, over Y3 width of the Spur longer than wide, with more or less valve, short, either rounded or trun­ hatchet-shaped end, separated from cate distally and separated from basi­ basi-scutal angle by its own width scutal angle by less than its own which is less than Y3 that of the width; crests for depressor muscles valve; crests for depressor muscles strongly developed, typically 5 (Fig. not so strong, exceeding 5 in num­ 2a ) ber (Fig. 2c) Labrum__ Multidenti culare on each side of a me- 3 teeth , or with a smaller additional one, dian notch (Fig. 1e) on each side (F ig. Id)

of this subspecies, compared with the typical Islands in the mid-Pa cific trop ical sea. From this communis, will help to a certain extent in later mid-Pacific area it may have spread much earlier identifications. and more rapidly in all seas than did the Aus­ tralasian barnacle, Elminius m odestus Darwin, NATIVE SEA AND HABITAT in European and South African waters. It is sup­ From these data of occurrence, it is very likely posed that the first establishment of this Ha­ that the subspecies hawaiiensis ( = denticulata) waiian barnacle took place from the tropical is not a local variant restricted to such a small west Pacific, such as the Philippines and For­ area of special environments as the Suez Canal, mosa, lying along the N orth Equatorial Curr ent, but nowadays is very widely distributed in cir­ where it is directed westerly, although the most cumequatorial tropical and temperate areas in probable means of migration may have been at­ all oceans. tachment on the hulls of ships. Its native habit at is probably the Ha waiian In these tropical and subtropical areas the 48 PACIFIC SCIENCE, Vol. XIV, January 1%0

FIG. 3.Geographical distribution of Balanus amph itrite hawaiiensis Broch.

barnacles grow abundantly in the intertidal zone organisms of naval ships, I am reminded that of sheltered coasts, usually below the mean sea B. amphitrite hawaiiensis was predominant in level; sometimes they occur in clusters on wharf Kure port, on the Seto Inland Sea, but was less piles, coastal rocks, and mangrove roots on the common in any of the bays facing open seas leeward side but not on seaward reefs with along the Pacific coast of Jap an. There the domi­ flourishing reef corals. nant form was B. amphitrite communis. In the more northern temperate areas, such In this connection it is interesting to find a as the Japanese and Californian coasts, the bar­ similar change of habitats in B. amphitrite den­ nacles are generally restrict ed to quiet bays or ticulata ( = B. a. hawaiiensis) established on the harbors of enclosed water and are usually at­ Atlantic coasts of France (Bishop et al., 1957). tached to floats, buoys, ship bottoms, or sub­ According to them, it is always submerged on merged panels under the low tide level. the northern coast but is rare or absent on Th ey seem to have a tolerance for a wide the southwestern coasts of highly estuarine range of salinities (on the average S 16-30 condition. o / 00) , for they are found in the main harbor From these ecological evidences, it seems most waters where the salinity is that of normal sea appropr iate to say that B. amphitrite hawaiiensis water but may be subject to at least occasional is a protected-water form rather than a poly­ diluti on with fresh water. In more estuarine haline form. B. amphitrite communis, on the situations they are replaced by B. amphitrite other hand, is a stenohaline subtidal form; and albicostatus Pilsbry or by B. amphitrite krugeri Elminius modestus is a more estuarine intertidal Nilsson-Cantell. On the other hand, in more form. Th is last-named barnacle was originally saline waters towards the mouth of bays, B. found in a temperate habitat, in European wa­ amphitrite communis as well as B. trigonus Dar­ ters, where it had established itself, so that it win dominate in the subtidal zone. A similar may extend farther northward than does B. am­ change in barnacle communities has been noted phitrite hawaiiensis. by Day and his collaborators (1956) during At any rate, marine animals living in an en­ their ecological survey of South African estuaries. vironment of variable salinity have their repro­ From my experience of studies on the fouling ductive cycle of spawning correlated with the H awaiian Barnacle-UTINOMI 49 temperatur e cycle rather than with salinity. tions of littoral marine invertebrates. J. Mar. Therefore, as might be expected, there is no Res. 15 (3) : 212-232. particular correlation between successful estab­ CRISP, D. J., and A. H . N . MOLESWORTH. 1951. lishment of fouling organisms and the variable Habitat of Balanus amphitrite var. denticulata salinity of sea water in the different regions in Britain . Nature 167: 489. where the barnacle makes a successful settlement. DARWIN, CH. 1854. A monograph on the sub­ class Cirripedia. Th e Balanidae, Verrucidae, SUPPLEMENTAL NOTE etc. viii + 684 pp., 30 pIs. Ray Society, After the present paper was submitted, an London. art icle by John D. Costlow and C. G. Bookhour, DAY, J. H ., and J. F. C. MORGAN. 1956. The entitled "Larval development of Balanus amphi­ ecology of South African estuaries. Part 7. trite var. denticulata Brach reared in the lab­ The biology of Durban Bay. Ann. Natal Mus. oratory," app eared in the Biological Bulletin, 13 (3): 259-312, pI. 4. 114 (3 ): 284-295, June, 1958. In this paper DEN HARTOG, C. 1953. Immigration, dissemi­ they confirmed the fact that B. amph. denticulata nation and ecology of Elminius modestus Dar­ occurs abundantly as an important fouling or­ win in the N orth Sea, especially along the ganism at Beaufort, N orth Carolina, on the east Dutch coast. Beauforti a 4 (33) : 9-20. coast of N orth America, and that it is wholly identical with B. amph . hawaiiensis in all EDMONDSON, CHARLES HOWARD. 1946. Reef naupli ar stages during metamorphosis. The oc­ and shore fauna of Hawaii. Spec. Publ. Bishop curre nce of this subspecies there may be sup­ Mus. 22. iii + 381 pp. ( Rev.) Bernice P. posed also from the illustrations given in W eiss Bishop Museum , Honolulu. (1948). EDMONDSON, C. H., and W . M. INGRAM. 1939. Fouling organisms in Hawaii. Occ. Pap. REFERENCES Bishop Mus. 14(14 ) : 251-300, 9 pls. FISCHER, ED. 1929. Le cirripede Balanus amphi­ ALLEN, F. E. 1953. Di stribution of marine in­ trite Darwin a Saint-Servan. Bull. Lab. Mus. vertebrates by ships. Aust. J. Mar . Freshw. H ist. N at. St-Servan 4: 10-11. Res. 4 (2 ) : 307-316. FISCHER-PIETTE, ED. 1932. Vitesse de croissance BISHOP, M.W . H. 1950. Distribution of Balanus de quelques organismes marins. Bull. Lab. amphitrite Darwin var. denticulata Brach. Mus. H ist. N at. St-Servan 10: 17-22. N ature 165: 409. GRAHAM, H. W., and H. GAY. 1945. Season of BISHOP, M. W .H ., D. J. CRISP, E. 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--- 1939. Studies on the Cirripedian fauna NORRIS, E., 1. W. G. JONES, T. LOVEGROVE, of Japan. IV. Cirripeds of Formosa (Taiwan), and D . J. CRISP. 1951. Variability in larval with some geographical and ecological re­ stages of cirripeds. Nature 167: 444-445. marks on the littoral forms. Mem. Cell. Sci. PILSBRY, HENRY A. 1916. The Sessile Barnacles Kyoto, Ser. B, 15(2): 245-284. (Cirripedia) Contained in the Collection of HUDINAGA, M., and H. KASAHARA. 1941. On the U. S. National Museum; Including a the rearing and metamorphosis of B. amphi­ Monograph of the American Species. Bull. trite hawaiiensis Broch. Zool. Mag., Tokyo U. S. Nat. Mus. 93. xi + 366 pp., 76 pls. 54(3): 108-118. (In Japanese, with English ---1928. Littoral barnacles of the Hawaiian resume.) Islands and Japan. Proc. Acad. Nat. Sci. Philad. KOLOSvARY, GABRIEL. 1947. Die Balaniden der 79: 305-317, pls, 24-26. Adria. Ann. Hist.-Nat. Mus . Hung. 40 (1) : POPE, E. C. 1945. A simplified key to the sessile 1-88. barnacles found on the rocks, boats, wharf KUHL, H. 1954. Uber das Auftreten von El­ piles and other installations in POrt Jackson Darwin in der Elbrnundung. minius modestus and adjacent waters. Rec. Aust. Mus. 21 (6) : Wiss. Meeresunters. Helgoland. 5( 1): 53-56. 351-372, pls, 28-30. MILLARD, N. A. H. 1950. On a collection of PRENANT, M. 1929. Balanus amphitrite Darwin sessile barnacles from Knysna estuary, South sur les cotes atlantique francaises. Bull. Soc. Africa. Trans. Roy. Soc. S. Afr. 32(3) : 265­ Zool. Fr. 54: 212-213 . 273, 1 pl. MONOD, TH. 1937. Missions A. Gruvel dans le RAJ, B. SUNDARA. 1927. The littoral fauna of Canal de Suez. 1. Crusraces, Mem, Insr, Egypt. Krusadai Island in the Gulf of Manaar. Cir­ 34: 1-19. ripedia (Barnacles). Bull. Madras Govt, Mus., NEU, WOLFGANG. 1939. Bemerkungen tiber New Ser. Nat. Hist. Sect. 1(1) : 111-115, einige balanomorphe Cirripedien der Isran­ pIs. 12-14. buler Gewasser, Zool. Anz. 125(9/10) : 209­ ROGERS, F. 1. 1949. Three new subspecies of 219. Bal~nus amphitrite from California. J. Ent. NEWMAN, WILLIAM A. 1955, 1956. Personal Zool. 41 (2): 23-32 , pl. 1. communications. WEISS, CHARLES M. 1948. Observations on the NILSSON-CANTELL, C.-A. 1921. Cirripeden­ abnormal development and growth of bar­ Studien, Zur Kenntnis der Biologie, Anat­ nacles as related to surface toxicity. 'Ecology omie und Systematik dieser Gruppe. Zool. 29(1): 116-119. Bidr. Uppsala 7: 1-403, 3 pls. ZEVINA, G. B., and N . 1. TARASOV. 1957. Bar­ --- 1938. Cirripedes from the Indian Ocean nacles newly found in the soviet Black Sea. in the collection of the Indian Museum, Cal­ Trudy Sevastopol Biol. Srantsii 8: 341-346. cutta. Mem. Indian Mus. 13(1): 1-81, 3 pls. (In Russian.)