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Home , Actiniidae, Adamsia, Adamsia palliata, Calliactis, Calliactis parasitica, Calliactis tricolor

Pacific Science (1980), vol. 34, no. 4 © 1981 by The University Press of Hawaii. All rights reserved

Stylohates: A Shell-Forming (, , ) 1

DAPHNE FAUTIN DUNN,2 DENNIS M. DEVANEY,3 and BARRY ROTH 4

ABSTRACT: and cnidae distinguish two of deep-sea ac­ tinians that produce coiled, chitinous shells inhabited by hermit of the Parapagurus. The actinian type species, Stylobates aeneus, first assigned to the , occurs around Hawaii and Guam with P. dofleini. Stylobates cancrisocia, originally described as Isadamsia cancrisocia, occurs off east Africa with P. trispinosus.

MANY MEMBERS OF THE ORDER Actiniaria pedal disk secretes a chitinous cuticle over attach obligately or facultatively to gas­ the small mollusk shell which the pagurid tropod shells inhabited by hermit crabs. had initially occupied and to which the small Some of these partnerships seem to be actinian had first attached, often extending strictly phoretic, the normally sedentary sea the cuticular material beyond the lip of the anemone being transported by the motile shell (Balss 1924, Faurot 1910, Gosse 1858). hermit (Ross 1971, 1974b). The re­ This arrangement affords the crab mainly lationships between other species pairs are mechanical protection (Ross 1971). mutualistic, the anemone gaining motility Carlgren (I928a) described as a new genus while protecting its associate from and species Isadamsia cancrisocia (family (Balasch and Mengual 1974; Hand 1975; Actiniidae), an actinian attached to a shell McLean and Mariscal 1973; Ross 1971, occupied by a , from four speci­ 1974b; Ross and von Boletsky 1979). As the mens collected by the Deutschen Tiefsee­ grows, it must move to increas­ Expedition (1898-1899) at 818 m in the ingly larger shells. Some species of hermits Pemba Channel off east Africa. The generic detach their sea from the smaller name recognizes two points of superficial shells and place them on the larger, whereas similarity with : (1) the in other associations, the anemones transfer is oriented on the shell so that its is themselves from shell to shell (Ross 1974a). beneath that of the crustacean; (2) the coiled The crustacean Pagurus prideauxi Leach shell, called by Carlgren (I928a, 1928b) a need never change shells when in association carcinoecium, is almost entirely produced by with the cloak anemone, the actinian. Carlgren (I928a, 1928b) did not Bohadsch (family ), which note that a shell-making actinian had been wraps around the hermit's abdomen and reported previously. Dall (1903: 61) des­ grows as the crab does. In the process, the cribed what he thought was a new genus and species of trochid gastropod, Stylobates aeneus, based on shells "of a flexible, horny I Manuscript accepted 12 August 1980. consistency ... " "occupied by hermit crabs 2 California Academy of Sciences, Department of Zoology, Golden Gate Park, San Francisco, and completely covered by the extended basal California 94118. membrane of a large " that had been 3 Bernice P. Bishop Museum, Division of Invertebrate collected at 220-436 fm (402-797 m) by the Zoology, Post Office Box 19000-A, Honolulu, Hawaii U.S. Fish Commission steamer Albatross in 96819. waters of the Hawaiian Islands. Sixteen 4 California Academy of Sciences, Department of Invertebrate Zoology, Golden Gate Park, San Francisco, years later, having seen a collection of simi­ California 94118. lar but irregularly shaped shells, Dall (1919: 379 380 PACIFIC SCIENCE, Volume 34, October 1980

80) published a note identifying them and and accurately characterized it as having S. aeneus as "secretions from the bases of smooth column, circumscribed endodermal the Actinias." Neave (1940) lists Stylabates as sphincter muscle, short , several a mollusk, although the catalog of Dall's cycles of perfect mesenteries of which the names (Boss, Rosewater, and Ruhoff 1968) first three orders are sterile, and weak, diffuse correctly includes it among the coelenterates. mesenterial retractor muscles. The name Stylabates, however, has priority over Isadamsia. The type species, by monotypy, is MATERIALS AND METHODS S. aeneus Dall 1903 (Boss, Rosewater, and Ruhoff 1968), described from Albatrass sta­ We have examined numerous carcinoecia tion 3893, in the Kaiwi Channel between formed by Stylabates from Hawaii and Molokai and Oahu, Hawaiian Islands, based Guam, most containing hermit crabs and on the shell. Although the name was based covered by actinians. Material from Guam is on "the work of an ," since it was in the collection of the Department of published prior to 1931, it is available for the Invertebrate Zoology, California Academy actinian [Article 16 (a) (viii), International of Sciences, catalog 011027-011030; Hawai­ Code ofZoological Nomenclature]. ian specimens are in the Division of Inverte­ brate Zoology, Bernice P. Bishop Museum, accession 1971.197 and 1979.529, the latter received from the National Marine Fisheries Service (Southwest Center), Honolulu. We have examined the holotype of Stylabates aeneus, U.S. National Museum molluscan lot 795117, which is represented only by the shell although it was collected with actinian and crustacean (Dall 1903). We were able to locate only one of the four specimens of Isadamsia cancrisacia from Deutschen Tiefsee-Expedition station 246. There is no indication in the catalog nor on the label that this specimen (Museum fur Naturkunde, Berlin, GDR, catalog 6957) is the holotype. A piece has been removed from the margin, apparently for histological study; the apex has been cracked; and it is illustrated in the original description (Carlgren 1928a: plate 2, fig. 9), so it certainly was studied by Carlgren. Histological sections (8 /lm thick) of speci­ mens from Hawaii and Guam were stained with hematoxylin and eosin. One of us (DFD) measured cnidae of seven Pacific specimens as well as of the specimen of Isadamsia cancrisacia from smash prepar­ ations. Not all tissues of every non-type specimen were examined. FIGURE I. Two views of Sly/obates aeneus and Parapagurus dofleini (specimen 011029, California Academy of Sciences). Shell aperture 35 mm greatest diameter. a, apex of shell at upper left; b, mesenterial RESULTS filaments visible through gaping mouth, typical of pre­ served specimens; longitudinal ridges due to mesenterial All the actinians belong to a single actiniid insertions pulling on thin column wall. (Photograph by genus. Carlgren (1949) called it Isadamsia, Lloyd E. Ullberg.) A Shell-Forming Sea Anemone-DuNN, DEVANEY, AND ROTH 381

FIGURE 2. Photomicrograph of column (a) and pedal disk (b) ectoderm of Sty/obates aeneus (specimen 011030, California Academy of Sciences).

The word stylabates, which may be trans­ Troy? The old master was not above this lated roughly as "a walker on stilts," is per­ sort of thing.] haps even more appropriate for the actinian The actinian is oriented with its oral disk than it was for DaII's presumed mollusk. behind and below the aperture of the car­ [And in naming the species aeneus (Latin, cinoecium, beside the umbilicus (Figure I). meaning "made ofcopper or bronze"), could The pedal disk completely covers the shell in Dall also have had in mind Aeneas, who life, although in preserved specimens it is carried his father on his back out of fallen drawn back from the lip (Figure I). Typical 382 PACIFIC SCIENCE, Volume 34, October 1980

sized Sty/obates is about 50 mm long on one side (at right in Figure Ib), extending almost completely around the shell, but only about 6 mm long on the opposite side (at left in Figure Ib). Thus, pedal disk and column wall are parallel over the shell, which they invest like a membrane. The animal is bila­ terally symmetrical, with its directive axis perpendicular to the median oral-aboral plane passing through the shortest and long­ est spans of the column (the directive axis is nearly vertical in Figure Ib). The column of the anemone is pale pinkish-tan to colorless, but the pedal disk is typically either deep brown or lighter in hue than the column. This gross difference is apparently attribu­ table to histological differences between column and pedal disk ectoderm (Figure 2). Cells of the former are loose and vacuolated; those of the latter are tightly packed, high columnar, and glandular. We infer, as did Dall (1919) and Carlgren (1928a, 1928b), that the pedal disk secretes the carcinoecium, and this histology supports that conclusion. The Sty/obates shell was well described by Dall (1903); all that we examined are tro­ choid with dextral coiling, three to four whorls, a broadly ovate aperture, and a deep umbilicus (Figure 3; Carlgren 1928b: figs. 8-10). They are brassy in color, opaque, of a papery texture, and readily dented or torn. Growth rings (Figures Ib, 3) concentric with the lip [also noted by Dall (1903) and Carlgren (1928b)] are typically the only sculpture, but the apical surface of one is papillose (Figure 3). The anemones from Hawaii and Guam belong to a single species, judging from sphincter form and cnidae. Their shells being of similar size and nature to DaB's holotype, we refer them to Sty/obates aeneus. The FIGURE 3. Three views of anemone shell (car­ single east African specimen that we ex­ cinoecium) made by Sly/abates aeneus (ex Bishop amined, illustrated in Carlgren's (1928a: Museum lot 1979.529). Note growth rings, papillose plate 2, fig. 9) original description of "protoconch" at angle to carcinoecium, umbilicus. Shell Isadamsia cancrisocia, represents a distinct aperture 30 x 38 mm; greatest width of shell 60 mm. (Photograph by Lloyd E. Ullberg.) species, the second of the genus, S. cancri­ socia. Cnidae of most categories are smaller in the latter than in the former species (Table actinians are cylindrical in shape, with oral 1), the difference being greatest in column and pedal disks perpendicular to the column basitrichs. It is unlikely that this is related to wall. However, the column of an average- the only slightly smaller size of the east A Shell-Forming Sea Anemone-DuNN, DEVANEY, AND ROTH 383

c

D

A B H

10 ~m

E F G

FIGURE 4. Cnidae of Slylobales {tenet/s. (Drawings by Steven Sechovec.) 384 PACIFIC SCIENCE, Volume 34, October 1980

TABLE I

CNIDAE OF Sty/obates aeneus AND S. cancrisocia

S. aeneus S. canerisocia

Tentacle basitrichs (A) 27.9-36.1 x 3.1-3.9, N = 43 27.1-30.3 x 2.5-3.3, N = II spirocysts (B) (25.4) 28.7-44.3 x 2.9-4.1 (4.9), N = 40 23.8-39.4 x 2.7-3.3, N = 10 Actinopharynx basitrichs (C) 17.2-21.3 x 2.5-3.1, N = 9 Not seen basitrichs (D) (26.2) 29.5-37.7 x 2.9-4.1, N = 38 27.1-31.2 x 2.5-3.3, N = 10 Mesenterial filaments microbasic p-mastigophores (E) 21.3-29.5 x 3.9-5.7, N = 27 18.9-23.0 x 4.1-5.7, N = II basitrichs (F) 29.5-36.1 x 2.5-3.9, N = 21 (25.4) 27.9-32.8 x 4.1-5.5, N = 14 basitrichs (G) 15.6-23.0 x 2.1-3.3, N= 58 14.8-18.0 (20.5) x 2.1-2.5, N = 8 Column basitrichs (H) 29.5-36.1 x 2.9-3.5, N = 24 23.0-28.7 x 2.7-3.3, N = 12

NOTE: N = number of cnidae measured; measurements in parentheses are for single cnidae falling well outside the usual range. Measurements are in micrometers. Letters refer to illustrations of cnidae from S. aeneus in Figure 4.

African specimen, and there are other dif­ Four of 21 Sty/obates aeneus shells from ferences as well. First, like us, Carlgren Hawaii that we dissected had remains of (1928a) did not find small basitrichs in the calcareous material in the apical region: two actinopharynx of S. cancrisocia. [The sizes of these were gastropods, and two were por­ ofcnidae measured by DFD do not precisely tions of serpulid worm tubes. In all cases, correspond with figures given by Carlgren the carcinoecium is of typical trochoid (1928a); he did not specify the number of shape. The gastropod shell underlying the and cnidae upon which his figures apex of the one specimen of S. cancrisocia were based.] Second, sphincter muscles of that we examined is relatively high spired, several specimens of S. aeneus that we ex­ producing a sharp apex on the anemone amined are similar (Figure 5) in that they shell (Figure 6; Carlgren I928a: plate 2, have a stronger mesogleal axis, and the secon­ fig. 9). Beyond this "protoconch," the car­ dary mesogleal lamellae are consistently cinoecium is trochoid. more pinnately arrayed than those of S. Large numbers of carcinoecia from cancrisocia illustrated by Carlgren (1928a), Sty/obates, most containing hermit crabs in which the secondary mesogleal lamellae and covered by actinians, are taken in radiate from a short primary axis. Third, the Hawaii and Guam by dredging and in fish shell of S. cancrisocia has a keel on the traps. The depth range of 93 trapped speci­ penultimate whorl just above the aperture mens from Guam was 250-500 m; the sub­ (Figure 6, arrow), but that may be only an stratum in the area of capture was primarily individual peculiarity. or entirely (except for one site with 57 Among the Hawaiian material are two percent silt and 43 percent sand) (L. G. very small trochoid shells containing Eldredge, personal communication, May specimens of Parapagurus, one of which is 1980). Depth and substratum correspond well ovigerous, that could not be specifically with what is known of the habitat in Hawaii. identified. The actinian on each does not completely cover the shell but is located beside the umbilicus on the shell's underside, DISCUSSION oriented as is Sty/obates, and the few nema­ tocysts we measured correspond in size to The evidence at hand suggests that those of S. aeneus. In both, a thin, narrow Sty/abates settles on a shell either before or sheet of brittle, translucent, yellow-brown after a hermit crab has occupied it. The chitinous material is attached to the lip of pedal disk of the actinian secretes a chitinous the shell and extends over the paguroid. membrane that gradually coats the shell. A Shell-Forming Sea Anemone-DuNN, DEVANEY, AND ROTH 385

FIGURE 5. Cross section of circumscribed endodermal sphincter muscle of Sty/abates aeneus (specimen 011030, California Academy of Sciences). (Drawing by Steven Sechovec.)

Sculpture on the original shell may be em­ Our observations indicate that the shape bossed into the apex of the anemone shell of the original hermit shell exerts little if any (Figure 3). We agree with Carlgren's (1928a) effect on the ultimate shape of the trochoid assumption that in cases where a large car­ carcinoecium. The anatomy of the actinian, cinoecium lacks a calcareous shell beneath especially the curvature of its pedal disk and the apex, the shell has dissolved. The solu­ the difference in column length from one side bility of calcium carbonate increases with to the other, may dictate the basic shell ocean depth (Correns 1955: 377). Species of form. The fact that the axis of the sculptured the sea anemone Paracalliactis living at "protoconch" of the shell illustrated in depths of several thousand meters com­ Figure 3 is set at an angle to the axis of the pletely cover the gastropod shells to which anemone shell supports this hypothesis. It is they are attached (Doumenc 1975), unlike also possible that the hermit crab, in one shallow-water actinians which usually cover way or another, mediates shell form. The only a portion of the shell's surface. We hermit crab's larger cheliped, which closes favor a functional interpretation of this cor­ the aperture of the shell much like a gas­ relation, viz. that coverage of a shell impedes tropod operculum, may serve as a template its dissolution at depth and thus is advan­ for the configuration of the growing aper­ tageous for both symbionts. tural edge. Furthermore, all Stylobates shells 386 PACIFIC SCIENCE, Volume 34, October 1980

shells. It is also possible that the shapes of some shells might totally prevent extension by Styfobates [e.g., that illustrated by Carlgren (1928a: plate 2, fig. 6; 1928b: fig. la)]. They may also be products of more than one actinian per shell, a situation that has been reported (Dall 1919, Miyake 1978) but that we have not seen. It seems highly improbable that two or three sea anemones could coordinate efforts to produce a single shell of such precise geometry. The pedal disks of some actinians other than Adamsia and Styfobates also produce chitinous secretions. The hormathiid Para­ is defined in part by its ability to form a cuticle that extends the shell to which the actinian attaches, and members of the monogeneric pelagic family Minyadidae FIGURE 6. Sty/abates cancrisacia and Parapagurus form a basal, chitinous, porous mass that trispinasus (specimen 6957, Museum fUr Naturkunde). acts as a float (Carlgren 1949). Arrow indicates keel. (Photograph by Lloyd E. Ullberg.) Another unanswered question is whether a single individual paguroid lives with a single actinian throughout the latter's life, or wheth­ that we have seen are dextrally coiled. We er a succession of individual paguroids of know of nothing intrinsic to the anemone's the same or different species occupy one anatomy to require preferential coiling in shell. Unless it were displaced by a more one direction, but the symbiotic hermit crabs aggressive individual, as occurs in typical are asymmetric and coiled to the right. hermit crabs (Hazlett 1966, 1970), there Perhaps the paguroid's abdomen is the should be no reason for the crustacean to guide, as it evidently is in Adamsia, where leave a shell it had not outgrown; this as­ the edges of the ovoid pedal disk wrap sumes that the actinian can extend the car­ around the crustacean, its symmetrically cinoecium fast enough to keep pace with the elongate edges meeting along the dorsal mid­ growth of the paguroid. Moreover, the del­ line, following the curve of the crab's abdo­ icacy of the shells, which might be torn men (Faurot 1910). during exit and entry, suggests that transfers In terms of Raup's (1966) geometric analy­ are rare or absent. A defihitive answer will sis of shell coiling, the Styfobates shell falls require observation ofliving animals through within the gastropod field, having low whorl time. expansion rate (w), moderate translation rate Shells of Styfobates aeneus that we ex­ (t), and little distahce between generating amined are inhabited by Parapagurus dof­ curve and axis (d). This correspondence is feini Balss (identified by M. Takeda). The doubly intriguing because Styfobates lays crab in the shell of S. canerisocia is the down its shell from an external position, smaller P. trispinosus Balss. [Balss (1912) whereas mollusks secrete their shells while listed three female specimens of Parapagurus ensconced within. The Styfobates umbilicus arcuatus var. trispinosa (but no actinians) is apparently produced by a portion of the from station 246. Saint Laurent (1972) anemone's pedal disk protruding inside it. recognized this variety as a distinct species, Perhaps the irregular shells that convinced P. trispinosus. Carlgren (1928a) said that two Dall (1919) that he was not dealing with a of the four actinians from station 246 lacked gastropod might be products of anemones ; he called them P. armatus var. that could not produce an appropriate gener­ trispinosus, clearly a fapsus.] This suggests ating curve from positions on nontrochoid species specificity between paguroids and ac- A Shell-Forming Sea Anemone-DuNN, DEVANEY, AND ROTH 387 tinians. It is possible that a single shell might Valuable assistance in preparing this paper be sequentially occupied by specimens of P. was provided by Steven Sechovec (draw­ trispinosus and P. dofleini, which are broadly ings), Lloyd E. Ullberg (photographs), sympatric (although P. trispinosus is absent Denise Ross (histological preparations), and in Hawaii; Saint Laurent 1972). However, Phoebe L. Grigg (typing), all of the Cali­ anatomy amd physiology of these anemones fornia Academy of Sciences. We also thank are so specialized for a shell-making way of M. Takeda, National Museum of Science, life that growth rate may well be adjusted to Tokyo, for locating Dall's holotype; Joseph that of a specific crustacean associate. Rosewater, U.S. National Museum, for Other published records allude to addi­ loaning us that specimen; and D. H. H. tional symbioses of this sort. Japanese speci­ Kuhlmann, Museum fur Naturkunde, for mens of Parapagurus dofleini reportedly live loan of Carlgren's east African specimen. J. in shells "usually encrusted with one or two Bailey-Brock, C. Hand, and P. A. McLaugh­ sea anemones" (Miyake 1978: 76), but the lin also contributed to the preparation of nature of the shells is not given, and there­ this paper. fore they may be gastropod shells. Japanese P. diogenes are said to "inhabit in thin­ shelled which are usually encrusted with one or two sea anemones. They were LITERATURE CITED occasionally found in a large thin cuticular snail and a small true snail was embedded in BALASCH, J., and V. MENGUAL. 1974. The the apex" (Miyake 1978: 74). If this were a behavior of Dardanus arrosor in associa­ third species of Stylobates, the hypothesis of tion with in artificial specificity between species of Parapagurus habitat. Mar. Behav. Physiol. 2(3): 251­ and species of Stylobates would be streng­ 260. thened. An actinian identified as Isadamsia BALss, H. 1912. Paguriden. Wiss. Ergeb. cancrisocia was reported from about 3500 m Deutsch. Tiefsee-Exp. 20(2): 85-122. in the North Atlantic (Doumenc 1975). It is 1924. Uber Anpassungen und most unlikely that this identification is cor­ Symbiose der Paguriden: eine zusammen­ rect, but it probably means that a shell­ fassende Ubersicht. Z. Morph. Okol. Tiere forming actinian occurs there. 1(4): 752-792. Small specimens of these symbiotic species Boss, K. J., J. ROSEWATER, and F. H. of Parapagurus inhabit gastropod and other RUHOFF. 1968. The zoological taxa of calcareous shells, and larger ones are also William Healy Dall. Bull. U.S. Nat. Mus. known to live in them (Miyake 1978). 287: 1-427. Presumably, the relationship with actinians CARLGREN, O. 1928a. Actiniaria der Deut­ is facultative for the crustacean. We also schen Tiefsee-Expedition. Wiss. Ergeb. believe that the carcinoecium must be oc­ Deutsch. Tiefsee-Exp. 22: 123-266. cupied continually by a hermit crab for the 1928b. Zur Symbiose zwischen actinian to survive. In the absence of its Actinien und Paguriden. Z. Morph. Okol. symbiont, the sea anemone would probably Tiere 12(1-2): 165-173. lack for food, and the light shell might ---. 1949. A survey of the Ptychodac­ tumble about and thereby abrade the tiaria, and Actiniaria. anemone. Thus, it is likely that the relation­ K. Svensk. Vetensk. Handl. Ser. 1(1): ship is obligate for the actinian. 1-121. CORRENS, C. W. 1955. Pelagic sediments of the North Atlantic Ocean. Pages 373­ ACKNOWLEDGMENTS 395 in P. D. Trask, ed. Recent marine sediments-a symposium. Soc. Econ. We are grateful to L. G. Eldredge, Marine Paleontol. Mineral. Spec. Publ. No.4. Laboratory, University of Guam, for speci­ DALL, W. H. 1903. A new genus of mens and data on animals from Guam. . Nautilus 17(6): 61-62. 388 PACIFIC SCIENCE, Volume 34, October 1980

1919. Stylobates, a warnmg. Nau­ gicus. Vo!. 4. Zoological Society, London. tilus 32(3) :79-80. 758 pp. DOUMENC, D. 1975. Actinies bathyales et RAUP, D. 1966. Geometric analysis of shell abyssales de I'Ocean Atlantique Nord. coiling: General problems. J. Paleonto!. Familles des Hormathiidae (genres Para­ 40: 1178-1190. calliactis et Phelliactis) et des Actinosto­ Ross, D. M. 1971. Protection of hermit lidae (genres Actinoscyphia et Sicyonis). crabs (Dardanus spp.) from by Bull. Mus. Nat. Hist. Nat. Ser. 3, Zoo!' commensal sea anemones (Calliactis 197(287): 157-204. spp.). Nature 230: 401-402. FAUROT, L. 1910. Etude sur les associations ---. 1974a. Behavior in associ­ entre les pagures et les actinies. Arch. ations and interactions with other Zoo!' Exp. Gen. Ser. 5(5): 421-486. animals. Pages 281-312 in L. Muscatine GOSSE, P. H. 1858. On the nature of the sub­ and H. M. Lenhoff, eds. Coelenterate basal membrane of Adamsia palliata. Ann. : Reviews and new perspectives. Mag. Nat. Hist. Ser. 3(2): 107-108. Academic Press, New York. HAND, C. 1975. Behaviour of some New ---. 1974b. Evolutionary aspects of as­ Zealand sea anemones and their mollus­ sociations between crabs and sea anem­ can and crustacean hosts. N. Z. J. Mar. ones. Pages 111-125 in W. B. Vernberg, Freshw. Res. 9(4): 509-527. ed. in the sea. University of HAZLETT, B. A. 1966. Social behavior of the South Carolina Press, Columbia, S. C. Paguridae and Diogenidae of Curac;ao. Ross, D. M., and S. VON BOLETSKY. 1979. Stud. Fauna Curac;ao and other Carib. lsI. The association between the pagurid 23(88): 1-143. Dardanus arrosor and the actinian Calli­ --. 1970. Interspecific shell fighting in actis parasitica. Recovery of activity in three sympatric species of hermit crabs in "inactive" D. arrosor in the presence of Hawaii. Pac. Sci. 24(4): 472-482. cephalopods. Mar. Behav. Physio!. 6: McLEAN, R. B., and R. N. MARISCAL. 1973. 175-184. Protection of a hermit crab by its sym­ SAINT LAURENT, M. DE. 1972. Sur la famille biotic sea anemone . des Smith, 1882. Descrip­ Experientia 29(1) : 128-130. tion de Typhlopagurus foresti gen nov., MIYAKE, S. 1978. The crustacean sp. nov., et de quinze especes ou sous­ of Sagami Bay. Biological Laboratory, especes nouvelles de Parapagurus Smith Imperial Household, Tokyo. 161 pp. (Crustacea, ). Bijd. Dierkunde NEAVE, S. A., ed. 1940. Nomenclator zoolo- 42(2): 97-123.