PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 115(4):868-877. 2002.

Taxonomy, anatomy, and histology of the Hned , Edwardsiella lineata (Verrill, 1873) (: : )

Marymegan Daly

Department of Biology, George Washington University, Washington, D.C. 20052, and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045

Abstract. —The lined sea anemone, Edwardsiella lineata (Verrill, 1873) is a common inhabitant of shallow waters of the North Atlantic from Cape Cod to Cape Hatteras. Its larval form, described as leidyi Verrill, 1898, is an endoparasite of the ctenophore Mnemiopsis leidyi and has been identified as the cause of sea-bather's itch. Although the biology and anatomy of the larval form are well documented, the internal anatomy and histology of adults has not been described. Confusion about the correct name for this species and about the identity of adults that produce the larvae have made identifying and

monitoring this species difficult. In this study, I describe the anatomy of adult

members of E. lineata, synthesize existing accounts of its development and life history, and detail the of E. lineata and the Edwardsiella.

Edwardsiella lineata is the valid name are undocumented. The lack of information for the species called Fagesia lineata (Ver- about internal morphology, and the absence rill, 1873) and E. leidyi Verrill, 1898. Ed- of clear differential diagnoses for E. lineata wardsia leidyi is the larval form of E. li- and other western North Atlantic species in neata (Crowell 1965, 1976). Although the the family Edwardsiidae make identifica- names of the larval and adult forms were tion and documentation of E. lineata pop- synonymized (Crowell 1965, 1976), the lar- ulations difficult. Because E. lineata has va continues to be cited as E. leidyi (e.g., been linked to sea-bather's itch in the North Gosner 1979, Kremer 1976, Meinkoth Atlantic (Freudenthal & Joseph 1992, 1995). Even publications that recognize the 1993), documenting the distribution of this synonymy fail to use the correct name for species is especially important. the species (e.g., Oviatt & Kremer 1977, Bumann & Puis 1996). Some of the no- Materials and Methods menclatural confusion is probably caused by Crowell's (1976) acknowledgment that For this study, I examined both live and the generic assignment given in the syn- preserved specimens of E. lineata. Live onymy was incorrect, as the genus Fagesia material was purchased from Marine Bio- was under revision at the time (see Manuel logical Laboratory (Woods Hole. Massa- 1981). chusetts); these specimens were collected Although the anatomy, ontogeny, and by SCUBA divers in Cape Cod Bay (E. ecology of the larval form are well char- Enos, pers. comm.). Coloration and external acterized (e.g., Mark 1884, Crowell 1976, anatomy were recorded prior to fixation. Crowell & Gates 1980, Bumann & Puis Freshly preserved material and museum 1996), and the external anatomy of the specimens from the Yale Peabody Museum adult is known (Verrill 1873), the internal (YPM 7642 Al and YPM 8802) and the anatomy and histology of adult E. lineata United States National Museum. Smithson- —

VOLUME 115, NUMBER 4 869 ian Institution (USNM 53363) were used Systematic Account for histological studies. Serial sections of Family Edwardsiidae Andres, 1883 paraffin-embedded preserved specimens Genus Edwardsiella Andres, 1883 were stained in Masson's Trichrome (Schreibman 1964). Edwardsiidae with column differentiated In the following discussion of anatomy, into capitulum, scapus, and scapulus, three "distal" refers to the oral disc and tenta- or more cycles of tentacles. Tentacles hex- cles, "proximal" to the aboral end. The amerously arranged, those of innermost cy- terms "dorsal" and "ventral" are used to cle longest. Capitulum with ridges; nema- describe the orientation of the retractor tocysts of capitulum concentrated on ridges. muscles and mesenteries (e.g., Carlgren Scapus bears periderm but lacks nemathy- 1921, Stephenson 1935, England 1987); bomes and tenaculi. Aboral end not differ- more recent descriptions of actiniarian anat- entiated into a physa. Ciliated tracts of fil- omy employ "sulcular" rather than dorsal, ament short, discontinuous (modified from and "sulcar," rather than ventral (e.g., Sta- Stephenson 1935, Carlgren 1949, Manuel chowitsch 1992). I use the older terminol- 1981). ogy to facilitate comparison with descrip- tions of other edwardsiids; in no way is the Edwardsiella lineata (Verrill, 1873) use of the terms dorsal or ventral meant to Figs. 1-3 homologize the longitudinal axis of actini- Edwardsia lineata Verrill, 1873:421.—Ver- arians with that of bilaterians. rill & Smith, 1874:445.—Parker, 1900: Cnidae preparations were made by 750.—Hargitt, 1912:248.—Miner, 1950: squashing approximately 1 .0 mm^ of tissue 198.—Williams, 1981:350. on a slide. Smears were prepared from ten- Edwardsia leidyi Verrill, 1898:496.—Park- tacle, actinopharynx, mesenterial filament, er, 1900:750.—Hargitt, 1912:248.—Min- and capitulum tissues from several individ- er, 1950:198.—Gosner, 1971:154.—Kre- uals. Tissue smears were examined at mer, 1976:213—Oviatt & Kremer, 1977: lOOOX magnification with differential in- 236.—Gosner, 1979:95.—Williams, terference optics. ScanPro measurement 1981:349. software (Jandel Scientific Software) was Edwardsia sp. Verrill & Smith, 1874: used to measure the length and width of 504.—Mark, 1884: pi. xii. undischarged cnidae capsules projected Edwardsiella lineata Pratt, 1935:151. onto a Summa Sketch digitizing pad tablet Freudenthal & Joseph, 1992:101; 1993: (Summagraphics). Cnidae nomenclature 542. follows Mariscal (1974). Edwardsia sp. Mark Carlgren, 1949:23. Cnidae presentation is modified from Fagesia lineata Carlgren, 1949:25.—Gos- Dunn et al. (1980). The capsule with the ner, 1971:153.—Crowell, 1976:247.— smallest (or greatest) width is generally not Crowell & Gates, 1980:139.—Sebens, the capsule with the smallest (or greatest) 1998:20.—Williams, 1981:350.—Shick, length. Summarizing the range using four 1991:253.—Meinkoth, 1995:352.— measurements, providing a length by width Weiss, 1995:4.14. range, as is typical in cnidarian taxonomy, is imprecise (Fautin 1988) and can imply Adult external morphology. —Body capsule sizes not observed. I present the translucent white, with pale tan lines indi- range by citing the size of the shortest, nar- cating mesenterial insertions. Mesenterial rowest, longest, and widest capsule sepa- compartments proximal to capitulum rately. The sizes of these capsules define a streaked with opaque white. Oral cone space that includes the size coordinates of prominent, opaque white. Tentacles margin- all observed cnidae. al, filiform, 18-36, in 2-3 cycles, 6 tenta- 870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. L External anatomy of preserved specimen of Echvardsiello lineata (USNM 53363). Scale bar = 3.0 mm. VOLUME 115, NUMBER 4 871 cles of innermost cycle longest. Tentacles mesenteries, forming capitular ridges. Ec- translucent white with opaque white flecks, toderm contains thin processes of mesoglea. each may bear longitudinal opaque mark on Cnidom. —Spirocysts, microbasic Z^-mas- oral side. tigophores, basitrichs, and microbasic

Column 10-30 mm long, 2-5 mm in di- /7-mastigophores (Fig. 3). See Table 1 for ameter, divisible into 2 distinct regions: distribution and size information. short, ridged, capitulum and long, smooth, Larval morphology. —Young larva pink- scapus covered by rough periderm of mu- ish-tan, ciliated. Shape very changeable, cus, detritus, and sand grains (Fig. 1). Peri- most commonly ovoid, can be spheroid or derm sheath-like, not attached to column by vermiform. Parasitic juvenile 2.0—3.0 mm suckers or tenaculi, extending from scapus long, approximately 1 mm in diameter, aboral contracted to end; specimens appear translucent pink, with 4-8 tentacles (Mark entirely enclosed by periderm. Aboral end 1884, Crowell 1976). Larva becomes short- not ampullaceous, instead tapering or er and thicker after leaving host (Mark slightly rounded, with rugae, but without 1884, Crowell 1976). cinclides or central invagination. Development, distribution, and natural Internal anatomy. —Actinopharynx tan, history. —Although position of each mesen- partially visible through body wall. Six tery in larval coelenteron prefigured by fur- pairs mesenteries in primary cycle: two row in endoderm (Mark 1884), mesenteries pairs of directives and four pairs, each of develop from evaginations of endoderm which has one member complete and fertile (Mark 1884). Mesenterial folds ciliated. (= macrocneme) and one member minute, Ventrolateral mesenterial pairs arise first, imperfect, sterile (= microcneme). Second- followed by dorsolateral, ventral and dorsal ary and tertiary cycles, when present, mi- mesenteries (Mark 1884). Mesenteries more crocnemous; all microcnemes restricted to developed distally than proximally. Muscles distal capitulum. Single ventral siphonog- develop after all pairs present (Mark 1884). lyph. Trilobed ciliated filaments short, dis- Found along North Atlantic coast of continuous, present in region proximal to North America, from Cape Cod, Massachu- actinopharynx and in reproductive tract; setts to Cape Hatteras, North Carolina. Lit- gonochoric, oviparous. toral to sublittoral. Adults commonly inhab- Retractor muscles large, circumscript, it rock crevices, Zostera root mats, or bore- convex (Fig. 2A, B), with closely spaced holes created by mollusks, worms, or branches of variable height on both sides of mesenterial lamella. Retractor branch sponges. Groups of E. lineata form mat-like height generally decreases toward coelen- "colonies," with individuals sharing peri- teron; branches both simple and ramified, derm. Adults asexually reproduce through with ramified branches toward edges of transverse fission (Crowell & Oates 1980). muscle (Fig. 2A, C). Parietal muscle strong, Juvenile E. lineata parasitize the cteno- circumscript, symmetrical, trefoil-shaped, phore Mnemiopsis leidyi (see Crowell 1965, with central mesogleal lamella thicker than 1976). Multiple larvae infest a single host. lateral branches. Fibers of parietal muscle Larvae enter the ctenophore through the ec- extend onto body wall, beyond profile of toderm and migrate through the canal sys- lower parietal branches. tem and mesoglea to the pharynx (Crowell Ectoderm of scapus columnar, glandular, 1976, Bumann & Puis 1996). The oral end with groups of basitrichs. Thickness of me- of the larva protrudes into the stomodeum soglea and ectoderm varies in capitulum: of host (Crowell 1976, Bumann & Puis mesoglea and ectoderm thickest in center of 1996); the larva removes food passing mesenterial compartment, tapering towards through host pharynx. 872 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ma>' ..

Fig. 2. Internal anatomy of Edwardsiella lineata. A. Cross section through column midline. B. Cross section through retractor muscle. C. Cross section through parietal muscle. Scale bars: A = 200 |xm; B, C = 50 jjim.

Discussion a type species. Carlgren (1892. 1893) de- scribed a new genus, Milne-Edwardsia (lat- Taxonomy. —The taxonomic history of er spelled Milneedwardsia), and altered the the genus Edwardsiella is extremely con- definitions of Edwardsia and Edwardsiella voluted. Andres (1883) erected Edward- to emphasize tentacle arrangement and the siella for edwardsiids with 20 or more ten- arrangement of nemathybomes (nemato- tacles. He listed the nominal species in no cyst-filled hollows in the ectoderm and me- apparent order and did not name or imply soglea). According to Carlgren (1892, VOLUME 115, NUMBER 4 873

Fig. 3. Representative cnidae from Edwardsiella lineata. (Refer to Table 1 for specific size information.) A. Spirocyst. B. Microbasic Zj-mastigophore. C. Basitrich. D. Microbasic p-mastigophore. E. Microbasic ii-mastig- ophore. F. Basitrich. Shape of these basitrichs varies from s-curve to straight. Scale bar = 20 iJim.

1893), Edwardsiella and Edwardsia were for Milneedwardsia, M. loveni is a nomen characterized by nemathybomes and octa- nudum. Edwardsia carnea Gosse, 1 856 was merously arrayed tentacles; Milneedward- the only valid species in Milneedwardsia at sia had no nemathybomes and had hexam- the time of description and is, therefore, its erously arrayed tentacles. Edwardsiella dif- type species by monotypy. fered from Edwardsia in tentacle number In the time between the description of and nemathybome arrangement (Carlgren Milneedwardsia and the discovery that Fa- 1893). Although Carlgren's (1892, 1893) gesia was preoccupied, the four species ini- definition of Edwardsiella was followed by tially assigned to Edwardsiella were trans- a few authors (e.g., McMurrich 1913, Ver- ferred to other genera or synonymized, and rill 1928), the name was generally not used. the name fell out of general use. In his Carlgren (1921) later submerged Edward- monograph on British anthozoans, Manuel siella within Edwardsia. (1981) resurrected Edwardsiella as a re- Milneedwardsia was found to be preoc- placement name for Fagesia, and named E. cupied, and the genus was renamed Fagesia carnea as the type of Edwardsiella Andres, (Delphy, 1938). In his catalog of Actiniaria, 1883. Although Manuel cited monotypy as Corallimorpharia, and Ptychodactiaria, the rationale for this action (because E. car- Carlgren (1949) listed Milne-Edwardsia nea was the last species left in Edwardsiel- loveni as the type of Fagesia. However, in la), it is actually type fixation due to sub- the earlier of Carlgren's two descriptions sequent elimination. Although subsequent 874 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

elimination is not sufficient for elevating a O t-~ q ^ ON 00 t^ m 00 \d 00 lo o\ ^ r^ VO ON (N species to type status, Manuel's declaration

^ ^ ^ ^ ^ ^ X) X) x> of E. carnea as the type, coupled with sub- r-; \q ON ON o\ ON >o q q sequent elimination, is 00 d NO 00 d NO 00 a valid type desig- m CnI o nation (Articles 6.9.1 and 6.9.4; ICZN o 1999). Because E. carnea is also the type of Milneedwardsia, Edwardsiella is the se- m ^ (N tN ON r^ 00 t' \o ^ r-^ CO vS NO NO cn! nior objective synonym for Milneedwardsia X3X>XlX)X)X)X)XiX) and Fagesia. Edwardsiella comprises five iO'-

r--;ND^_ON — r-;0qON00 E. loveni (Carlgren, 1893). The combination Edwardsiella lineata o^oooNcot^inO'* has been used three times. This combina- tion has not appeared in any general taxo- nomic, ecological, or experimental publi- cation. The first citation of the correct com- r-;NpoNON(N—

U 0) I) a variable number of imperfect, minute mi- U ;-H (-. o o o o o x; X3 x; x; x; crocnemes just below the oral disc. Ed- Cu Cli ex Q< o o o o o 00 00 60 00 wardsiella is the only genus of Edwardsi- idae whose members generally do not bur- 6 E £ row; members of this genus typically live 3 ^ < -a-^ u ^ _o ii o in crevices, in dead coral skeletons, or in ^ Crt C/3 V) — f= ca w M -f^ f? XI .a X> y XI XI x> y mats on the substrate. Edwardsiella lineata n O h O ^ o o o ^- .tli U U U .tl .b! t->

1869, vectensis Stephenson, in squash preparations, making it less likely 1935, and Drillactis pallida (Verrill, 1880). that they will be recognized and recorded. Edwardsia elegans normally has 16 tenta- The life history of Edwardsiella lineata cles; Edwardsiella lineata has 18 or more. has two unusual features: parasitism and These two species can be further distin- transverse fission. Parasitism on medusae is guished by differences in column morphol- seen in several burrowing anemones. Spe- ogy. Edwardsia elegans has an unridged ca- cies in the genus Peachia (family Haloclav- pitulum, conspicuous nemathybomes on the idae) parasitize hydromedusae (Nyholm middle and lower column, and a naked, bul- 1949, Spaulding 1972); in fact, Carigren bous aboral end; Edwardsiella lineata has (1949) regarded this as a diagnostic feature a ridged capitulum, a smooth mid and lower of Peachia. Edwardsiella lineata is the only column, and a tapering, periderm-covered edwardsiid known to parasitize medusae, aboral end. Nematostella vectensis and E. although larvae of E. carnea may be asso- lineata differ in tentacle number, size, and ciated with the ctenophore Bolinopsis (see color: A^. vectensis has 16 tentacles and is Stephenson 1935). generally small (<10 mm), opalescent Transverse fission is reported in only a white or translucent. Furthermore, the coe- few actiniarians, including the edwardsiid Nematostella vectensis (Crowell Oates lenteron of A^. vectensis contains nemato- & somes, ciliated spherical nematocyst bodies 1980, Shick 1991, Hand & Uhlinger 1992). of no other are (Stephenson 1935). Drillactis pallida, Members edwardsiid species known to undergo regular asexual repro- which has been reported only once (the duction. Individuals of both Edwardsiella specimens cited in Verrill 1928 and Carl- lineata and N. vectensis often live in dense gren 1950 are from the type collection, de- aggregations (pers. obs., Crowell & Oates scribed in Verrill 1880), is similar in size 1980, Hand & Uhlinger 1994), presumably and external appearance to E. lineata. How- the result of asexual reproduction. The dis- ever, in D. pallida, the inner tentacles are persal ability afforded E. lineata larvae by shorter than the outer tentacles and the re- the association with ctenophores may be es- tractor muscles are smaller and less com- pecially important because of the tendency plex than those of E. lineata. Furthermore, of this species to form dense mats that cov- D. pallida lacks nematocysts on the capit- er locally available substrate. ulum, and possesses nematosomes (pers. Edwardsiella lineata has been suggested obs.). as a possible biocontrol agent for Mne- Unusual aspects of the species. The — miopsis leidyi introduced into the Black mesenterial filaments of E. lineata contain Sea. Edwardsiella lineata compromises long, thin basitrichs (Fig. 3F, called pencilli growth and survival of M. leidyi and may B, by den Hartog 1987). have a long-term effect on the population Although these nematocysts are recorded size of M. leidyi (Crowell 1976, Kremer from other actiniarians (e.g.. Hand 1955, 1976, Bumann & Puis 1996). However, the den Hartog 1987, Belem & Pinto 1990, life history of E. lineata suggests that an Riemann-Ziimeck & Gallardo 1990), they introduction of this species may have un- have not been reported in Edwardsiidae, desirable consequences. The larvae cause and their distribution in the Actiniaria is sea-bather's eruption, an unpleasant der- poorly understood. Because these basitrichs matitis that may render beaches temporarily are more rare than other types of cnidae in unfit for recreational use (Freudenthal & Jo- the filament, and are unlike other nemato- seph 1993). Predators of adult E. lineata are cysts in general shape, they may have been unknown, as is the role this species plays overlooked in other species. Furthermore, in the ecology of benthic communities. Be- these nematocysts are often broken or bent cause E. lineata is capable of asexual re- — —

876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON production and tends to occur in high num- description of a new genus and species from Peru. Archiv fiir Zoologie 71:1—38. bers when present, this species could itself — Crowell, S. 1965. An anemone parasitic as a larva in become invasive, compounding the prob- a comb jelly [Abstract].—American Zoologist in lem of introduced species the Black Sea, 5:645. it. rather than solving . 1976. An edwardsiid larva parasitic in Mne- miopsis. Pp. 247-250 in G. O. Mackie, ed., Acknowledgments Coelenterate Ecology and Behavior. Plenum Press, New York, 744 pp.

, & S. Oates. 1980. Metamorphosis and repro- This research was funded by grants NSF duction by transverse fission in an edwardsiid DEB 9801370 to D. L. Lipscomb and M. anemone. Pp. 139-142 in P. Tardent and R. Tar- Daly and NSF-PEET DEB 9978106 to D. dent, eds.. Development and Cellular Biology

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