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Home , Coeloplana, Ctenophora, Platyctenida, Rottnest Island

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 127(2):423–436. 2014. waltoni, a new species of minute benthic ctenophore (: ) from south Florida

Peter W. Glynn*, Frederick M. Bayer†, and D. Abigail Renegar (PWG) Department of and Ecology, Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149, U.S.A., e-mail: [email protected]; (FMB) Department of , National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013, U.S.A.; (DAR) National Coral Institute, Nova Southeastern University Oceanographic Center, 8000 North Drive, Dania Beach, Florida 33004, U.S.A., e-mail: [email protected]

Abstract.—A minute and abundant species of benthic ctenophore is described, resurrected from an unpublished manuscript of Frederick M. Bayer (deceased). It is an ectosymbiont of octocorals in south Florida. Members of the ctenophoran Platyctenida were unknown from American waters until 1945, when specimens from Miami, Florida were reported by F. G. Walton Smith and referred to the Coeloplana. That species subsequently was found in Jamaica, and Rankin (1956) described it under the new generic name Vallicula. A second species of creeping ctenophore was found in the 1960s and rediscovered in 2011, living as an ectosymbiont on octocorals in the coastal waters near Miami. This species belongs to the original genus Coeloplana and is here described as Coeloplana waltoni, new species. This description is based on specimens examined by F. M. Bayer in the 1960s and additional material from collections obtained in 2012–2013. Critical differences between C. waltoni and Vallicula multiformis are noted, based on recently collected specimens of the latter in Biscayne Bay, Miami, Florida.

Keywords: ectosymbiont, Florida, platyctenid ctenophore, subtropical western Atlantic

In 1880, the Russian zoologist A. wellastocreepinthemannerof Kowalevsky reported the discovery of a Coeloplana; Gastrodes Korotneff (1888) remarkable planariform ctenophore from (because of its double homonymy, i.e., the Red Sea, to which he gave the name with two previously named taxa bearing Coeloplana metschnikowii.Intheensuing this name, it was later renamed Gastra 70 yr, additional species of these peculiar Stechow, 1921, see also Stechow 1923), creeping ctenophores were discovered in an internal parasite in which is so various parts of the Indo-Pacific region, modified that it was at first not recog- in particular, Japan and Southeast Asia, nized as a ctenophore; Tjalfiella, a sessile and several related genera were de- form living upon the pennatulid Umbel- scribed. These are Ctenoplana Korotneff lula off Greenland and first reported by (1886), which retains vestiges of ctenes in Mortensen (1910); Planoctena,from the adult stage and is able to swim as Southeast Asian waters, a genus extreme- Ctenoplana * Corresponding author. ly close to , established by † Deceased. Dawydoff (1936); the remarkable Lyroc- 424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON teis from Japan, a giant among the Materials and Methods Platyctenida, first found in 1896 but formally named by Komai (1941) who This description of a new species of recognized its true affinities; and, finally, Coeloplana isbasedontwosetsof the curious genus Savangia, which has six collections, the first studied by F. M. tall, tuberculate aboral papillae and Bayer in the 1960s, and a second series of resembles a mollusk, de- collections obtained in 2012 and 2013. The scribed from the China Sea by Dawydoff early collection examined in this study (USNM 52454) contains several specimens (1950). It is noted that Gastrodes and adhering to octocoral branches from Sol- Gastra are not listed in Mills (1998); dier Key, upper Florida Keys. This collec- however, in Mills’ updated log (2014) tion was made by F. M. Bayer on 10 Gastrodes komai Dawydoff, 1937 is listed October 1960 and was fixed in Bouin’s as a junior synonym of Lampea komai ; it is in excellent condition. Recent (Dawydoff, 1937). collections (2012–2013) have followed the No representatives of the Platyctenida conditions and provisions outlined in the were known from American waters until Special Activity License (No. SAL-12- 1945, when some translucent, creeping, 1365-SR), Florida Fish and Wildlife Con- flatworm-like collected from sea- servation Commission. water aquaria at Miami by W. H. Branch clippings from various octocoral Sutcliffe, Jr., were recognized by F. G. species inhabiting the upper Florida Keys Walton Smith (1945), Marine Laboratory and off Dania Beach, Florida were cut (now Rosenstiel School of Marine and with scissors underwater to lengths of 10– Atmospheric Science), University of Mi- 20 mm. Bayer’s (1961) monograph was ami, and reported as a species of Coelo- used to identify octocoral species. Sam- plana. These metazoans proved to be pling was confined to the higher peripheral abundant in Biscayne Bay, living on branches of colonies in order to minimize and colonies of the arborescent ctenos- injury. Each branch was sealed in a water- tome bryozoan Zoobotryon.Regrettably, filled polypropylene bag and transferred to no opportunity to study them in detail a 20-liter bucket. The bag were presented itself, and the species remained opened to facilitate circulation. The undescribed systematically until 1956 branches were then transported to the when J. J. Rankin (1951, 1956) published laboratory under continuous aeration an account of some specimens found in where they were examined microscopical- Jamaica. For these, she established the ly. Live animals collected off Dania Beach new genus Vallicula, based upon the (22 July 2013) were observed and photo- shape of the tentacular sheaths, nature graphed with a Leica MZFLIII dissecting of the aboral papillae, presence of oral microscope fitted with a Carl Zeiss camera grooves, and position of the body when attachment. Three video clips showing full detached from the substrate and floating body views of the ctenophore (MVI_0780), freely. There is no doubt of the identity of close up views of symbiotic flatworms the Jamaican animals when compared (MVI_0784), and a clip of a specimen with those reported from Florida (Wal- moving slowly over an octocoral host ton-Smith 1945). The new species de- (MVI_0791) are available as supplementa- scribed herein, by virtue of its smooth ry material. Specimens were carefully aboral surface, seems more similar to the removed with forceps in order to avoid original Coeloplana than do any of the damage or collected while floating freely at species subsequently ascribed to this the water’s surface. Specimens were re- genus. laxed for 2 h in isotonic MgSO4 and VOLUME 127, NUMBER 2 425

National Museum of Natural History (NMNH), Smithsonian Institution. Systematics Order Platyctenida Bourne, 1900 Coeloplanidae Willey, 1896

Diagnosis.—Creeping or sessile cteno- phores, ctene rows absent in adults. , with tentilla, retractable into sheaths. Meridional canals branched and anastomosed. permanently evert- ed; present and aboral; body length ,6 cm. Coeloplana Kowalevsky, 1880

Diagnosis.—With the features of the family and often with a high host specific- ity. May or may not have an oral groove; can form temporary chimneys. Coeloplana waltoni, new species Figs. 2–6

Material examined.—Coeloplana walto- Fig. 1. Map of upper Florida Keys showing ni. 1960s: The following information is collection sites of Coeloplana waltoni. Triangles ¼ taken from F. M. Bayer’s unpublished 1960s, circles ¼ 2012–2013. Vallicula multiformis was manuscript. Several specimens living on collected at Crandon Park Marina, Key Biscayne the gorgonian Eunicea palmeri Bayer, 1961 (open rectangle). collected at Soldier Key, Biscayne Bay, Florida, 10 Oct 1960, by F. M. Bayer, fixed preserved in 70% and 95% ethanol or fixed in Bouin’s. One specimen prepared as a in 2% glutaraldehyde in 0.05 M sodium whole-mount on a slide. Twelve specimens, cacodylate buffered sea water. The alco- up to 4 mm in length, preserved in ethanol hol-preserved specimens would be suitable (USNM 52454). Several specimens living for molecular analysis. Site locations and on various gorgonians, including Eunicea additional information for recent collec- palmeri Bayer, 1961, Eunicea tourneforti tions of Coeloplana waltoni and Vallicula Milne Edwards & Haime, 1857, Plexaur- multiformis are contained in Figure 1 and ella dichotoma (Esper, 1791), Ragged Table 1. The sites of two collections of C. Keys, Biscayne Bay, Florida, Sep 1964, waltoni made near the pier at Dania Beach, collected by F. M. Bayer, J. J. McClelland, about 30 km north of Miami Beach, are and Samuel Stout; fixed in Bouin’s. Four not shown in Figure 1. prepared as a whole-mount on one slide; Abbreviations: UMML, University of one serially sectioned. Paratypes. Several Miami Marine Laboratory (now the Ma- specimens living on various gorgonians, rine Invertebrate Museum, Rosenstiel Ragged Keys, Biscayne Bay, Florida, Sep School of Marine and Atmospheric Sci- 1964, collected by John A. Jones and R. J. ence, University of Miami); USNM, Unit- Daly; preserved in formalin. Paratypes. ed States National Museum (now the Several specimens living on gorgonians, 2 RCEIG FTEBOOIA OIT FWASHINGTON OF SOCIETY BIOLOGICAL THE OF PROCEEDINGS 426 Table 1.—Benthic ctenophore collection information, Coeloplana waltoni and Vallicula multiformis Rankin, 1956, southeast Florida, 2012–2013. Numbers (n)of individuals denote those found on one to three, 10–20 cm-long branches of microhabitat. Values in brackets under Microhabitat denote wet settled volumes of algae sampled.

Date Species n Microhabitat Location Latitude, Longitude

18 Feb 2012 Coeloplana waltoni 30 Gorgonia ventalina Linnaeus, 1758 Broad Key 25820001.9600 N, 80812025.1700 W C. waltoni 35 Plexaura homomalla (Esper, 1792) C. waltoni 11 Pseudoplexaura porosa (Houttuyn, 1772) 11 May 2012 – 0 Pseudopterogorgia acerosa (Pallas, 1776) Emerald Reef 25839027.9500 N, 80807043.3400 W C. waltoni .48 Eunicea tourneforti Milne Edwards & Haime, 1857 –0Pseudopterogorgia acerosa –0Pseudopterogorgia hystrix Bayer, 1961 C. waltoni 1 Gorgonia ventalina 24 Oct 2012 Vallicula multiformis 3 Acanthophora spicifera þ Crandon Park Marina 25843032.0700 N, 80809019.0400 W clumps unidentified macroalgae [6 liters, 0.4 liters] 16 Nov 2012 – 0 Pterogorgia sp. Patch Reef 25843035.2800 N, 80805046.600 W –0Plexaurella nutans (Duchassaing & Michelotti, 1860) –0Eunicea laciniata Duchassaing & Michelotti, 1860 –0Eunicea calyculata (Ellis & Solander, 1786) –0Gorgonia ventalina 16 Nov 2012 – 0 Pterogorgia sp. Half Moon Wreck 25843038.6100 N, 80808004.0800 W C. waltoni 11 Eunicea laciniata 20 Feb 2013 V. multiformis 10 Filamentous , Enteromorpha sp. [6 liters] Crandon Park Marina 25843032.0700 N, 80809019.0400 W 27 Feb 2013 V. multiformis 2 Filamentous green algae, Enteromorpha sp. [2.5 liters] Crandon Park Marina 25843032.0700 N, 80809019.0400 W 20 Mar 2013 V. multiformis 1 Clumps of unidentified algae Crandon Park Marina 25843032.0700 N, 80809019.0400 W 12 Apr 2013 – 0 Pseudopterogorgia bipinnata (Verrill, 1864) Half Moon Wreck 25843038.6100 N, 80808004.0800 W –0Pseudopterogorgia acerosa –0Pseudoplexaura porosa –0Eunicea succinea (Pallas) forma plantaginea (Lamarck, 1815) 14 May 2013 C. waltoni 3 Plexaura flexuosa Lamouroux, 1821 Dania Beach 26803030.000 N, 80806031.500 W C. waltoni 12 Muriceopsis flavida (Lamarck, 1815) C. waltoni 3 Eunicea palmeri Bayer, 1961 22 Jul 2013 – 0 Plexaurella nutans Dania Beach 26803025.700 N, 80806032.400 W –0Gorgonia ventalina C. waltoni 3 Muricea muricata C. waltoni 4 Eunicea succinea C. waltoni 4 Eunicea calyculata C. waltoni 6 Muriceopsis flavida VOLUME 127, NUMBER 2 427

Fig. 2. Coeloplana waltoni. Live outline sketches. A, attached and resting; B–D, floating free; E–H, attached and creeping. Centrally located statocyst is visible in aboral views (A–C, E–G), the is visible in D, H. The long pinnate tentacles are visible in A–D, G, H. Drawings by F. M. Bayer.

Ragged Keys, Biscayne Bay, Florida, 22 1222177). Several specimens living as Oct 1964, collected by R. J. Daly and R. E. ectocommensals on Muriceopsis flavida, Gore; preserved in Zenker’s. 2–3 m depth off Dania Beach, 14 May 2012–2013: Re-examination of speci- 2013, collected by B. Coffman. Two mens collected at Soldier Key, Florida specimens, slides A and B, paratypes (USNM 52454). Several specimens living (USNM 1222178). Several specimens liv- as ectocommensals on Muriceopsis flavida ing as ectocommensals on Muriceopsis (Lamarck, 1815), 2–3 m depth off Dania flavida, 2–3 m depth off Dania Beach, Beach, 22 Jul 2013, collected by J. Levy Florida, 14 May 2013, collected by B. and B. Coffman. One specimen selected as Coffman. Three specimens selected as the holotype (USNM 1222176). Two paratypes (UMML 9.1). One specimen specimens selected as paratypes (USNM from Muriceopsis flavida, 2 m depth, off 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Coeloplana waltoni. Aboral views of live specimens with tentacles in various degrees of extension (A–F) and retraction (G–I), and illustrating the disposition of pigmentation. Smooth, non-papillate upper surface and mid-body thickening along tentacular axis are evident. Drawings by F. M. Bayer.

Dania Beach, Florida, 14 May 2013, sp. predominant, collected by J. Levy, 8 collected by B. Coffman; serial longitudi- specimens selected for study (UMML 9.3). nal sections, slides A4 and A5, paratype Site location and numbers of specimens (UMML 9.2). collected are noted in Figure 1 and Table Other material examined.—Vallicula 1. multiformis Rankin, 1956. 2012–2013: Description.—The newly designated ho- Crandon Park Marina, Biscayne Bay, lotype and paratype specimens collected in Florida, mixed macroalgal assemblages 2012–2013, both living and preserved (Acanthophora spicifera predominant), sus- individuals, were compared critically with pended from boat slips, 0.1–0.4 m depth, F. M. Bayer’s original description and collected by J. Levy and P. W. Glynn, 24 illustrations in his unpublished manuscript Oct 2012, one specimen selected for study of material examined in the 1960s with (UMML 9.4); 20 Feb 2013, Enteromorpha additional new information noted. VOLUME 127, NUMBER 2 429

Fig. 4. Coeloplana waltoni. Histological longitudinal section, 3 lm, stained with hematoxylin and eosin. Abbreviations: e, ; g.v.c., gastrovascular canals; o.g., oral groove; p., pharynx; s.t., ; t.f., tentacular filament; t.o., opening of tentacular sheath; t.s., tentacular sheath.

Animals of small size, at most about 5 floating free (Fig. 2B–D). When the animal mm in greatest length when fully extended is attached to a smooth surface and and creeping, usually 2–3 mm when resting, the outline is roughly oval, longer attached and passively resting (Fig. 2A), in the tentacular axis, but the shape is and only 1.5–2.0 mm when detached and extremely fluid, changing momentarily by

Fig. 5. Coeloplana waltoni. A–D, oral views of mouth and pharyngeal folds. Drawings by F. M. Bayer. 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Coeloplana waltoni. Aboral view of living free-floating specimen (~ 2 mm length). Collected off Dania Beach, 22 July 2013. local advances and retractions of the exterior at a slight notch in the margin margin. When creeping, the outline be- that can, however, be pulled back some comes quite elongated and flatworm-like distance from the edge of the animal (Figs. (Fig. 2E–H), and in this condition the 2–4). The tentacles are long – twice to three greatest length is diagonal to the tentacular times the diameter of the body – and axis, as the animals rarely advance with the serially pinnate; however, portions of the tentacular margin foremost; usually a distal tip are readily autotomized, which point to one side of the tentacular aperture together with their great motility, makes it leads. In this state, the width of even the almost impossible to obtain meaningful largest specimens may be as little as 0.5 measurements of their length from living mm. With the exception of the aboral material. The apical sense is well sense organ, which forms a small central developed and clearly visible with low protuberance, the aboral surface is entirely magnification as a pale central spot. Under smooth and completely lacking aboral higher power, the statolith can be clearly papillae or ‘‘dorsal tentacles.’’ The body viewed and appears to be in constant rapid is thickest in the central region over the vibration. The sense organ closes by stomach and pharynx, and along the contraction of the margin in an iris-like tentacular sheaths, so there is a slightly fashion; however, closure rarely occurs, raised elongated bulge situated in the and then only momentarily. Because of tentacular axis (Fig. 3A–I). The several irregular marginal lobulations or sheaths are pyriform and open to the foldings, the aperture is not circular. The VOLUME 127, NUMBER 2 431 functional mouth is roughly circular, and winter months, and in July, suggesting bounded by oral folds, and capable of round occurrence. More recently great distension. If the entire oral surface (2012–2013) gorgonian hosts included spe- of the animal is considered the everted cies of Eunicea, Gorgonia, Pseudoplexaura, outer pharynx, as in Vallicula (Rankin Pseudopterogorgia, Plexaura, and Muri- 1956:61), the mouth represents the opening ceopsis (Table 1). The highest numbers of of the inner pharynx. There seem to be C. waltoni,i.e.,.48 individuals, were distinct ‘‘oral grooves’’ along the tentacu- found on Eunicea tourneforti collected at lar axis (Fig. 4) as observed in Vallicula. Emerald Reef. Gorgonia ventalina and The wall of the inner pharynx, exposed Plexaura homomalla at Broad Key also when the mouth is open, is thrown into hosted numerous ctenophores, totaling 30 numerous irregular convolutions called and 35 individuals, respectively. Multiple pharyngeal folds (Figs. 4, 5A–D). The short branches (4–13 cm in length) of an canal system is much simpler than in individual of Plexaura flexuosa collected at Vallicula and other species of Coeloplana. Soldier Key on 10 October 1960 (USNM It is composed of simple spacious subdivi- 52454) still contain 12 individuals of C. sions of the meridional canals that radiate waltoni firmly attached to the coenen- outward from the stomach to the margin, chyme. Relatively large numbers also were reminiscent of the organization in the collected from Eunicea laciniata at the Half parasitic burrowing form of Lampea found Moon Wreck site (11 individuals) and in salps (¼ Gastrodes Korotneff, 1888, ¼ from Muriceopsis flavida off Dania Beach Gastra Stechow, 1921). The canals are in (18 individuals). When abundant on any such close proximity that there are appar- given host colony, the ctenophores became ently no connecting canals but only crowded and were often in contact. openings through the walls between adja- Etymology.—Named in honor of F. G. cent canals. In , the animals are Walton Smith, founder of the Rosenstiel translucent (not transparent). Some indi- School of Marine and Atmospheric Sci- viduals display a pale yellowish green tint ence, who first recognized the occurrence (Fig. 6), whereas others are yellowish of benthic ctenophores in American wa- brown and darker in the central area over ters. the stomach and along the tentacular Behavior.—The ctenophores creep slow- sheaths. There is a fine irregular reticula- ly on the surface of the gorgonian rind and tion of white on all individuals, and most are usually very close to the polyps. Often of them also have brown pigment granules, they are found encircling the calyces of the which in some are coalesced to form a polyps, even extending into the mouth. reticulation somewhat coarser than the The translucent body, with its flecks of white pattern. white and brown pigment, blends with the Habitat.—Coeloplana waltoni live as surface of the gorgonian, which is sprin- commensals on various species of shal- kled with white where spicules show low-water gorgonians. In the 1960s, they through the epidermis and also with were found on several species in the genera brownish or purplish pigment spots. The Eunicea, Plexaurella, and Muricea, and the ctenophores are extremely difficult to see incidence of infestation in some localities in situ and can be found only by close was almost 100%. In fact, on the several examination of the living gorgonian sub- occasions when collections were made, merged in seawater. Unlike specimens of every gorgonian examined was found to Coeloplana bocki, which Komai (1922) host these ctenophores. Specimens were removed from their alcyonacean hosts by found from September through May in the means of a pipette, individuals of Coelo- 1960s and more recently during the fall plana waltoni adhere tightly to the gorgo- 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2D). In this condition, the animals usually trail their tentacles, possibly as a flotation device and/or to catch in the branches of any gorgonian that might serve as a new host. If stimulated by moving water, the creeping animals also stream their tenta- cles in a food-catching attitude. They readily eat young nauplii of Artemia in the laboratory, so it seems likely that their natural food consists of minute zooplank- ters and the abundant that live as epizoites on the surface of the gorgoni- ans. Fig. 7. Vallicula multiformis Rankin. Resting on Comparisons.—It is unlikely that Coelo- unidentified host. Three clusters of papillae visible, plana waltoni could be confused with one around apical organ and one behind each Vallicula multiformis (Figs. 7, 8), which is tentacular aperture. Drawing by F. M. Bayer. the only other platyctene reported from the western Atlantic region. Its small size nian and can be dislodged only by (one-third to one-quarter that of Vallicu- carefully lifting them off with fine-pointed la), smooth aboral surface, simple, sac-like forceps. The oral surface often assumes a tentacle sheaths, simple canal system, and concave shape, which may aid in forming a translucent (rather than transparent) qual- suction cup-like seal. They are rather firm ity, render C. waltoni quite distinct. The and elastic and suffer no damage if habitat of C. waltoni, occurring on shallow handled gently. They produce quantities water gorgonians, is quite limited. Vallicu- of sticky that aids in clinging tightly la has never been found living upon to gorgonians living in swift currents. This gorgonians but occurs generally among mucus apparently is secreted by the highly algae, on the bryozoan Zoobotryon, and glandular that contains both on other fouling organisms, or under clear and granular cells, closely resembling stones (Marcus 1957). Vallicula has been the epidermis of Coeloplana mitsukurii and collected only in the boat harbor of the Lyrocteis imperatoris (Komai 1922:pl. 4, Crandon Beach Marina, where there is fig. 1; Komai 1942:fig. 4A). Once detached limited water motion, turbid conditions, from the host, the copious adhesive mucus and low light penetration. of C. waltoni makes the animals very Coeloplana sophiae Dawydoff, 1938 is difficult to dislodge from the forceps. the only other species in the genus known When unattached, they assume a ‘‘floating to associate with a gorgonian [Solenocau- phase’’ (Fig. 6; video clip MVI_0780) lon jedanensis in the South China Sea similar to that described in Vallicula by (Dawydoff 1938)]. Also, Coeloplana walto- Rankin (1956:58). Although it is presum- ni is markedly different from C. sophiae in ably the chief, if not the only, means of color and morphology. The latter is brick moving from one host to another, it is not red with white flecks, and possesses nu- known whether this ‘‘phase’’ is ever vol- merous distinctively arranged dorsal pa- untarily assumed by Coeloplana,asitisby pillae, and a unique median oral groove Vallicula.UnlikeVallicula, Coeloplana extending along the entire length of the does not fold itself lengthwise along the oral surface and ascending aborally to the tentacular axis but instead hollows the oral tentacular sheaths. surface and folds the margins inward on Relationships.—The characteristics of either side of the tentacular apertures (Fig. this new species seem clearly to support VOLUME 127, NUMBER 2 433

Fig. 8. Vallicula multiformis Rankin. Major papillae visible in lateral-aboral views (A, B), and mouth and pharyngeal folds in oral views (C–E). Drawings by F. M. Bayer. its placement in the original genus Coelo- pretations differ sufficiently from theirs to plana, rather than in Vallicula as conceived raise some doubt about the status of by Rankin (1956). The defining character- Vallicula. A form like the ‘‘floating phase’’ istics of the latter genus are its ‘‘anchor of Vallicula also occurs in Coeloplana shaped’’ tentacular sheaths, variable num- waltoni and may be widespread in Coelo- ber and position of aboral papillae, pres- plana. In spite of intensive ence of ‘‘food grooves’’ along the sampling around the Miami area where tentacular axis leading from the margin both genera are abundant, neither has to the pharyngeal opening, and occurrence been taken in the plankton. of a ‘‘floating phase’’ in which the animal In all of the material of Vallicula from folds itself lengthwise along the tentacular Florida, the aboral papillae seem to have a axis, turns up (aborally) the openings of rather consistent distribution, and regard- the tentacular sheaths to form short less of their retractility, do not change ‘‘chimney-like’’ structures. Also, Vallicula position. The major papillae commonly is able to drift in the plankton. Although occur in three groups of four: one group specimens collected in Florida appear to be around the apical organ, and one behind the same as those found in Jamaica by each tentacular opening (Fig. 7); some- Rankin (1956) and in Brazil by Marcus times the tentacular openings are flanked (1957), the present observations and inter- by two papillae instead of four (Fig. 8A, 434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

B), and additional laterally-placed papillae additional platyctenid-cnidarian associa- have also been observed (Fig. 8B). Smaller tions with the discovery of Coeloplana projections may appear along the gastro- anthostella and Coeloplana bocki inhabit- vascular canals, and these are so small that ing Dendronephthya spp. (alcyonaceans) in they can be fully retracted. Korea (Song & Hwang 2010, Song et al. The anchor-shaped tentacular sheaths, 2011) and Coeloplana sp. associated with suggested as characteristic of Vallicula the aboral surfaces of fungiid corals in (Figs. 7, 8A, C, E), probably are not Malaysia (Hoeksema et al. 2013). Three limited to V. multiformis. A similar con- species were listed as planktonic. Addi- figuration in Coeloplana bocki is suggested tional records are Coeloplana bannwarthi, in illustrations published by Komai inhabiting the echinoid Diadema (Eeck- (1922:pl. 2, fig. 1). Coeloplana scaberiae haut et al. 1997), and Coeloplana willeyi also has large tentacular sheaths with that adheres to the of the green alga varying cross-bar development, not so Caulerpa racemosa var. cylindracea (Cavas elaborately branching, but similar to Val- & Yurdakoc 2005). licula in some specimens (Matsumoto & Since the gorgonian colonies sampled Gowlett-Holmes 1996). When exposed, the typically ranged between 0.5 to 1.0 m in inner pharynx of Vallicula,asinCoelopla- height and possessed numerous branches na waltoni, consists of prominent folds and branchlets, producing a high surface (compare Figs. 4, 5, 8E). area, it is likely that single colonies could Coeloplana In view of the fact that host hundreds if not thousands of individ- metschnikowii Kowalevsky, 1880, the type ual Coeloplana waltoni. Because platyctene species of Coeloplana, has no dorsal ctenophores are capable of both sexual papillae, and that tentacular sheaths with and asexual reproduction (Hyman 1940), lateral branches occur in Coeloplana and Matsumoto & Gowlett-Holmes (1996) Vallicula, the various nominal species hypothesized that these dual reproductive should be reviewed and the validity of modes could explain their high population these characters reappraised. Pending such a review, generic alignment is not dis- numbers. The latter workers noted that a turbed, and the new species herein estab- platyctene species in Hawai’i, Vallicula multiformis, reaches population densities lished is assigned to Coeloplana. 2 Remarks.—Coeloplana waltoni repre- of over 3000 individuals m (C. P. Galt sents a new, minute species of platyctenid pers. comm.). Coeloplana scaberiae,a ctenophore that exhibits an ectosymbiotic species described by Matsumoto & Gowl- lifestyle. In Matsumoto’s (1999) summary ett-Holmes (1996) as inhabiting a brown of known Coeloplana host associations, alga in , attains densities of from five species were listed as being associated one to 50 individuals per alga. These with marine (mainly green, red, and workers found brooded embryos under brown algae), six species with the margins of 18 of 22 specimens and (echinoids and asteroids), and nine species observed asexual fragmentation. No sign epizooic on cnidarians. Coeloplana willeyi of sexual or asexual reproduction was Abbott, 1907 is found on a variety of observed in C. waltoni in the present study. plants and echinoderms. Of the cnidarian Some specimens collected off Dania hosts, seven species of Coeloplana were Beach, however, contained one to two or associated with alcyonaceans, and one three unidentified symbiotic flatworms, species each with a pennatulid and gorgo- possibly species of Dalytyphloplanida. nian cnidarian. At present, Coeloplana These moved freely within the pharynx sophiae is the only species associated with and stomach and are visible in video clip a gorgonian. It is now possible to add three MVI_0784. VOLUME 127, NUMBER 2 435

Acknowledgments de Chine Me´ridionale (Savangia atentaculata nov. gen. nov. spec.). Comptes Rendus Stephen Cairns is responsible for resur- Hebdomadaires des Se´ances de l’Acade´mie recting this unfinished study, and we thank des Sciences (Paris) 231(17):814–816. Eeckhaut, I., P. Flammang, C. Lo Bue, & M. him for the opportunity to describe this new Jangoux. 1997. Functional morphology of the species. Collections benefitted from the help tentacles and tentilla of Coeloplana bannworthi of Brian Coffman, Joshua Levy, Samuel A. (Ctenophora, Platyctenida), an ectosymbiont May, Ryan C. McMinds, and Joyce A. of Diadema setosum (Echinodermata, Echi- Yager. George I. Matsumoto, Claudia E. noida). Zoomorphology 117:165–174. Hoeksema, B. W., Z. Waheed, & A. Alamaru. 2013. Mills, and Nancy Voss helped with system- Out of sight: aggregations of epizoic comb atic questions. Various laboratory needs jellies underneath corals. Coral were provided by Patricia Blackwelder, Jay Reefs 32:1065. Fisch, and Chris Langdon. Herman Wirsh- Hyman, L. H. 1940. The . Part I. ing is thanked for his help in identifying Protozoa through Ctenophora. McGraw-Hill, New York, New York, 726 pp. octocoral species, and Julian P. S. Smith, Komai, T. 1922. Studies on two aberrant cteno- III for help in recognizing the suspected phores, Coeloplana and Gastrodes. Published dalytyphloplanid flatworms. This publica- by the author, Kyoto, Japan, 102 pp. tion benefitted substantially from the Komai, T. 1941. A new remarkable sessile cteno- phore. Proceedings of the Imperial Academy review process and editorial recommenda- Tokyo 17:216–220. tions offered by Claudia E. Mills, Stephen Komai, T. 1942. The structure and development of L. Gardiner, Rick Hochberg, and an the sessile ctenophore Lyrocteis imperatoris anonymous reviewer. Komai. Memoirs of the College of Science, Kyoto Imperial University, Ser. B 17(1):1–36. Korotneff, A. 1886. Ctenoplana kowalevskii. Zeits- Literature Cited chrift fu¨r Wissenschaftliche Zoologie 43:242– 251. Abbott, J. F. 1907. The morphology of Coeloplana. Korotneff, A. 1888. Cunoctantha und Gastrodes. Zoologische Jahrbu¨cher. Abteilung fu¨r Anat- Zeitschrift fu¨r Wissenschaftliche Zoologie omie und Ontogenie der Tiere 24:41–70. 47:650–657, pl. XL. Bayer, F. M. 1961. The shallow-water Octocorallia Kowalevsky, A. 1880. Coeloplana metschnikowii. of the West Indian region: a manual for Zoologischer Anzeiger 3:140. marine biologists. Martinus Nijhoff, The Marcus, E. d. B.-R. 1957. Vallicula multiformis Hague, The Netherlands, 373 pp. Rankin, 1956, from Brazil. Boletim do In- Bourne, G. C. 1900. Chapter VII, The Ctenophora. stituto Oceanogra´fico Sao Paulo 7(1–2):87–90. Pp. 1–25 in E. R. Lankester, ed., A treatise on Matsumoto, G. I. 1999. Coeloplana thomsoni sp. zoology. Part II, The Porifera and Coelenter- nov., a new benthic ctenophore (Ctenophora: ata. Adam & Charles Black, London, United Platyctenida: Coeloplanidae) from Western . Australia. Pp. 385–393 in D. I. Walker & F. Cavas, L., & K. Yurdakoc. 2005. An investigation E. Wells, eds., The seagrass flora and fauna of on the antioxidant status of the invasive alga Rottnest Island, : proceed- Caulerpa racemosa var. cylindracea (Sonder) ings of the Ninth International Marine Verlaque, Huisman, et Boudouresque (Cau- Biological Workshop, held at Rottnest Island, lerpales, ). Journal of Experimen- Western Australia, January 1996. Western tal Marine Biology and Ecology 325:189–200. Australian Museum, , Western Austral- Dawydoff, C. 1936. Les Ctenoplanidae des eaux de ia. l’Indochine franc¸aise, e´tude syste´matique. Matsumoto, G. I., & K. L. Gowlett-Holmes. 1996. Bulletin Biologique de la France et de la Coeloplana scaberiae sp. nov., a new benthic Belgique 70:456–486. ctenophore (Ctenophora: Platyctenida: Coe- Dawydoff, C. 1938. Deux Coeloplanides remarqu- loplanidae) from . Records of ables des eaux indochinoises. Comptes Re- the South Australian Museum 29:33–40. ndus Hebdomadaires des Se´ances de Mills, C. E. 1998–present. Ctenophora: list l’Acade´mie des Sciences (Paris) 206:1143– of all valid species names. Electronic internet 1145. document available at ,http://faculty. Dawydoff, C. 1950. La nouvelle forme de Cte´no- washington.edu/cemills/Ctenolist.html.;. phores planarise´s sessiles provenant de la Mer Published by the author, web page established 436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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