BOOK 4: THE CTENOPHORES

Transcription

1 BOOK 4: THE CTENOPHORES

2 CSIRO Marine and Atmospheric Research 2014 This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be reproduced by any process or stored in any retrieval system or database without prior written permission from the copyright holder. The intent of this report is to be used as a field guide for the identification of midwater invertebrates of South-eastern Australia. It is envisioned that this report will lead to further editions as a published field guide; comments toward improving the presentation and usability are appreciated. This work was supported by CSIRO Wealth from Oceans Flagship. This work may be cited as: Gershwin, L., Lewis, M., Gowlett-Holmes, K., and Kloser, R The Ctenophores. In: Pelagic Invertebrates of South-Eastern Australia: A field reference guide. Version 1.1. CSIRO Marine and Atmospheric Research: Hobart. National Library of Australia Cataloguing-in-Publication entry Gershwin, Lisa-ann, author. The ctenophores / Lisa-ann Gershwin ; Mark Lewis ; Karen Gowlett-Holmes ; Rudy Kloser. ISBN: (paperback) ISBN: (ebook) Series: Pelagic invertebrates of south-eastern Australia. Field reference guide. Includes bibliographical references. Ctenophora--Australia, Southeastern--Identification. Marine invertebrates--australia, Southeastern--Identification. Lewis, Mark, author. Gowlett-Holmes, Karen L., author. Kloser, R. J., author v. 1.1 ~ April 2014 Page 1

3 Acknowledgments Figures were drawn by Ellen Lund Jensen / copyright CSIRO, unless otherwise noted. Trawl images were taken by Mark Lewis aboard the Southern Surveyor in trans-tasman research voyages and are the copyright of CSIRO Marine and Atmospheric Research. We sincerely thank the following people for use of their images: Lisa-ann Gershwin: cover, and pp. 5, 6, 13, 15. Underwater World: cover, p. 8. Martin George/Lisa-ann Gershwin: cover, p. 10. Karen Gowlett-Holmes: cover, pp. 11, 12. Denis Riek: cover, pp. 7, 9, 12, 14. David Paul / DPI Images: p. 5. South Australian Museum / Bob Hamilton-Bruce: p. 15. Cover montage Clockwise, from upper left: Pukia falcata (copyright L. Gershwin), Beroe forskalii (copyright D. Riek), Velamen parallelum (copyright K. Gowlett-Holmes), Ocyropsis vance (copyright K. Gowlett-Holmes), Bolinopsis ashleyi (copyright Underwater World), Leucothea filmersankeyi (copyright M. George/L. Gershwin). Ctenophore from the Tasman Sea, image taken by CSIRO remote optical system. v. 1.1 ~ April 2014 Page 2

4 GENERAL INFORMATION ABOUT THE CTENOPHORA Ctenophores, or Comb Jellies, are genuinely strange beasts: they have unusual habits, weird adaptations, and more often than not, unbelievably odd morphology. The phylum contains about 150 species. All are marine, most are bioluminescent, all are carnivorous, many are capable of blooming in huge numbers where they may become a problem for other species. But love them or hate them, ctenophores are pretty amazing, and often pretty. Long thought to be an evolutionary link between the Cnidaria (jellyfish, corals, and kin) and the worms, but recent DNA analysis has placed ctenophores as the sister group to the bilaterians (Podar et al., 2001), although this is still disputed (Philippe et al., 2009). Anatomy of a cydippid. Redrawn from Anatomy of a lobate. Redrawn from Anatomy of a beroid. Redrawn from Mayer ANATOMY All ctenophores have eight bands of cilia known as comb rows, some longer, some shorter... some lost in adults, others growing as the animal develops. These cilia rows are used in locomotion and often give the animal an iridescent appearance. Ctenophores cluster into three quite readily identifiable groups (orders): the cydippids (sea gooseberries), the lobates (the sea walnuts), and the beroids (jellies with jaws). Cydippids have a more or less spherical or oval body with two long tentacles with side branches called tentilla. Lobates are shaped like a blob with two large lobes at one end. The beroids are shaped like a pocket, open at one end. Like cnidarian jellyfish, the ctenophore body is gelatinous. But instead of having harpoon-like stinging cells as in the cnidarians, comb jellies have sticky cells called colloblasts which act more like a rope covered in honey. They are not toxic. Ctenophores also lack the polyp stage found in many cnidarians: fertilisation is external and the larvae develop directly into juveniles. Most ctenophores are hermaphroditic. v. 1.1 ~ April 2014 Page 3

5 ANNOTATED CLASSIFICATION PHYLUM CTENOPHORA CLASS TENTACULATA: With tentacles in at least some part of the life history Order Cydippida: Body spherical to globular, with two retractile posterior tentacles Euplokamis: Tentacle bulbs tadpole-shaped, along the sides of the stomach... 5 Pukia: Tentacle bulbs crescentic, wrapping around the end of the stomach... 6 Order Lobata: Body globular with two large anterior lobes Bolinopsis: The lobes comprise about half the body length... 8 Eurhamphaea: Two tentacles emit from corners at the aboral end... 9 Leucothea: Body covered in tactile gelatinous tubercles...10 Ocyropsis: The lobes comprise almost the entire body length...11 Unidentified black ctenophore.12 Order Cestida: Body extremely compressed and belt-shaped Cestum: canals parallel the stomach before departing; gonads continuous...13 Velamen: canals arise straight off the axial canal system; gonads discontinuous...13 CLASS NUDA (beroids): Lacking tentacles; the body is extremely compressed and pocket-shaped Beroe: Stomach occupying entire body; posterior end rounded or tapering to a point Neis: Stomach vase-shaped; posterior end drawn out into conspicuous lobe-like extensions...16 FURTHER INFORMATION The ctenophores of Australia were recently reviewed by Gershwin et al. (2010a), along with a dichotomous key to the species. Much of the information used herein was gleaned from the following sources: Gershwin, L., W. Zeidler, and P. J. F. Davie. 2010a. "Ctenophora of Australia." Memoirs of the Queensland Museum 54:1-45. Mayer, A. G Ctenophores of the Atlantic coast of North America. Vol. No Washington D.C.: Carnegie Inst. Washington Publ. Mills, C Ctenophores. Wrobel, D. Jellies Zone. v. 1.1 ~ April 2014 Page 4

6 CTENOPHORA: TENTACULATA: EUPLOKAMIDIDAE: The Sea Gooseberries Euplokamis evansae Gershwin, Zeidler, & Davie, 2010 Euplokamis sp. CAAB FIELD MORPHOLOGY: A small, cylindrical ctenophore with a conical forward end (mouth; right arrow) and two highly retractile tentacles coming out of sidepockets near the other end (left arrow). The species is characterised by its tadpoleshaped tentacle bulbs along the middle of the stomach. The side branches (tentilla) on the tentacles are held in tightly-coiled beehive-like bundles. Length to ~20mm. Euplokamis evansae Copyright David Paul / DPImages MAY BE CONFUSED WITH: The other Sea Gooseberry known from SE Australian waters is Pukia falcata, which is characterised by having crescent-shaped tentacle bulbs that curl around the base of the stomach (arrow, right), whereas in Euplokamis the bulbs are tadpole-shaped or elongate and alongside the stomach. It is also possible that the more or less spherical Sea Gooseberries might be confused with the spherical siphonophores, e.g., Sphaeronectes or Euplokamis sp. Copyright Lisa-ann Gershwin/CSIRO Amphicaryon, but the presence of either 8 external comb rows (ctenophores) or an internal yellowish stem (siphonophores) would immediately distinguish them. NOTES: Very little is known about the biology or ecology of Euplokamis evansae, as it was only discovered as new to science in It has only been reported twice originally from Tasmania and more recently from Port Phillip Bay but it is highly likely that the species is also found offshore and has simply escaped notice thus far. The Sea Gooseberries have the unfortunate feature of being born with their tentacles for food capture on the opposite end of the body from their mouth. They solve this problem with a spectacular somersaulting behaviour when they feed: when a copepod or other food particle is captured on a tentacle, the animal immediately shortens the tentacle and begins twirling towards it, eventually with enough spins the animal is able to draw the tentacle across the mouth and the lips grab the food particle. It s a lot of work for the animal, but quite entertaining to watch. v. 1.1 ~ April 2014 Page 5

7 CTENOPHORA: TENTACULATA: PUKIIDAE The Sea Gooseberries Pukia falcata Gershwin, Zeidler, & Davie, 2010 CAAB FIELD MORPHOLOGY: A small, more or less spherical to apple-shaped ctenophore with a protruding mouth on one end (upper arrow) and two highly retractile tentacles coming out of side-pockets near the other end. The species is characterised by its crescentic tentacle bulbs which curve around the bottom of the stomach (bottom arrow). The side branches (tentillae) on the tentacles are often held in tightly-coiled beehive-like bundles. Body length 15-20mm. Pukia falcata Copyright L. Gershwin MAY BE CONFUSED WITH: The preceding Sea Gooseberry, Euplokamis evansae, is superficially similar to Pukia, i.e., both are like small clear marbles with two tentacles. But they are easily distinguished from one another on the form of their tentacle bulbs: In Pukia, the bulbs are crescentic and curved around the bottom of the stomach, whereas in E. evansae the bulbs are tadpole-shaped running alongside the stomach. It is also possible that they might be confused with the spherical siphonophores, e.g., Sphaeronectes or Amphicaryon, but the presence of either 8 external comb rows (in ctenophores) or an internal yellowish stem (in siphonophores) would immediately distinguish the two groups. Pukia falcata Copyright L. Gershwin NOTES: Pukia is Australia s commonest ctenophore, being found most of the year in fairly large numbers, less abundant in the cooler months and cooler latitudes. Not much is known about its biology or ecology; its identity as a species was only just discovered in 2010! Like its other Sea Gooseberry cousins, Pukia must tumble and twirl in the water to draw food particles off its tentacles (see explanation in Euplokamis section). Pukia is a voracious predator on small crustaceans such as copepods it is thought that the jerking motion of the crustaceans when they are ensnared by the tentacles is what triggers the somersaulting action of the ctenophore. v. 1.1 ~ April 2014 Page 6

8 Juvenile Pukia falcata (images below). Note the coiled tentilla and the small, crescentic tentacle bulbs forming around the base of the stomodaeum (stomach). In the first photo (above), a polychaete larva can be seen inside the body of the animal. Images copyright Denis Riek. v. 1.1 ~ April 2014 Page 7

9 CTENOPHORA: LOBATA: BOLINOPSIDAE The Sea Walnuts Bolinopsis ashleyi Gershwin, Zeidler, & Davie, 2010 Bolinopsis spp. CAAB SS2011 TO2 OP39, MIDOC6 Net1 Bolinopsis ashleyi copyright Underwater World FIELD MORPHOLOGY: A blob about the size of an egg, maybe a bit larger, with about half the body length comprised of two massive lobes. Eight comb rows are conspicuous: two running the entire length of the body onto each lobe, and two on each short side of the body. Several different described and undescribed forms are found in SE Australian waters: Bolinopsis ashleyi has bright red pigmented racing stripes over the comb rows, and is found in southern Queensland and northern New South Wales A completely transparent and colourless form is common throughout the region A transparent form with fine magenta-pink internal canals is common in Great Australian Bight waters and likely extends both east and west MAY BE CONFUSED WITH: Bolinopsis species are easily confused with the highly invasive (and not yet reported in Australian waters) Mnemiopsis leidyi. In Bolinopsis the lobes start around the level of the mouth, whereas in Mnemiopsis the lobes start near the aboral end of the body. NOTES: In Spencer Gulf (South Australia) the occurrence of blooms of Bolinopsis appears to be on the increase; because of the close genetic relationship and similar trophic roles between Bolinopsis and the highly invasive Mnemiopsis, this situation should be monitored closely. Bolinopsis itself has been directly correlated with bloom problems in Japan. Bolinopsis is a voracious predator on a wide variety of zooplankton, from copepods to fish eggs and larvae, and crustacean and mollusc larvae. Few studies have been done on the fecundity and ecological parameters of Bolinopsis, but it is likely that they are similar to those of Mnemiopsis: the primary difference between the two genera appears to be an inch of lobe rather than a functional or ecological difference. v. 1.1 ~ April 2014 Page 8

10 CTENOPHORA: LOBATA: EURHAMPHAEIDAE The Filamented Lobate Eurhamphaea cruenta, new species CAAB Eurhamphaea cruenta, holotype in life, unpreservable Copyright Denis Riek FIELD MORPHOLOGY: The animal is broad in the tentacular axis, narrow in the stomodael axis. At the oral end it has two small-ish lobes and two short rabbit-earshaped auricles. At the aboral end are two solid filaments Eurhamphaea cruenta Copyright Denis Riek emitting from short triangular extensions of the body. The combs only extend as far forward as the base of the lobes, although the canals underlying them continue up into the lobes, then curl back around in a characteristic shape. The statocyst is very deeply embedded in the aboral end of the body, about 1/3 of the distance forward. The body and lobes are extremely transparent and colourless, except for a series of red pigment dots alternating with the ctene plates in each comb row. Total body length to about 80 mm, tentacles about 50mm. MAY BE CONFUSED WITH: Eurhamphaea is most similar to species of Bolinopsis, another type of lobate ctenophore, in having the lobes attached to the end of the body, small auricles, and a smooth body exterior. However, Eurhamphaea is more compressed than Bolinopsis, and it is the only ctenophore that has aboral filaments. NOTES: This form differs conspicuously from E. vexilligera, which has long triangular extensions of the body to the tentacles, whereas they are barely perceptible in E. cruenta. It has been found only once. It occurred in a thicket of other gelatinous zooplankton. The following observations were made in life: Apart from bobbing around at the surface it did very little. It didn t make any great moves of the lobes like an Ocyropsis would... It was able to right itself very quickly... the two "tentacles" seemed to act like a counterbalance or the tail on a kite. The tentacles were very reflective of the flash and if you look closely you can see that the white part is surrounded by a clearer layer, at least where they leave the body. (D. Riek, pers. comm.). v. 1.1 ~ April 2014 Page 9

11 CTENOPHORA: LOBATA: LEUCOTHEIDAE The Spiky Sea Walnuts Leucothea filmersankeyi Gershwin, Zeidler, & Davie, 2010 CAAB FIELD MORPHOLOGY: The animal is more or less eggshaped and covered in soft gelatinous bristles. Two massive lobes are about half the total length, connecting to the body at the forward end; the opposite end is where the comb rows meet and contains the statocyst (for balance). The colouration is completely transparent and colourless, but the gut may be faintly yellowish. Total length to about 13cm. Leucothea filmersankeyi Copyright Martin George / Lisa Gershwin MAY BE CONFUSED WITH: Leucothea is most similar to species of Bolinopsis, another type of lobate ctenophore, in having the lobes attached to the end of the body; Ocyropsis, another local lobate, has the lobes attached farther down on the body near the statocyst. However, this species is distinctive as no other local ctenophore has the gelatinous spikes. NOTES: The curious gelatinous spikes or bristles that cover the body have a sensory function that is not well understood. Experiments demonstrate that all spikes in a region will immediately point toward a stimulus; when not disturbed the spikes gently wave with no obvious pattern. Leucothea is incredibly fragile and extremely difficult to catch undamaged; only careful individual collection by dipping with a rigid container (e.g., aquarium or glass jar) will yield a whole specimen. The species is impossible to preserve, exploding at first contact with chemicals such as formaldehyde or alcohol. Leucothea is brilliantly bioluminescent, displaying flashes of blue light from the lobes and canals. Very little is known about the biology or ecology of Leucothea filmersankeyi; the species was only discovered in It has been found in abundance in Bass Strait waters off Tasmania, and is likely to be found elsewhere in southeastern Australia. v. 1.1 ~ April 2014 Page 10

12 CTENOPHORA: LOBATA: OCYROPSIDAE The Sea Clappers Ocyropsis vance Gershwin, Zeidler, & Davie, 2010 CAAB Ocyropsis vance, viewed from lateral (left) and aboral (right) Copyright Karen Gowlett-Holmes FIELD MORPHOLOGY: The body itself is quite short, with two massive wings or lobes that dominate the animal. The wings are almost the entire length of the animal, being attached to the body near its base, similar in appearance to a pair of praying hands. The stomach is a deeply-incised hourglassshaped structure easily visible through the body wall. MAY BE CONFUSED WITH: Other lobate ctenophores such as Bolinopsis, Leucothoe, and Mnemiopsis (the latter of which has not yet been reported in Australia), have a similar lobate form, but in those species the body is much larger and takes up a greater percent of the total length of the animal. NOTES: Most ctenophores do not pulsate or flap for locomotion, like the more familiar jellyfish do, but instead use their ciliated comb rows for tractor motion. When disturbed, however, Ocyropsis claps like a scallop with vigorous flapping motion of the two lobes as a means of escape. Very little is known about the biology or ecology Ocyropsis vance, as it was only discovered recently. It is one of the few lobates that can be preserved with formalin, whereas most others explode or disintegrate immediately upon contact with the chemical. This durable nature of the animal led it to being named after Cyclone Vance, the most powerful and longest lasting cyclone to hit Australia in recorded history. Like many other ctenophores, Ocyropsis is brilliantly bioluminescent. v. 1.1 ~ April 2014 Page 11

13 CTENOPHORA: LOBATA: Unidentified Unidentified Unidentified ctenophore from Tasman Sea, viewed from lateral (left) lateral-aboral (centre) and oral (right) Copyright CSIRO. Unidentified ctenophore from Tasman Sea, viewed from aboral (right and left) and aboral-lateral (centre) Copyright CSIRO. FIELD MORPHOLOGY: The body is blackish throughout, which is a highly unusual colour for ctenophores and other pelagic organisms. MAY BE CONFUSED WITH: The distinctive blackish colouration would no doubt make this species instantly recognisable. NOTES: The massive lobes suggest that this form may be related to Ocyropsis. Beyond that, nothing is known about its biology or ecology. It will quite likely be new to science when found and characterised. v. 1.1 ~ April 2014 Page 12

14 CTENOPHORA: CESTIDAE The Venus Girdles Velamen parallelum (Fol, 1869) Cestum veneris Lesueur, 1813 CAAB CAAB FIELD MORPHOLOGY: The diaphanous and fragile body is extremely compressed and drawn out into a beltshaped wing. The mouth is at the midpoint of the leading edge, while four long and four short comb rows are arranged along the posterior edge. Completely transparent, sometimes with yellowish or reddish pigment highlights along the margins. Velamen parallelum Copyright Karen Gowlett-Holmes Velamen: the canals arise straight off the axial canal system toward the lobes; the gonads appear discontinuous; length to 20cm. Cestum: the canals parallel the stomach for a short distance then curve sharply outward along the lobes; the gonads are in one long unbroken chain; length to 2m. MAY BE CONFUSED WITH: Unlikely to be confused with any other creature. Cestum veneris Copyright Denis Riek NOTES: These species explode on contact with preservatives, so they are unlikely to be found in preserved samples. However, they are relatively common in tropical and subtropical waters of the world, and may occasionally be found quite close inshore while snorkelling or scuba diving. Both species prey on small crustaceans such as copepods. They are occasionally infested with hyperiid amphipods. Cestids swim with the long oral edge leading, i.e., what appears to be sideways, with thin tentacles trailing over the body. The animal makes graceful sweeps back and forth through the water column, disturbing its prey with the turbulence of its first pass then capturing them with the tentacles on the next pass. When disturbed, the animal escapes by undulating vigorously like a snake in a direction perpendicular to normal movement. v. 1.1 ~ April 2014 Page 13

15 CTENOPHORA: NUDA The Swimming Mouths Beroe cucumis Fabricius, 1780 CAAB FIELD MORPHOLOGY: Body flimsy, gelatinous, highly compressed, sac-like, tapering aborally and squaredoff orally; the sides of the mouth are even with the widest part of the body. Total length to about 200mm, but more typically 60-80mm. The eight comb rows nearly reach the mouth. Colouration: overall with a pinkish, purplish, or even pale orange hue. MAY BE CONFUSED WITH: Beroe cucumis may be confused with B. gracilis or even with Neis cordigera. In B. cucumis, the mouth is straight or pursed relative to the body, and the overall colouration is faintly pinkish or purplish, whereas in B. gracilis the mouth is conspicuously flared relative to the body and the colouration is whitish with reddish granules sometimes present on the lips or comb rows; Neis, in comparison, is characterised by having wing-like extensions to the aboral end (away from the mouth), and typically bright red or yellowish streaks on the two narrower edges and the aboral lobes. Beroe cucumis -- SS2011_T02 OP4_MIDOC1 NOTES: Beroe is highly luminescent, flashing brilliant blue-green along the comb rows and sometimes throughout the canals of the body. Beroe is a voracious predator of other species of Beroe cucumis Copyright L. Gershwin ctenophores, and occasionally, of other gelatinous zooplankton. Almost immediately upon encountering a prey item, Beroe will just open its mouth and engulf the prey. For large or semi-rigid prey, the very floppy Beroe will take on the shape of the organism in the gut feeding experiments with gelatine cubes produced some rather strange looking cuboid Beroes! For organisms too large to fit into the gut, Beroe uses its teeth to bite off chunks. The teeth are modified cilia that line the lips. Because of the pocket-like shape of Beroe, it might seem that swimming forward may fill the body with water. However, Beroe has adhesive cells lining the mouth, which it uses to keep its mouth pursed shut during swimming. v. 1.1 ~ April 2014 Page 14

16 CTENOPHORA: NUDA The Swimming Mouths Beroe forskalii Milne-Edwards, 1841 CAAB Beroe forskalii (with an ingested Pukia) - Copyright Denis Riek Beroe forskalii SS2011_T02, OP25 MIDOC4 _Net2 FIELD MORPHOLOGY: Body flimsy, gelatinous, highly compressed, sac-like, with a constricted neck and flared mouth Beroe forskalii Copyright Chun, 1880 opening (arrow). The eight comb rows do not reach the mouth, running only from the statocyst (at the closed end of the body) to about 2/3 of the body length. Colouration: the lips and comb rows are typically magenta. Total length to 20cm. MAY BE CONFUSED WITH: Beroe forskalii may be confused with B. cucumis or even with Neis cordigera. In B. forskalii, the mouth is conspicuously flared relative to the body and the colouration is whitish with reddish or magenta granules sometimes present on the lips or comb rows, whereas in B. cucumis, the mouth is straight or pursed relative to the body, and the overall colouration is faintly pinkish, purplish, orangish, or yellowish; Neis, in comparison, is characterised by having wing-like extensions to the aboral end (away from the mouth), and typically bright red or yellowish streaks on the two narrower edges and the aboral lobes. NOTES: Like other species of Beroe, B. forskalii feeds by rapidly and aggressively engulfing its prey or biting it with modified cilia teeth that line the mouth. While most ctenophores are hermaphrodites, and most Beroe are both male and female simultaneously, a close relative of B. forskalii, namely B. ovata, can also self-fertilise; it seems likely that B. forskalii may also do this. This is the first record of this species in Australia. v. 1.1 ~ April 2014 Page 15

17 CTENOPHORA: NUDA The Swimming Mouths Neis cordigera Lesson, 1829 CAAB Neis cordigera, adult Copyright South Australian Museum/ photo by Bob Hamilton-Bruce Neis cordigera, juvenile Copyright L. Gershwin FIELD MORPHOLOGY: Body bag-like, soft gelatinous, flimsy; shaped like a match-box with a squaredoff mouth and two conspicuous lobes projecting aborally in adult specimens (lower arrow). The stomach is vase-shaped to funnel-shaped (upper arrow), flared at the mouth and tapering down narrowly to the statocyst with a bulge at the centre. The eight comb rows nearly reach the mouth. Colouration transparent to whitish, with crimson or yellowish pigmentation along the narrow sides extending down onto the lobes. Total length to about 15cm. MAY BE CONFUSED WITH: Similar morphologically to Beroe, but conspicuously differing in the posterior wings or lobes, i.e., in Beroe the posterior is U-shaped or V-shaped terminating at the statocyst, whereas in Neis the wings extend well beyond the statocyst, with the posterior being Mshaped. Furthermore, the stomach is vase-shaped and tapered in Neis, but in Beroe it occupies the whole body area. NOTES: Neis has not yet been reported from Tasmanian waters, but it has been sporadically found in the waters off Queensland, New South Wales, South Australia, and southern Western Australia, and therefore it seems inevitable that it will be found in Tasmanian and Bass Strait waters. It is unknown whether Neis is capable of emitting bioluminescence: under experimental manipulation the production of light could not be stimulated, however, members of the Beroida are typically highly luminescent so it is possible that more tests will be enlightening. v. 1.1 ~ April 2014 Page 16

18 GLOSSARY TO THE CTENOPHORES Aboral literally, away from the mouth ; the term is used for orientation. See also oral. Auricles in lobates, four slender straight or coiled appendages at the base of the lobes; the ciliated edges are thought to help in feeding and locomotion. Colloblasts microscopic non-venomous sticky-cells used in feeding, comparable with nematocysts (stinging cells) found in the phylum Cnidaria (jellyfish, corals, and sea anemones); they do not sting. Comb rows the eight rows of ciliary plates (or ctenes) that run longitudinally, giving the animal propulsion. Ctene rows see comb rows. Ctene the individual swimming plates that make up the ctene rows, each composed of many fused macrocilia. These structures are unique to the phylum Ctenophora ( comb bearer ). Infundibulum the broad canal running between the stomodaeum and the statocyst; also sometimes called the aboral canal. Meridional canals the circulatory canals running under the comb rows, connected to the stomodaeum by smaller perpendicular radial canals. Oral the mouth-end of the animal; see also aboral. Paragastric canals (pharyngeal canals) a pair of internal canals that run from the base of the stomodaeum up along its flattened sides toward the mouth. Polar plate the figure-8 shaped structure on the aboral end of the animal; believed to be sensory. Radial canals short canals that run laterally through the jelly, connecting the meridional canals (under the comb rows) to the base of the stomodaeum. Statocyst the balance organ embedded in the aboral end between the two lobes of the polar plate. Stomodaeum the gastric cavity, appearing as a flat canal between the mouth and the infundibulum. Sometimes also called the pharynx. Substomodaeal canals the four meridional canals (under the comb rows) that lie closest to the plane of the stomodaeum. Subtentacular canals the four meridional canals (under the comb rows) that lie adjacent to the tentacle sheath openings and to the plane running longitudinally through the tentacle bulbs. Tentacles in cydippids, two highly extensile filaments used in feeding; may be entirely retracted into the body through aborally-directed sheaths, or deployed externally. The tentilla are embedded with colloblasts. Tentacle bulbs the swollen bases of the tentacles; their size, shape, and position are taxonomically diagnostic. v. 1.1 ~ April 2014 Page 17

19 Tentacular sheaths two aborally-directed, funnel-shaped tubes through which the tentacles emerge when relaxed or deployed. Tentilla (tentillum, sing.) the feathery side branches on the tentacles of most tentaculates. v. 1.1 ~ April 2014 Page 18

20 CHECKLIST FOR THE CTENOPHORES Euplokamis Pukia Pukia (juvenile) Bolinopsis Eurhamphaea Leucothea Ocyropsis Unidentified Cestum/Velamen Beroe Beroe Neis Neis (juvenile) v. 1.1 ~ April 2014 Page 19

21 REFERENCES ON THE CTENOPHORES Arai, M. N Predation on pelagic coelenterates: a review. Journal of the Marine Biological Association of the United Kingdom 85(3): Baker, L. D. and M. R. Reeve Laboratory culture of the lobate ctenophore Mnemiopsis mccradyi with notes on feeding and fecundity. Marine Biology 26: Båmstedt, U Trophodynamics of Pleurobrachia pileus (Ctenophora, Cydippida) and ctenophore summer occurrence off the Norwegian north-west coast. Sarsia 83(2): Benham, W. B New Zealand ctenophores. Trans. N.Z. Inst. 39: , pl. 7. Bigelow, H. B Reports on the scientific results of the expedition to the eastern tropical Pacific... Albatross... XXVI. The ctenophores. Bulletin of the Museum of Comparative Zoology at Harvard College 54(12): Bishop, J. W Feeding rates of the ctenophore, Mnemiopsis leidyi. Chesapeake Science 8(4): Bishop, J. W A comparative study of feeding rates of tentaculate ctenophores. Ecology 49(5): Bourne, G The Ctenophora. Lankester's Treatise on Zoology. Part 2, Chapter 7: Burrell, V. G. and W. A. v. Engel A means of coping with Mnemiopsis leidyi in plankton samples. Chesapeake Science 11(2): Ceccaldi, H. J Observations biologiques de Cestus veneris. Tethys 4: Chun, C Die ctenophoren des Golfes von Neapel und der angrenzenden Meeres-abschnitte. Fauna and Flora des Golfes von Neapel 1: 313 pp., 16 plates. Chun, C Die Ctenophoren der Plankton-expedition. Kiel, Lipsius & Tischer. Coonfield, B. R. 1936b. Regeneration in Mnemiopsis leidyi, Agassiz. Biological Bulletin (Woods Hole) 71: Costello, J. H. and R. Coverdale Planktonic feeding and evolutionary significance of the lobate body plan within the Ctenophora. Biological Bulletin (Woods Hole) 195(2): Costello, J. H., R. Loftus and R. Waggett Influence of prey detection on capture success for the ctenophore Mnemiopsis leidyi feeding upon adult Acartia tonsa and Oithona colcarva copepods. Marine Ecology Progress Series 191: de Lima, M. C. G. and J. L. Valentin New records of Amphipoda Hyperiidea in associations with gelatinous zooplankton. Hydrobiologia(448): Deason, E. E. and T. J. Smayda Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relative to ctenophore-zoopankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA. Journal of Plankton Research 4(2). Easterly, C. O A study of the occurrence and manner of distribution of the Ctenophora of the San Diego region. University of California Publications in Zoology 13: Engell-Sørensen, K., P. Andersen and M. Holmstrup Preservation of the invasive ctenophore Mnemiopsis leidyi using acidic Lugol s solution. Journal of Plankton Research 31(8): Finenko, G. A., B. E. Anninsky, Z. A. Romanova, G. I. Abolmasova and A. E. Kideys Chemical composition, respiration and feeding rates of the new alien ctenophore, Beroe ovata, in the Black Sea. Hydrobiologia(451): Finenko, G. A., A. E. Kideys, B. E. Anninsky, T. A. Shiganova, A. Roohi, M. R. Tabari, H. Rostami and S. Bagheri Invasive ctenophore Mnemiopsis leidyi in the Caspian Sea: feeding, respiration, reproduction and predatory impact on the zooplankton community. Marine Ecology Progress Series 314: v. 1.1 ~ April 2014 Page 20

22 Finnerty, J. R., V. A. Master, S. Irvine, M. J. Kourakis, S. Warriner and M. Q. Martindale Homeobox genes in the Ctenophora: Identification of paired-type and Hox homologues in the atentaculate ctenophore, Beroe ovata. Molecular Marine Biology and Biotechnology 5(4): Franc, J. M The mesoglea of Ctenophores. Bulletin de la Societe Zoologique de France 113(4): Frank, K. T Ecological significance of the ctenophore Pleurobrachia pileus off southwestern Nova Scotia [Canada]. Canadian Journal of Fisheries and Aquatic Sciences 43(1): Gershwin, L., W. Zeidler and P. J. F. Davie Ctenophora of Australia. Memoirs of the Queensland Museum 54(3): Gomez-Aguirre, S [Notes on the biology and culture of Pleurobrachia Fleming 1822 (Ctenophora: Cydippida)]. Bol. Mus. Mar, Bogota 8: Govoni, J. J. and J. E. Olney Potential predation on fish eggs by the lobate ctenophore Mnemiopsis leidyi within and outside the Chesapeake Bay [USA] plume. Fishery Bulletin 89(2): Greene, C. H., M. R. Landry and B. C. Monger Foraging behavior and prey selection by the ambush entangling predator Pleurobrachia bachei. Ecology 67(6): Greve, W Cultivation experiments on North Sea ctenophores. Helgolander wiss. Meeresunters. 20: Greve, W., J. Stockner and J. Fulton Towards a theory of speciation in Beroe. Coelenterate Ecology and Behavior. G. Mackie. New York, Plenum Publishing Company: Gudger, E. W Some ctenophore fish-catchers. Scientific Monthly 57(1): Gyllenberg, G. and W. Greve Studies on oxygen uptake in ctenophores. Ann. Zool. Fenn 16: Haddock, S. H. D Comparative feeding behavior of planktonic ctenophores. Integrative and Comparative Biology 47(6): Haddock, S. H. D. and J. F. Case Not all ctenophores are bioluminescent: Pleurobrachia. Biological Bulletin (Woods Hole) 189(3): Hamer, H. H., A. M. Malzahn and M. Boersma The invasive ctenophore Mnemiopsis leidyi: a threat to fish recruitment in the North Sea? Journal of Plankton Research 33(1): Hamner, W. M., S. W. Strand, G. I. Matsumoto and P. P. Hamner Ethological observations on foraging behavior of the ctenophore Leucothea sp. in the open sea. Limnology and Oceanography 32(3): Harbison, G. R On the classification and evolution of the Ctenophora. The origins and relationships of lower invertebrates. S. C. Morris, J. D. George, R. Gibson and H. M. Platt. Oxford, Oxford University Press: Harbison, G. R Toward a study of the biogeography of pelagic ctenophores. UNESCO TECHNICAL PAPERS IN MARINE SCIENCE 49(1985 [1986]): Harbison, G. R Ctenophora. Introducción al Estudio del Zooplancton Marino. R. Gasca and E. Suárez. Chetumal, Mexico, ECOSUR: Harbison, G. R. and L. P. Madin Ctenophora. Taxonomy and classification of living organisms, vol. 1. S. P. Parker. New York, McGraw-Hill: , plates 68, 69. Harbison, G. R., L. P. Madin and N. R. Swanberg On the natural history and distribution of oceanic ctenophores. Deep-Sea Research 25(3): Harbison, G. R. and R. L. Miller Not all ctenophores are hermaphrodites. Studies on the systematics, distribution, sexuality and development of two species of Ocyropsis. Marine Biology 90: Hargitt, C. W Ctenophora, systematic synopsis, in: The medusae of the Woods Hole region. Bull. Bureau Fish. Washington xxiv: v. 1.1 ~ April 2014 Page 21

23 Hirota, J Laboratory culture and metabolism of the planktonic ctenophore, Pleurobrachia bachei A. Agassiz. Biological oceanography of the northern North Pacific Ocean. Tokyo, Idemitsu Shoten: Hirota, J Quantative natural history of Pleurobrachia bachei in La Jolla Bight. Fish. Bull. U.S. 72: Hoffmeyer, M. S Some feeding observations on Mnemiopsis mccradyi Mayer, 1900 (Ctenophora, Lobata). Iheringia Série Zoologia 70: Horridge, G. A Recent studies on the Ctenophora. Coelenterate Biology. Muscatine and Lenhoff: Johnsen, S Transparent animals. Scientific American 282(2): Kamshilov, M. M Feeding of ctenophore Beroe cucumis. Fabr. Dokl. (Proc.) Acad. Sci. U.S.S.R. (Sect. Biol. Sci.) 130(1/6): [Transl. from Dokl. Akad. Nauk SSSR 130(5): ]. Kasuya, T., T. Ishimaru and M. Murano Feeding characteristics of the lobate ctenophore Bolinopsis mikado Moser. Bulletin of Plankton Society of Japan 41(1): Kasuya, T., T. Ishimaru and M. Murano Metabolic characteristics of the lobate ctenophore Bolinopsis mikado (Moser). Plankton Biology and Ecology 47(2): Kasuya, T., T. Ishimaru and M. Murano In situ Measurement of the Lobate Ctenophore Bolinopsis mikado (Moser) Abundance by the Video Recording System. Journal of Oceanography in Taiwan Strait 57: Kideys, A. E., A. Roohi, S. Bagheri, G. Finenko and L. Kamburska Impacts of Invasive Ctenophores on the Fisheries of the Black Sea and Caspian Sea. Oceanography 18(2): Komai, T On ctenophores of the neighborhood of Misaki. Annotationes Zoologicae Japonenses 9: , 1 plate. Komai, T A note on the phylogeny of the Ctenophora. Lower Metazoa, Comparative Biology and Phylogeny. E. C. Dougherty and e. al., Univ. California Press: Kremer, P., M. R. Reeve and M. A. Syms The nutritional ecology of the ctenophore Bolinopsis vitrea: comparisons with Mnemiopsis mccradyi from the same region. Journal of Plankton Research 8(6): Larson, R. J In-Situ Feeding Rates of the Ctenophore Mnemiopsis mccradyi. Estuaries 10(2): Larson, R. J Feeding and functional morphology of the lobate ctenophore Mnemiopsis mccradyi. Estuarine Coastal and Shelf Science 27(5): Lee, G., S. Y. Park, J. Hwang, Y.-H. Lee, S. Y. Hwang and e. al Development of DNA chip for jellyfish verification from South Korea. BioChip Journal 5(4): Madin, L. P Feeding behavior of tentaculate predators: in situ observations and a conceptual model. Bulletin of Marine Science 43(3): Madin, L. P. and G. R. Harbison Bathocyroe fosteri gen. nov., sp. nov.: a mesopelagic ctenophore observed and collected from a submersible. Journal of the Marine Biological Association of the United Kingdom 58: Madin, L. P. and G. R. Harbison Thalassocalyce inconstans, new genus and species, an enigmatic ctenophore representing a new family and order. Bull. Mar. Sci. 28(4): Main, R. J Observations on the feeding mechanism of a ctenophore, Mnemiopsis leidyi. Biological Bulletin (Woods Hole) 55: Martindale, M. Q Larval reproduction in the ctenophore Mnemiopis mccradyi (order Lobata). Marine Biology (Berlin) 94(3): Martorelli, S. R Digenea parasites of jellyfish and ctenophores of the southern Atlantic. Hydrobiologia(451): Matsumoto, G. I A new species of lobate ctenophore, Leucothea pulchra sp. nov., from the California Bight. Journal of Plankton Research 10(2): v. 1.1 ~ April 2014 Page 22

24 Matsumoto, G. I Ctenophores of the Monterey Canyon (including a general key, photographs, and slides). Unpublished Text, Monterey Bay Aquarium Research Institute. Pacific Grove, CA: 34. Matsumoto, G. I Phylum Ctenophora (Orders Lobata, Cestida, Beroida, Cydippida, and Thalassocalycida): Functional morphology, locomotion, and natural history (morphology). Los Angeles, U.C.L.A.: 266. Matsumoto, G. I Functional morphology and locomotion of the arctic ctenophore Mertensia ovum (Fabricius) (Tentaculata: Cydippida). Sarsia 76(3): Matsumoto, G. I Swimming movements of ctenophores, and the mechanisms of propulsion by ctene rows. Hydrobiologia 216/217: Matsumoto, G. I. and W. M. Hamner Modes of Water Manipulation by the Lobate Ctenophore Leucothea sp. Marine Biology (Berlin) 97(4): Matsumoto, G. I. and G. R. Harbison In Situ Observations of Foraging, Feeding, and Escape Behavior in Three Orders of Oceanic Ctenophores: Lobata, Cestida, and Beroida. Marine Biology 117: Matsumoto, G. I. and B. H. Robison Kiyohimea usagi sp. nov., a new lobate ctenophore from the Monterey Submarine Canyon. Bulletin of Marine Science 5(1): Mayer, A. G Ctenophores of the Atlantic coast of North America. Washington D.C., Carnegie Inst. Washington Publ. McCrady, J On the development of two species of Ctenophora found in Charleston Harbor. Proceedings of the Elliott Society of Natural History of Charleston, South Carolina 1: , plate 14. Mianzan, H. W Ctenophora. South Atlantic Zooplankton. D. Boltovskoy. Buenos Aires, SPB Academic Publishing BV, Netherlands: Mianzan, H., M. Pajaro, G. A. Colombo and A. Madirolas Feeding on survival-food: Gelatinous plankton as a source of food for anchovies. Hydrobiologia(451): Mianzan, H. W., N. Mari, B. Prenski and F. Sanchez Fish predation on neritic ctenophores from the Argentine continental shelf: A neglected food resource? Fisheries Research (Amsterdam) 27(1-3): Mills, C. E Revised classification of the genus Euplokamis Chun, 1880 (Ctenophora: Cydippida: Euplokamidae n. fam.) with a description of the new species Euplokamis dunlapae. Canadian Journal of Zoology 65(11): Mills, C. E Medusae, siphonophores, and ctenophores as planktivorous predators in changing global ecosystems. ICES Journal of Marine Science 52(3-4): Mills, C. E Jellyfish blooms: Are populations increasing globally in response to changing ocean conditions? Hydrobiologia 451: Mills, C. E. and N. McLean Ectoparasitism by a dinoflagellate (Dinoflagellata: Oodinidae) on 5 ctenophores (Ctenophora) and a hydromedusa (Cnidaria). Diseases of Aquatic Organisms 10(3): Mills, C. E. and R. L. Miller Ingestion of a medusa (Aegina citrea) by the nematocyst-containing ctenophore Haeckelia rubra (formerly Euchlora rubra): phylogenetic implications. Marine Biology 78: Mortensen, T Ctenophora. Danish Ingolf-Expedition 5(Part 2): 1 95, plates 1 10, 1 chart. Mortensen, T Papers from Dr. Th. Mortensen's Pacific Expedition XXXIX. Two new ctenophores. Videnskabelige Meddelelser Dansk Naturhistorisk Forening 83: Mortensen, T Ctenophora. Report on the Scientific Results of the "Michael Sars" North Atlantic Deep Sea Expedition, (Part 2, No. 1): 1 12, 1 plate. Moser, F Die Ctenophoren der Siboga-Expedition. Leiden, E. J. Brill. Moser, F Ctenophores de la Baie d'amboine. Revue Suisse de Zoologie 16: v. 1.1 ~ April 2014 Page 23

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