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FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: © 1989 Marine Biological Association of the United Kingdom. This manuscript is an author version with the final publication available and may be cited as: Larson, R. J., Mills, C. E., & Harbison, G. R. (1989). In situ foraging and feeding behavior of narcomedusae (Cnidaria: Hydrozoa). Journal of the Marine Biological Association of the United Kingdom 69(4), 785-794. doi: 10.1017/S002531540003215X {yjO J. mar. bioI. Ass. U.K. (1989), 69, 785-794 785 Printed in Great Britain IN SITU FORAGING AND FEEDING BEHAVIOUR OF NARCOMEDUSAE (CNIDARIA: HYDROZOA) RONALD J. LARSON*, CLAUDIA E. MILLSt AND G. RICHARD HARBISON* 'Harbor Branch Oceanographic Institution, 5600 Old Dixie Highway, Fort Pierce, Florida 34946 USA tFriday Harbor Laboratories, University of Washington, 620 University Road, Friday Harbor, Washington 98250, USA (Figure 1) Narcomedusae are a small and mostly oceanic group of hydromedusae whose tentacle morphology and comportment sets them off behaviourally and perhaps ecologically from most other medusae. Their tentacles are relatively few in number (2-40), stiff, and non contractile, with points of insertion located well above the bell margin. Eleven species representing eight narcomedusan genera (Aegina, Aeginura,an undescribed aeginid, Cunina, Pegantha, Solmaris, Solmissus, and Solmundello) were observed and collected in situ in the NW Atlantic, Arctic and Antarctic, using scuba and manned submersibles. In life, the tentacles of narcomedusae are nearly always held upwards over the bell or projected laterally. The major prey were other gelatinous zooplankton, especially salps and doliolids. In the laboratory, these relatively large prey were caught on the tentacles which bend inward and coil at the tips to bring food to the mouth. By extending the tentacles perpendicular to the swimming path, these medusae achieve a relatively large encounter area, thus increasing the probability of contact with prey, for the amount of protein invested in tentacles. INTRODUCTION Narcomedusae are common oceanic hydromedusae that have been little studied because of their inaccessibility. Although there are only about 50 species, some of these are widely distributed (e.g. Aegina citrea Eschscholtz, Aeginura grimaldii Maas, Pegantha clara R.P. Bigelow, Cunina octonaria McCrady, Solmtssus incisa (Fewkes), Solmissus marshalli Agassiz & Mayer and Solmundeila biientaculaia (Quoy & Gaimard) (Kramp, 1961). For example,S. biteniaculata occurs in the Antarctic, tropics and boreal regions of the world oceans, and at shallow and great depths. Although most species are found above 1000 m, some species (e.g. Solmissus spp.) occur at greater depths [Hartman & Emery, 1956; Peres, 1958, 1959 (as 501maris)J. Narcomedusae are holoplanktonic medusae. In some species (e.g. Cunina peregrina, Pegantha triloba) the larvae are known to parasitize other medusae (Bouillon, 1987), whereas other species develop non-parasitically in the plankton. The morphology and histology of narcomedusae have been described by Vanhoffen (1908), Bigelow (1909), Russell (953), and others. They typically have a fragile lens- or dome-shaped umbrella a 786 RJ. LARSON, C.E. MILLSAND G.R. HARBISON 1-10em in diameter that is generally transparent, but may be pigmented in mesopelagic species (e.g. Aeginuragrimaldii). Narcomedusae have from 2 to 40 solid tentacles project ing from the bell margin. The non-extensile tentacles are equally spaced around the bell and may be equal in length to the bell radius or up to about twice as long as the bell diameter. Grooves extending downward from the tentacles allow each tentacle to bend inward across the lower portion of the bell, bringing attached prey into contact with the circular mouth. The stomach is flat and broad, covering most of the subumbrella surface and can accommodate large prey. The cnidome of narcomedusae has been described for only a few species. Where known, it consists exclusively of apotrichous isorhizas, which are sometimes most abundant on the upper (abaxial) side of the tentacles and can have an extremely long (9 mm) thread (Mackie & Mackie, 1963; Mills & Miller, 1984; Mills & Goy, 1988; Purcell & Mills, 1988). Because most of these medusae are oceanic, fragile, and transparent, they have rarely been observed alive in an undamaged state. Bigelow (909) noted that freshly netted specimens had tentacles that extended stiffly outwards. He also remarked that some species were powerful and active swimmers, while others were inactive owing to reduced swimming muscles. Gladfelter (973), who studied the locomotion of Solmissus marshalli, noted that the bell was divided into two zones, a rigid central disc and a flexible lateral wall. Upon contraction of the circular swimming muscle, the lateral wall bends inward and upward, providing propulsive thrust. Madin (988) described ten tacle postures of three narcomedusan genera observed by scuba diving or from sub mersibles. Mills & Goy (988) described very active swimming and extensive vertical migration of Solmissus albescens (Cegenbaur), as seen from a submersible. Narcomedusae can be abundant at mesopelagic depths. In fact, they were among the first mid water animals to be photographed in situ. Emery (952) and Hartman & Emery (956) photographed many Solmissus incisa off California using a remote 'benthograph' camera. One of their photographs shows five specimens swimming in various direc tions with the tentacles arched outwards (Hartman & Emery, 1956, plate 1, incorrectly identified as a trachyline medusa). Other photographs of narcomedusae have been taken by manned submersibles. Bernard (958), Peres 0958, 1959), Tregouboff (1959), Houot (960), and Laban et al. (963) observed and photographed several narcomedu san species in the Mediterranean and off Japan, as deep as 2000 rn, from the French bathyscaphe FNRS III. Most of their observations concerned vertical distributions but they did note that the tentacles of Solmissus albescens and Aegina citrea (incorrectly identified as Aeginopsis Iaureniii Brandy) are usually held stiffly outward or upward from the bell. Benovic (1973) and Mills & Goy (1988) have described the 300-600 m (50 144 m h') diel vertical migration of S. albescens in the southern Adriatic and Ligurian Sea in the Mediterranean. Mackie & Mackie (1963), at Friday Harbor, Washington, observed Solmissus marshalli, a weak swimmer that holds its tentacles either up over the bell or laterally from the sides. One specimen that was maintained in an aquarium, ingested a hydromedusa (Euphysa sp.). Mills observed a S. marshalli in situ at the same location feeding on Euphysa sp. FEEDING AND FORAGING IN NARCOMEDUSAE 787 Mills & Miller (1984) have described predation on Aegina citrea by the ctenophore Haeckelia rubra (Kolliker), In the same paper they report feeding by the narcomedusa Cunina sp. on H. rubra in the laboratory. Mills & Goy (988) reported in situ observations from the submersible Cyana of several Solmissus albescens in the Ligurian Sea with the pteropod Cavolinia sp. in their guts, and of one S. albescens capturing, but subsequently dropping, the ctenophore Bathocyroe sp. In this paper, we describe swimming and tentacle posture, feeding, and diet for representatives of eight genera of narcomedusae that have been observed, photo graphed, and collected using scuba and manned submersibles in the western North Atlantic Ocean and elsewhere. .. METHODS Some of the medusae were observed in situ and were collected with the aid of scuba equipment, using 'blue water' diving techniques (Madin et a1., 1986) during a series of scuba transects made on a trans-Atlantic cruise of the RV 'Alantis' I [cruise no. 101, June-July 1978: see Bidigare & Biggs (980) for cruise track]. Other medusae, were observed in the Antarctic (McMurdo Sound) in December 1987 and in the Arctic (between Iceland and Spitsbergen) during August 1988. For observations of feeding behaviour, medusae were collected individually in jars and were maintained in ship board aquaria at near sea-surface temperatures (see Table 1). Mesopelagic narcomedusae were observed, photographed, video recorded, and col lected using the manned submersibles 'Johnson-Sea-Link' I and II [equipment fully described by Youngbluth (1984)] during cruises off the Bahamas in October 1984 and October 1988, off New England in September 1986 and August 1987, and off the Dry Tortugas in August September 1987. RESULTS In situ observations Representatives of eight narcomedusan genera (Aegina, Aeginura, an undescribed aeginid, Cunina, Pegantha, Solmaris, Solmissus, and Solmundella) were observed in situ and collected (Table 1). Narcomedusae in the sea were oriented in apparently random directions, but were usually swimming. Bell pulsation rates for epipelagic medusae were generally an inverse function of size, ranging from 0·08-0·17 Hz (pulsations per second) in relatively large (>30 mm bell diameter) sluggish species (Cunina duplicata and Cunina proboscidea), to 0·75-1·67 Hz in small medusae 00-20 mm bell diameter) (Aegina citrea and Solmaris corona). However, the small· epipelagic species, Cunina globosa, was relatively inactive, with pulsation rates of about 0·33-0·50 Hz. Pulsation rates of mesopelagic species ranged from 0·31-0·79 Hz for Solmissus spp. (bell diameter 30-60 mm) to 0·50-1·00 Hz for Aeginuragrimaldii (bell diameter 20-30mm) to 1·25 Hz for an undescribed aeginid (bell diameter