FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: ©2009 Inter-Research Science Center. This manuscript is available at http://www.int-res.com/ and may be cited as: Jacoby, C. A., Youngbluth, M. J., Frost, J. R., Flood, P. R., Uiblein, F., Båmstedt, U., Pagès, F., & Shale, D. (2009). Vertical distribution, behavior, chemical composition and metabolism of Stauroteuthis syrtensis (Octopoda:Cirrata) in the Northwest Atlantic. Aquatic Biology, 5(1), 13-22. doi:10.3354/ab00117. _- - Vol. 5: 13-22, 2009 AQUATIC BIOLOGY Printed March 2009 Published online Pebrusrv s. 2009 doi: 10.3354/ab00117 Aquat BioI 1 Vertical distribution, behavior, chemical composition and metabolism of Stauroteuthis svrtensis (Octopoda: Cirrata) in the northwest Atlantic 3 Charles A. Jacoby':", Marsh J. Younqbluth", Jessica R. Prost':", Per R. Flood , 4 Franz Uiblein , Ulf Bamstedt", Francese Paqes'':", David Shale? 'University of Florida, Gainesville, Florida 32653, USA '!IIMhor Branch Oceanoqraphlc InsliLution at Florida Atlantic University, 5600 Highway 1 North, Fort Pierce, Florida 34946, USA "Bathybtoloqica A/S, Gerhard Grans vei 58, 5081 Bergen, Norway "Institute of Marine Research, Nordnesgaten 33, PB 1870 Nordnes, 5817 Bergen, Norway sUmeiI Marine Sciences Centre, Norrbyn, 910 20 Hornetors, Sweden "Institut de Ciencies del Mar (CSIC), Passeig Marilim de Ia Barceloneta 37-49,08003 Barcelona, Spain 714 Victoria Avenue, Swanage, Dorset BH19 IAN, UK "Present address: Institute for Hydrohiology and Fisheries Science, University of Hamburg, Olbersweg 24, 22767 Hamburg, Germany ABSTRACT: The cirrate octopod Stauroteuthis syrtensis is a mesopelagic species commonly collected in the North Atlantic. Individuals were observed at depths >600 m and typically within 100 m of the bot­ tom in three -900 m deep canyons indenting the southern edge of Georges Bank. When first sighted, most octopods were floating passively with their webbed arms gathered into a small ball. When dis­ turbed, they expanded their webs to form a 'balloon' shape, swam slowly by sculling their fins, pulsed their webs like medusae and, in some cases, streamlined their arms and webs and moved away smoothly by rapidly sculling their fins. The bodies of 9 octopods comprised 92 to 95 '}';, water, with tissue containing 9 to 22 'X, carbon (C) and 2 to 4 % nitrogen (N). These values were similar to those reported 1 for medusae and ctenophores. Oxygen (02) consumption rates of 4.6 to 25.8 umol O 2 rr C h- were within ranges reported for medusae, ctenophores, and deep-water cephalopods. The stomachs of S. syrtensis, dissected immediately after capture, contained only the calanoid copepod Calanus tinmarchi­ cus. Calculations indicated that S. syrtensis need 1.3 to 30.1 indo d- 1 of C. titunatcbicus to meet their ~lg measured metabolic demand. Excretion rates (0.3 to 12.4 NH4+ s' C h' and Cl.06 to 4.83 pg 1'04] ­ U C· II !) we'll; dt ledsL <111 order of muqnitudo lower th.in Idtes reported for other octopods or gelatinous zooplankters. O:N ratios (11 to 3(6) suggested that S. svrietisis catabolized lipids, which may be sup­ plied by C. Iinnuucbicus. Vertical distribution, relatively torpid behavior and low metabolic rates char­ actcrizcd S. svrtetisis as d benthopelaqic and relatively passive predator on copepods. KEY WORDS: Stauroieuthis svrtensis . Respiration . Excretion ' Chemical composition . Vertical distribution' Behavior --------------------- Resnlc or republication not permitted without written consent of the publisher ----"""-- -------- INTRODUCTION mens caught in bottom trawls and higher abundances reported at depths of 1500 to 2500 ill (e.g. Collins & Stauioteuthis syrtensis Verrill, 1879 is a deep-water Henriques 2000, Collins et aL 2001, 2002, Vecchione & octopod that occurs throughout the North Atlantic Galbraith 2001, Vecchione & Pohle 2(02). Specimens (Collins & Henriques 2000, Collins et ell. 2002, Collins captured at depths of 250 to 500 rn off Greenland and & Villanueva 20(6). Most specimens have been col­ Iceland provided evidence that the vertical distribution lected at depths of 500 to 4000 In, with many speci- of this species was influenced by water temperature as • Email: [email protected] © Inter-Research 2009' www.int-res.corn t Decedsf~d 14 Aquat BioI 5: 13- 22, 2009 Iceland provided evidence th at the vertical distribution of th is species was influenced by water temperature as much as, or more so th an, depth (Collins 2002) . Stsutote utbis syrtensis ha s been observed to display a variety of behaviors, including swimming by sculling its paired fins , swimmi ng via medusoid contractions of its web complex , inflating its web to varying degrees, and inve rting its arms and web to expose suckers that differ from those of most octopods in that they contain bioluminescent photophores and ar e not adhesive (Vecchione & Young 1997, Johnsen et al. 1999, Collins et al. 2002, Collins & Villanueva 2006) . Inflation and inv ersion of the web have been hypothesized to be defensive responses to disturbance or behaviors involved in capturing small cru stacean pr ey attracted by the biol uminescence emanating from the sucke rs (Vecchione & Young 1997, Johnsen et al. 1999, Collins et al. 2002). A se condary web that links the arms to the primary web in S. syrtensis and other Cirr ote uthidae facilitat es behaviors su ch as medusoid swimming, inflation and inv ersion. Sim ilar to other members of the suborder Cirra ta, Stauroteuthis syrtensis possesses a fragile, se mi-gela ti­ Fig . 1. Locat ion s of sampling for cirra te oc topods Stauro­ nous body (Collins & Henriques 2000 , Collins et al. teutbis syrten sis 2002), which suggests that th e met abolic rat es of S. syrtensis should resemble those of medusae and cteno­ in Oceanographer Canyon during September 2004 . At phores, rather than the higher respiratory demands of the sampling stations, dates, times, depths, tempera­ more mu scular cepha lopods . In addition, if S. syrten sis tures, sali nities, and conce ntrations of dissolved O2 am bushes pr ey, perhaps by using its bioluminescent wer e recorded with Seabird SBE 25 Sealoggers. suc ke rs as lur es, th en its O 2 consumption rates should Octopods and water samples without visible organ­ be relatively low and similar to other deep-water isms (i.e. controls) were collected in 6.5 I acrylic sam­ ce phalopods that have replaced visually guided pur­ plers that had been washed with a 1 mol l"! HCI solu­ suit of pr ey with an ambush strategy (Childress & tion prior to use (Tietze & Clark 1986) . An individual Mickel 1985, Seibel et al. 2000, Seibel & Carlini 2001). sampler contained either 1 individual of Stauroteuthis The present study provides the first measurements of syrtensis or a control water sample. 3 O 2 consumption and NH 4+ and P04 - excretion for Measu rements of respiration an d excretion. Upon Stauroteuthis syrtensis collected in Oceanographer return to the surface, samplers were transferred Ca nyon, a deep-water canyon that indents the south­ quickly from the submersible to a dark room main­ ern bo unda ry of Ge orges Bank. Furthermore, in situ tained at an air temperature of 4 to 5°C, which mim­ observa tions from cre wed subme rsibles documented icked in situ conditions. Within 28 h of capture, all the vertical distribution and behavior of these octopods octopods excep t the largest individual (wet weight, in this and 2 adj acent canyons, Hydrographer and WW = 158 g) were transferred gently into 0.5 to 2.2 I Lydonia Canyons. ac rylic chambers. Th es e smaller chambers, which acted as respirometers, were filled and sealed in a manner that avoided the formation or trapping of air MATERIALS AND METHODS bubb les . When additional water was needed to fill a sma lle r chamber, seawater collecte d in Niskin bottles Observation and collection. Stauroteuthis syrtensis from 800 to 900 m water depth was added. Th e largest were observed from 'J ohnso n-Sea-Link' submersibles, individual was too large for th e smaller chambers, so it which operat e to a depth of 914 m (Youngbluth 1984). was incubated in the original 6.5 I sampler, which Visual obse rva tions were made in Hydrographer, se rved as a respirometer. Oceanographer and Lyd on ia Canyo ns during June An experiment consisted of on e or more respirome­ 1995, September 2001 , June 2002 , September 2003, ters, each holding a single octopod, and a respirometer September 2004, and October 2005 (Fig . 1). Individuals that contained water without visible organisms, which for studies of respiration and excre tion wer e collected yielded measures of background changes in concen- Jacoby el al.: Distribution, behavior and metabolism of Steuroteuthis syrtensis 15 trations of dissolved 02, NH4 + and 1'04 :1- . Antibiotics to the nearest 0.1 g over a period of 2 to 4 d. Each dried were not added to any of the rcspirometors. During specimen was thoroughly homogenized, and aliquots experiments, all respiromctcrs were kept in darkness were sent to the Analytical Laboratory of the Marine at 4 to 5°C and ambient pressure (1 atm) for 9 to 22 h. Science Institute at the University of California, Santa (}, consumption WdS mou surod concurrently in con­ Barbara, where proportions of C, Hand N were deter­ trol and experimental respirometers with a noninva­ mined using a Perkin-Elmer Model 240 elemental ana­ sive, rnicro-optode system (PreSens. http://microoptode. lyzer and an acetanilide standard. com/html/stert.htrnl). O 2 concentrations (prnol O 2 1 J) were loqqcd at 1 s intervals. In some experiments, re­ I (II dlllJl S \V( 'II' cond uctocl tor :30 min periods, separated RESULTS by :30 to 60 min intervals, so that O 2 consumption could be trucked in multiple respirometers using a single Vertical distribution micro-optode system.