Abstract
In September 2003 specimens of the sympagic amphipod Gammarus wilkitzkii were sampled in drifting pack ice above 50 m water depth and in the pelagic/benthic environment beneath in the coastal waters off Svalbard (Norway). Abundance values ranged between 1.5 and 8 individuals m−2 for the pelagic/benthic and the sympagic environments, respectively, and showed corresponding sex ratios of 1:1.4, favoring males. In the ice a significantly higher number of juveniles prevailed. In contrast, ovigerous females were more abundant among pelagic/benthic living specimens. Approximately 25% of the individuals hosted ciliated epibionts of the genera Ephelota, Cryptacineta, Acineta, Podophrya (all suctoria), and Epistylis (peritrichia). Cryptacineta and Ephelota were the most abundant epibionts on this amphipod species. Female specimens of G. wilkitzkii showed the highest degree of infestation (>2,100 individual epibionts per amphipod specimen: indE/A) followed by juveniles (>1,200 indE/A) and males (>220 indE/A). Highest densities of epibionts were found on anterior body parts with the antennae bearing up to 130 individuals. This is the first sighting of epibionts on crustacea from the sympagic environment. We interpret them as biomarkers that indicate the existence of sympago–benthic coupling processes between the ice and the underlying waters and the seafloor. The population structure and the proportion of infested specimens are equal for the amphipods sampled from both the sympagic and the pelagic/benthic environment, indicating the existence of exchange (coupling) processes between the two habitats. Considering the annual ice cycle, local ice drift patterns, and the shallow water depth in the sampling area, we hypothesize that ice amphipods spend ice-free periods near the seafloor where they may serve as basibionts for protozoans and eventually re-colonize the ice with the onset of ice formation. Our observations strongly emphasize that shallow coastal areas serve as both retention (rather than sink) areas for ice fauna during ice-free periods, and as stepping-stones for re-colonizing the ice when the ice is formed in winter.
Similar content being viewed by others
References
Aarset AV (1991) The ecophysiology of under-ice fauna. Polar Res 10:309–324
Arndt CE (2002) Feeding ecology of the Arctic ice-amphipod Gammarus wilkitzkii—physiological, morphological and ecological studies. Ber Polar Meeresforsch 405:1–74
Arndt CE, Lønne OJ (2002) Transport of bioenergy by large scale Arctic ice drift. In: Squire V, Langhorne P (eds) Ice in the environment: Proc 16th IAHR Int Symp on Ice 2002:382–390
Arndt CE, Pavlova O (2005) Origin and fate of ice fauna in the Fram Strait–Svalbard area. Mar Ecol Prog Ser (in press)
Arntz WE, Brey T, Gallardo VA (1994) Antarctic zoobenthos. Ocean Mar Biol 32:241–304
Beuchel F, Lønne OJ (2002) Population dynamics of the sympagic amphipods Gammarus wilkitzkii and Apherusa glacialis in sea ice north of Svalbard. Polar Biol 25:241–250
Clamp JC (1993) A new species of Lagenophrys (Ciliophora: Peritrichia) symbiontic on marine amphipods. Trans Am Microsc Soc 112:62–68
Clarke A (1982) Temperature and embryonic development in polar marine invertebrates. Int J Invert Reprod 5:71–82
Cook JA, Chubb JC, Veltkamp CJ (1998) Epibionts of Asellus aquaticus (L.) (Crustacea, Isopoda): an SEM study. Freshw Biol 39:423–438
Cross WE (1982) Under-ice biota at the Pond Inlet ice edge and in adjacent fast ice areas during spring. Arctic 35:13–27
Curds CR (1985) A revision of the Suctoria (Ciliophora, Kinetofragminophora) 3. Tokophrya and its morphological relatives. Bull Brit Mus Nat Hist 49:167–193
Curds CR (1986) A revision of the Suctoria (Ciliophora, Kineotfragminophora) 4. Podophrya and its morphological relatives. Bull Brit Mus Nat Hist 50:59–91
Dunn AM, Dick JTA (1998) Parasitism and epibiosis in native and non-native gammarids in freshwater in Ireland. Ecography 21:593–598
Fenchel T (1965) On the ciliate fauna associated with the marine species of the amphipod genus Gammarus J. G. Fabricius. Ophelia 2:281–303
Fernandez-Galinao D (1976) Silver impregnation of ciliated protozoa: procedure yielding good results with the pyridinated silver carbonate method. Trans Am Microsc Soc 95:557–560
Fernandez-Leborans G (2000) A review of the species of protozoan epibionts on crustaceans III. Chonotrich ciliates. Crustaceana 74:581–607
Fernandez-Leborans G, Castro de Zaldumbide M (1986) The morphology of Anophrys arenicola sp. nov. (Ciliophora, Scuticociliatida). J Nat Hist 20:713–721
Fernandez-Leborans G, Tato-Porto ML (2000a) A review of the species of protozoan epibionts on crustaceans I. Peritrich ciliates. Crustaceana 73:643–683
Fernandez-Leborans G, Tato-Porto ML (2000b) A review of the species of protozoan epibionts on crustaceans II. Suctorian ciliates. Crustaceana 73:1205–1237
Friedrich C, Hendelberg J (2001) On the ecology of Acoala living in the Arctic sea ice. Belg J Zool 131:213–216
Gilbert JJ, Schröder T (2003) The ciliate epibiont Epistylis pygmaeum: selection for zooplankton hosts, reproduction and effect on two rotifers. Freshw Biol 48:878–893
Gulliksen B (1984) Under-ice fauna from Svalbard waters. Sarsia 69:17–23
Gulliksen B, Lønne OJ (1989) Distribution, abundance, and ecological importance of marine sympagic fauna in the Arctic. Rapp P-v Réun Cons Int Explor Mer 188:133–138
Haarpaintner J, Gascard JC, Haugan PM (2001) Ice production and brine formation in Storfjorden, Svalbard. J Geophys Res 106:14001–14103
Hausmann K, Hülsmann N, Radek R (2003) Protistology. Schweizerbartsche, Berlin
Hop H, Poltermann M, Lønne OJ, Falk-Petersen S, Korsnes R, Budgell WP (2000) Ice amphipod distribution relative to ice density and under-ice topography in the northern Barents Sea. Polar Biol 23:357–367
Lizotte MP (2003) The microbiology of sea ice. In: Thomas DN, Dieckmann GS (eds) Sea ice: an introduction to its physics, chemistry, biology and geology. Blackwell Science, Oxford, pp 184–210
Lønne OJ (1988) A diver-operated electric suction sampler for sympagic (= under-ice) invertebrates. Polar Res 6:135–136
Lønne OJ, Gulliksen B (1991) On the distribution of sympagic macro-fauna in the seasonally ice covered Barents Sea. Polar Biol 11:457–469
Lynn DH, Small EB (2000) Phylum Ciliophora. In: Lee JJ, Leedale GF, Bradbury P (eds) An illustrated guide to the protozoa. Allen Press, Lawrence, KS, pp 371–656
Matthes D, Guhl W, Haider G (1988) Suctoria und Urceolariidae. Protozoenfauna Band 7/1. Gustav Fischer, Stuttgart
Melnikov IA, Kulikov AS (1980) The cryopelagic fauna of the central Arctic Basin (in Russian). In: Vinogradov ME, Melnikov IA (eds) Biology of the central Arctic Basin. Nauka, Moscow, pp 97–111
Mohr JL, Tibbs J (1963) Ecology of ice substrates. Proceedings of the Arctic Basin Symposium, October 1962. Arctic Institute of North America, Calgary, pp 245–248
Morado JF, Small EB (1995) Ciliate parasites and related diseases of Crustacea: a review. Rev Fish Sci 3:275–354
Pöckl M (1992) Effects of temperature, age and body size on moulting and growth in the freshwater amphipods Gammarus fossarum and G. roeseli. Freshw Biol 27:211–225
Poltermann M (1997) Biologische und ökologische Untersuchungen zur kryopelagischen Amphipodenfauna des arktischen Meereises. Ber Polarforsch 225:1–170
Poltermann M, Hop H, Falk-Petersen S (2000) Life under Arctic sea ice—reproduction strategies of two sympagic (ice-associated) amphipod species, Gammarus wilkitzkii and Apherusa glacialis. Mar Biol 136:913–920
Steele DH, Steele VJ (1974) The biology of Gammarus (Crustacea, Amphipoda) in the northwestern Atlantic VIII. Geographic distribution of the northern species. Can J Zool 52:1115–1120
Swarczewsky B (1928) Zur Kenntnis der Baikalprotistenfauna. Die an Baikalgammariden lebenden Infusorien IV. Acinetidae. Arch Protistenk 63:25–37
Threlkeld ST, Chiavelli DA, Willey RL (1993) The organization of zooplankton epibiont communities. Trends Ecol Evol 8:317–321
Werner I, Auel H, Garrity C, Hagen W (1999) Pelagic occurrence of the sympagic amphipod Gammarus wilkitzkii in ice-free waters of the Greenland Sea—dead end or part of life-cycle? Polar Biol 22:56–60
Werner I, Auel H, Kiki R (2004) Occurrence of Anonyx sarsi (Amphipoda: Lysianassoidea) below Arctic pack ice: an example for cryo-benthic coupling? Polar Biol 27:474–481
Weslawski JM (1994) Gammarus (Crustacea, Amphipoda) from Svalbard and Franz Josef Land. Distribution and density. Sarsia 79:145–150
Acknowledgements
We would like to thank the crew on R.V. “Jan Mayen,” The University Centre on Svalbard, and Total E & P for logistical and financial support. We further acknowledge the helpful comments given by one anonymous reviewer and I. Werner.
Author information
Authors and Affiliations
Corresponding author
Additional information
Communicated by M. Kühl, Helsingor
Rights and permissions
About this article
Cite this article
Arndt, C.E., Fernandez-Leborans, G., Seuthe, L. et al. Ciliated epibionts on the Arctic sympagic amphipod Gammarus wilkitzkii as indicators for sympago–benthic coupling. Marine Biology 147, 643–652 (2005). https://doi.org/10.1007/s00227-005-1599-4
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00227-005-1599-4