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Home , Pyrosoma atlanticum, Pyrosome, Thaliacea

The Scientific Naturalist

Ecology, 0(0), 2018, pp. 1–3 © 2018 by the Ecological Society of America appearance and persistence over multiple seasons as well as potential ecosystem impacts. Pelagic , which include , dolioloids, and , are Urochordates that spend their whole life cycle in the and feed using fine meshes. Range expansion of tropical Pyrosomes are colonies of zooids that are connected in a pyrosomes in the northeast chitinous tunic and resemble colonial benthic ascidians (Class ). Genetically identical blastozooids are Pacific Ocean added to the colony via asexual budding. colonies can reach lengths of several meters, with pyrosomes in the northeastern Pacific reaching up to 80 cm in length (Bro- I have just watched the moon set in all her glory, and deur et al. 2018). Ciliary beating within the zooids achieves looked at those lesser moons, the beautiful Pyrosoma, both suspension feeding and locomotion (Alldredge and shining like white-hot cylinders in the water. Madin 1982). Each zooid contains luminous organs that —T. H. Huxley, 1849, Diary of the Voyage of H.M.S. may be used to communicate with zooids further away Rattlesnake within the colony in response to mechanical or light stimuli (Bowlby et al. 1990). Pyrosomes are colonial pelagic tunicates that have fasci- Pyrosomes remain one of the least-studied planktonic nated marine biologists for over a century. Their name grazers, in spite of their widespread distribution and ecologi- comes from the “fiery” that luminous cal significance. Like other pelagic tunicates, pyrosomes are organs produce at night time. Blooms of pyrosomes, identi- known to form high density blooms reaching tens of individ- fied as (Peron, 1804), have recently uals per cubic meter, with swarms of P. atlanticum removing appeared in the North Pacific Ocean (Fig. 1), prompting >50% of phytoplankton standing stock in the 0–10 m layer questions about environmental factors that triggered their (Drits et al. 1992). Most , including P. atlanticum,

a b

c

d

FIG. 1. Pyrosome bloom in the northeast Pacific Ocean. (a) Large densities of Pyrosoma atlanticum observed at ~40 m depth during a February 2018 cruise off of Newport, Oregon, USA (also see Video S1), (b) pyrosome colonies from a net tow on the same cruise (photo by M. Farley), (c) pyrosomes washed up on beach in Newport, November 2017 (photo by A. Rainey) and, (d) dead pyrosomes on the sea floor (~50 m) at Cape Perpetua reef, Oregon, May 2017 (photo by S. Marion).

1 2 THE SCIENTIFIC NATURALIST Ecology, Vol. xx, No. xx have been considered tropical to subtropical in their distri- (Graham et al. 2001). Like other pelagic tunicates, pyro- bution (Van Soest 1981) with blooms previously reported in somes are filter feeders that use cilia to pump water into the southeast Atlantic (Drits et al. 1992) and the northwest their mucous filters to consume planktonic microorganisms Mediterranean (Andersen and Sardou 1994). (Mayzaud et al. 2007). P. atlanticum have been recorded to Pyrosomes are relatively common off of the California have some of the highest clearance rates of any pelagic gra- coast south of Cape Mendocino; for example, in a time ser- zer, with up to 35 L/h per colony (Perissinotto et al. 2007). ies of planktonic abundance, P. atlanticum was reported Their high filtration rates allow them to feed efficiently on about half the time during annual sampling off the coast of small planktonic microorganisms (Lavaniegos and Ohman southern California from 1951 to 2002 (Lavaniegos and 2003) down to the submicron scale (Sutherland et al. 2010). Ohman 2003), within its known latitudinal range. In 2014, These high filtration rates coupled with rapid reproduction scientists, fishermen, and beachgoers first started reporting and growth capabilities enable pelagic tunicates to be highly the appearance of P. atlanticum in coastal waters near responsive to environmental fluctuations (Alldredge and northern California, Oregon, and Washington north of Madin 1982). During blooms, they may have significant their previously reported range. The pyrosomes appeared impacts on food web dynamics through grazing and fecal again in 2015 and 2016. By the summer of 2017, they pellet production (Drits et al. 1992). P. atlanticum can appeared in unprecedented numbers along the entire west undertake extensive diel vertical migrations, migrating up to coast, reaching the Western Gulf of Alaska, but showed depths of 700 m (Angel 1989), potentially accelerating verti- some of their highest abundances off Oregon (Brodeur cal flux to the benthos. et al. 2018) clogging nets and disrupting marine activities The impacts of a bloom of this density and extent on eco- such as commercial and sport fisheries (Kaety Jacobson, logical interactions are unknown. In spite of their widespread personal communication). During two research cruises off distributions (Van Soest 1981), there are very few studies on the Pacific Northwest coast, USA in the summer of 2017, the dietary impact of pyrosomes. Analysis of pyrosome fecal colonies were 4–26 cm long and occurred in densities up to pellets suggested they consume small phytoplankton (3– 3 colonies/m3. Vertical video camera profiles and corre- 5 lm), coccolithophores, centric diatoms, and silicoflagellates sponding environmental data (temperature, salinity, and (Drits et al. 1992). However, based on their measured mesh fluorescence) indicated that layers of pyrosomes were dis- opening dimensions (0.6 lm; Bone et al. 2000) and the data tributed at ~60 m depth, at the base of the surface mixed supporting that other pelagic tunicates with larger meshes layer. These observations were made during ecosystem sur- consume submicron particles (Sutherland et al. 2010), it is vey cruises in May 2017 from Bodega Bay, California likely that they consume organisms in the nano- and (38° N) to Cape Meares, Oregon (45.5° N), and August picoplankton size range with relatively high efficiencies. This 2017 from Newport, Oregon, to the north end of Vancouver may allow them to persist during relatively oligotrophic con- Island, British Columbia, Canada (44° N–49° N). ditions that favor smaller cells as evidenced by their presence The appearance of such high densities of tropical pyro- during the “warm blob” event and also during winter off the somes in the temperate northeast Pacific presents interesting Oregon coast. Both of these phenomena represent periods of research questions about the physical oceanographic fea- increased stratification and reduced upwelling favoring smal- tures that led to their northerly expansion, and the environ- ler phytoplankton and bacterioplankton. It is presently mental drivers that have allowed their populations to persist unknown how these recent pyrosome blooms may interact for multiple years. Beginning in 2014, an unusually warm with or affect the dynamics of nutritionally valuable, lipid- and stable water mass termed the “warm blob” formed in rich plankton in the California Current. the North Pacific and lasted several years, and then was Once present in the ecosystem, pyrosomes may be pre- replaced by the northward progression of a strong El Nino~ dated upon or eventually sink to the bottom, serving as a in 2016 (Di Lorenzo and Mantua 2016), which may have benthic food source. A number of organisms, including sea facilitated the survival of this tropical species well north of turtles and sea birds, have been observed feeding on their normal range. Furthermore, onshore flow may have pyrosomes and other pelagic tunicates (e.g., Harbison 1998, pushed them closer to the shore than normal, leading to Perissinoto et al. 2007) and they can comprise a primary numerous reports of beached pyrosomes from 2014 to 2018 prey source for a number of fish species (Harbison 1998). (Fig. 1c). During a February 2018 cruise, we again observed However, reports are mostly descriptive owing to the chal- pyrosomes in large numbers during day and night at three lenges of conducting quantitative feeding studies with such stations (10, 25, and 45 km off shore) along the Newport, rapidly digested organisms. The sinking of dead and dying Oregon Hydrographic line (44.40° N), where temperatures pyrosomes occurred on a large scale during summer 2017 ranged from 10° to 10.8°C (Fig. 1a,b; Video S1). Similar to and were captured in benthic trawls and images of the sea the summer cruise observations, video footage showed that floor (Fig. 1d). Observations in the Gulf of Mexico and off pyrosomes were absent in surface waters but aggregated in a the coast of British Columbia have shown invertebrates, layer near the base of the surface mixed layer at ~40 m including anemones, sea urchins, and crabs directly consum- depth. ing pyrosomes (Archer et al. 2018). Until recently, jelly falls More generally, physical environmental parameters have been mostly overlooked as a source of carbon deposi- including temperature, light, salinity, dissolved oxygen tion to the sea floor (Lebrato et al. 2012). (DO), and currents have significant impacts on the biology The appearance of pyrosomes in temperate and subpolar and behavior of gelatinous aggregations latitudes challenges assumptions about their temperature Xxxxx 2018 THE SCIENTIFIC NATURALIST 3 tolerance. Moreover, their appearance in multiple years and Drits, A. V., E. G. Arashkevich, and T. N. Semenova. 1992. Pyro- the capacity to reach bloom proportions suggests that they soma atlanticum (Tunicata, ): grazing impact on phyto- may even thrive in colder waters, especially during more plankton standing stock and role in organic carbon flux. Journal of Plankton Research 14:799–809. oligotrophic conditions, and could become more permanent Graham, W. M., F. Pages, and W. M. Hamner. 2001. A physical con- residents in the California Current marine ecosystem. Their text for aggregations: a review. Hydrobi- continued presence will likely become a nuisance for certain ologia 451:199–212. fishing activities, causing fishers to relocate or spend extra Harbison, G. R. 1998. The parasites and predators of Thaliacea. time sorting their catch. Large pyrosome aggregations have Pages 187–214 in Q. Bone, editor. The biology of pelagic tunicates. the potential to restructure energy flows through food webs Oxford University Press, Oxford, UK. via efficient removal of photosynthetic plankton and subse- Lavaniegos, B. E., and M. D. Ohman. 2003. Long-term changes in pelagic tunicates of the California Current. Deep-Sea Research II quent fecal pellet production, consumption by higher 50:2473–2498. trophic levels, or sinking to depth. Lebrato, M., K. A. Pitt, A. K. Sweetman, D. O. Jones, J. E. Cartes, A. Oschlies, R. H. Condon, J. C. Molinero, L. Adler, and C. Gail- ACKNOWLEDGMENTS lard. 2012. Jelly-falls historic and recent observations: a review to – We thank the crew of the RV Bell Shimada and the RV Sikuliaq drive future research directions. Hydrobiologia 690:227 245. for all of their help. We thank Scott Marion for feedback and Linsey Mayzaud, P., M. Boutoute, R. Perissinotto, and P. Nichols. 2007. Isala for help with pyrosome identification. Funding for this work Polar and neutral lipid composition in the pelagic Pyro- – was provided by Oregon Sea Grant (K. R. Sutherland, A. W. Gal- soma atlanticum. Lipids 42:647 657. loway, R. D. Brodeur, H. L. Sorensen), the National Science Foun- Perissinotto, R., P. Mayzaud, P. D. Nichols, and J. P. Labat. dation (OCE-1737364 to K. R. Sutherland), the Northwest 2007. Grazing by Pyrosoma atlanticum (Tunicata Thaliacea) in Fisheries Science Center (R. D. Brodeur), and the NOAA Ernest F. the south Indian Ocean. Marine Ecology Progress Series 330: – Hollings Scholarship Program (O. N. Blondheim). 1 11. Sutherland, K. R., L. P. Madin, and R. Stocker. 2010. Filtration of submicrometer particles by pelagic tunicates. Proceedings of LITERATURE CITED the National Academy of Sciences USA 107:15129–15134. Van Soest, R. W. M. 1981. A monograph of the order Pyroso- Alldredge, A. L., and L. P. Madin. 1982. Pelagic tunicates: unique – matida (Tunicata, Thaliacea). Journal of Plankton Research 3: herbivores in the marine plankton. BioScience 32:655 663. 603–631. Andersen, V., and J. Sardou. 1994. Pyrosoma atlanticum (Tunicata, Thaliacea): diel migration and vertical distribution as a function 1,4 1 – KELLY R. SUTHERLAND , HILARIE L. SORENSEN, of colony size. Journal of Plankton Research 16:337 349. 2 3 Angel, M. V. 1989. Vertical profiles of pelagic communities in the OLIVIA N. BLONDHEIM, RICHARD D. BRODEUR , AND 1 vicinity of the Azores Front and their implications to deep ocean AARON W. E. GALLOWAY – ecology. Progress in Oceanography 22:1 46. 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E-mail: [email protected] Brodeur, R. D., R. I. Perry, J. L. Boldt, L. Flostrand, M. Galbraith, J. King, J. Murphy, K. Sakuma, and A. Thompson. 2018. An unu- sual gelatinous plankton event in the NE pacific: the great pyro- Additional supporting information may be found in the online ver- some bloom of 2017. PICES Press 26:22–28. sion of this article at http://onlinelibrary.wiley.com/doi/10.1002/ecy. Di Lorenzo, E., and N. Mantua. 2016. Multi-year persistence of the 2429/suppinfo 2014/15 North Pacific marine heatwave. Nature Climate Change 6:1042–1047.