Dol-LAYER and Dol-MICROBE Publications

Frischer, M.E., L.M. Lamboley, T.L. Walters, J.A. Brandes, E. Arneson, L.E. Lacy, N.B. López-Figueroa, A.E. Rodriguez-Santiago, D.M. Gibson (2021). Selective feeding and linkages to the microbial food web by the doliolid Dolioletta gegenbauri in the South Atlantic Bight. Limnology & Oceanography 66(5): 1993-2010. doi: 10.1002/lno.11740

Greer, A.T., L.M. Chiaverano, L.M. Treible, C. Briseño-Avena, F.J. Hernandez (2021). From spatial pattern to ecological process through imaging zooplankton interactions. ICES Journal of Marine Science 78(8):2664–2674. doi: 10.1093/icesjms/fsab149

Greer, A.T., M.S. Schmid, P.I. Duffy, K.L. Robinson, M.A. Genung, J.Y. Luo, T. Panaïotis, C. Briseño-Avena, M.E. Frischer, S. Sponaugle, R.K. Cowen (2023). In situ imaging across ecosystems to resolve the fine-scale oceanographic drivers of a globally significant planktonic grazer. Limnology & Oceanography 68(1):192-207. doi: 10.1002/lno.12259

Videos and Documents for Students/Teachers/Community

Doliolid selective feeding (October 2018)

What do doliolids eat for breakfast?

Georgia teachers participate in UGA research cruises

Frischer, M.E. (2019). The tiny animal at the centre of the marine ecosystem.

Sullivan, M. and M.E. Frischer. Tiny but all-consuming marine organism focus of UGA Skidaway Institute study. Skidaway News Archives. March 2017


Blogs from Teachers for “Rivers to Reefs”

Educators are nearly unanimous in agreement that participating in authentic research and working closely with active scientists are effective means of understanding the culture of science (PCAST 2012). To address this need, the Dol-LAYER and Dol-MICROBE projects are providing opportunities for K-12 educators to participate in research cruises and professional development workshops.

Dol-LAYER and Dol-MICROBE support two programs to provide these opportunities.

I. Opportunities to participate in research cruises. If you are interested in participating, please contact someone on our team.

II. Participate in the Gray’s Reef national Marine Sanctuary Foundation Rivers to Reefs Teacher Professional Development Workshop.

The R2R workshop is a six-day program designed to provide teachers with an opportunity to be active participants in the process of learning scientific concepts related to watershed processes. The workshop, approved by the State of Georgia for 4 Professional Learning Units, involves immersing teachers in a guided exploration of the Altamaha River system from its headwaters near Atlanta, GA out to the continental shelf. For more information about the R2R workshop, please visit:

The Dol-MICROBE project supported the 2021 R2R workshop offered June 24 – July 1, 2021. Each of the educator participants wrote a short blog about their experiences, along with photos of their activities.

Ben Wells

Dr. Karen D. Chassereau

Elspeth Pirrie

Linda Rule

Mary Thaler

Raegan Dillon

Kania Greer


Previous Publications about Doliolids from the Science Team

Frischer, M.E., C.A. Sanchez, T.L. Walters, M.E. Thompson, L.M. Frazier, G.A. Paffenhöfer (2014). Reliability of qPCR for quantitative gut content estimation in the circumglobally abundant pelagic tunicate Dolioletta gegenbauri (Tunicata, Thaliacea). Food Webs 1: 18-24. doi: 10.1016/j.fooweb.2014.11.001

Greer, A.T., A.D. Boyette, A.D., V.J. Cruz, M.K. Cambazoglu, B. Dzwonkowski, L.M. Chiaverano, S.L. Dykstra, C. Briseño-Avena, R.K. Cowen, J.D. Wiggert (2020). Contrasting fine-scale distributional patterns of zooplankton driven by the formation of a diatom-dominated thin layer. Limnology & Oceanography 65: 2236-2258. doi: 10.1002/lno.11450

López-Figueroa, N.B. (2017) Spatial and temporal trends of zooplankton communities in the South Atlantic Bight (SAB). MS Thesis. The Graduate College of Hampton University. ProQuest Number 10620671.

Walters, T.L., D.M. Gibson, M.E. Frischer (2019). Cultivation of the marine pelagic tunicate Dolioletta gegenbauri (Uljanin 1884) for experimental studies. Journal of Visualized Experiments 150: e59832, doi:10.3791/59832.

Walters, T.L., L.M. Lamboley, N.B. López-Figueroa, A.E. Rodríguez-Santiago, D.M. Gibson, M.E. Frischer (2018). Diet and trophic interactions of a circumglobally significant gelatinous marine zooplankter, Dolioletta gegenbauri (Uljanin, 1884). Molecular Ecology 28: 176-189 doi: 10.1111/mec.14926


Other Relevant Publications (Doliolids and more)

Alldredge, A.L. and L.P. Madin (1982). Pelagic tunicates, unique herbivores in the marine plankton. BioScience 32: 655-633.

Bone, Q. (1997) The biology of pelagic tunicates. Oxford University Press, Oxford; New York.

Conley, K.R., F. Lombard, K.R. Sutherland (2018). Mammoth grazers on the ocean’s minuteness, a review of selective feeding using mucous meshes. Proceedings of the Royal Society B. 285: 20180056.

Cowen, R.K. and C.M. Guigand. (2008). In situ ichthyoplankton imaging system (ISIIS): System design and preliminary results. Limnology & Oceanography-Methods 6: 126-132.

Daniels, C. and M. Breitbart (2012). Bacterial communities associated with the ctenophores Mnemiopsis leidyi and Beroe ovata. FEMS Microbiology Ecology 82: 90-101.

Deibel, D. and B. Lowen (2012). A review of the life cycles and life-history adaptations of pelagic tunicates to environmental conditions. ICES Journal of Marine Science 69: 358-369.

Deibel, D. and G.A.  Paffenhöfer (2009). Predictability of patches of neritic salps and doliolids (Tunicata, Thaliacea). Journal of Plankton Research 31: 1571-1579.

Deibel, D. (1990). Still-water sinking velocity of fecal material from the pelagic tunicate Dolioletta gegenbauri. Marine Ecology Progress Series 62: 55-60.

Deibel, D. (1998) The abundance, distribution and ecological impact of doliolids. In, B. Q (ed) The biology of pelagic tunicates. Oxford University Press, Oxford, pp. 171-186.

Deibel, D. (1985) Blooms of the pelagic tunicate, Dolioletta gegenbauri, Are they associated with Gulf Stream frontal eddies? Journal of Marine Research 43: 211-236.

Ellen, J.S., C.A. Graff, M.D. Ohman (2019). Improving plankton image classification using context metadata. Limnology & Oceanography-Methods 17: 439-461.

Faillettaz, R., M. Picheral, J.Y. Luo, C. Guigand, R.K. Cowen, J. Irisson (2016). Imperfect automatic image classification successfully describes plankton distribution patterns. Methods in Oceanography 15-16: 60-77.

Gibson, D. M. and G.A. Paffenhöfer, G. A. (2000) Feeding and growth rates of the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea). Journal of Plankton Research 22: 1485-1500.

Greer, A.T., C. Briseno-Avena, A.L. Deary, R.K. Cowen, F.J. Hernandez, W.M. Graham (2017). Associations between lobster phyllosoma and gelatinous zooplankton in relation to oceanographic properties in the northern Gulf of Mexico. Fisheries Oceanography 26: 693-704.

Greer, A.T., R.K. Cowen, C.M. Guigand, J.A. Hare (2015). Fine-scale planktonic habitat partitioning at a shelf-slope front revealed by a high-resolution imaging system. Journal of Marine Systems 142: 111-125.

Greer, A.T. and C.B. Woodson (2016). Application of a predator – prey overlap metric to determine the impact of sub-grid scale feeding dynamics on ecosystem productivity. ICES Journal of Marine Science 73: 1051-1061.

Iversen, M.H., E.A. Pakhomov, B.P.V. Hunt, H. van der Jagt, D. Wolf-Gladrow, C. Klass (2017). Sinkers or floaters? Contribution from salp pellets to the export flux during a large bloom event in the Southern Ocean. Deep-Sea Research II 38: 116-125.

Köster, M. and G.A. Paffenhöfer (2017). How efficiently can doliolids (Tunicata, Thaliacea) utilize phytoplankton and their own fecal pellets? Journal of Plankton Research 39: 305-315.

Köster, M., G.A. Paffenhöfer, R. Schlüter, A. Meuche (2014). Time-series observations of prokaryotic colonization of zooplankton fecal pellets. Journal of Plankton Research 36: 1451-1475.

Lebrato, M., Pahlow, M., Frost, J.R., Küter, Mendes, P.dJ., Molinero, J-C., Oschlies, A. (2019) Sinking of gelatinous zooplankton biomass increases deep carbon transfer efficiency globally. Global Biogeochemical Cycles 33: 1765-1783. doi: 10.1029/2019GB006265.

Lucas, C. and M. Dawson (2014). What Are Jellyfishes and Thaliaceans and Why Do They Bloom? Jellyfish Blooms pp. 9-44. 10.1007/978-94-007-7015-7_2.

Luo, J.Y., J.O. Irisson, B. Graham,  C. Guigand, A. Sarafraz, C. Mader,  R.K. Cowen (2018). Automated plankton image analysis using convolutional neural networks. Limnology & Oceanography-Methods 16: 814-827.

MacKenzie, K.M., C.N. Trueman, C.H. Lucas, J. Bortoluzzi (2017). The preparation of jellyfish for stable isotope analysis. Marine Biology 164: 219.

Madin L.P. and Deibel D. (1998) Feeding and energetics of Thaliacea. In, Bone Q (ed) The biology of pelagic tunicates. Oxford University Press, Oxford, pp 81–104.

Möller, K.O., M. St John, A. Temming, J. Floeter, A.F.  Sell, J. Herrmann, C. Möllmann (2012). Marine snow, zooplankton and thin layers: indications of a trophic link from small-scale sampling with the Video Plankton Recorder. Marine Ecology Progress Series 468: 57-69.

Nakamura, Y. (1998). Blooms of tunicates Oikopleura spp. and Dolioletta gegenbauri in the Seto Inland Sea, Japan, during summer. Hydrobiologia 385: 183-192.

Paffenhöfer, G.A. (2013). A hypothesis on the fate of blooms of doliolids (Tunicata, Thaliacea). Journal of Plankton Research 35: 919-924.

Paffenhöfer, G.A. and D.M. Gibson (1999). Determination of generation time and asexual fecundity of doliolids (Tunicata, Thaliacea). Journal of Plankton Research 21: 1183-1189.

Paffenhöfer, G. and Köster, M. (2005) Digestion of diatoms by planktonic copepods and doliolids. Marine Ecology Progress Series 297: 303-310.

Paffenhöfer, G.A, B. Wester, W. Nicholas (1984). Zooplankton abundance in relation to state and type of intrusions onto the southeastern united-states shelf during summer. Journal of Marine Research 42: 995-1017.

Paffenhöfer, G.A., L.P. Atkinson, T.N. Lee, P.G. Verity, L.R. Bulluck (1995). Distribution and abundance of thaliaceans and copepods off the southeastern USA during winter. Continental Shelf Research 15: 255-280.

Paffenhöfer, G.A., L. Atkinson, J. Blanton, T. Lee, L. Pomeroy, J. Yoder (1987). Summer upwelling on the southeastern continental-shelf of the USA during 1981 – summary and conclusions. Progress in Oceanography 19: 437-441.

Patonai, K., H. El-Shaffey, G.A. Paffenhöfer (2011). Sinking velocities of fecal pellets of doliolids and calanoid copepods. Journal of Plankton Research 33: 1146-1150.

Pomery, L.R. and D.  Deibel (1980). Aggregation of organic matter by pelagic tunicates. Limnology & Oceanography 25: 643-652.

Remsen, A., T.L Hopkins, S. Samson (2004). What you see is not what you catch, a comparison of concurrently collected net, Optical Plankton Counter, and Shadowed Image Particle Profiling Evaluation Recorder data from the northeast Gulf of Mexico. Deep Sea Research Part I Oceanographic Research Papers 51: 129-151.

Takahashi, K., T. Ichikawa, H. Saito, S. Kakehi, Y. Sugimoto, K. Hidaka, K. Hamasaki (2013). Sapphirinid copepods as predators of doliolids: Their role in doliolid mortality and sinking flux. Limnology & Oceanography 58: 1972-1984.

Takahashi, K., T. Ichikawa, C. Fukugama, M. Yamane, S. Kakehi, Y. Okazaki, H. Kubota, K. Furuya (2015). In situ observations of a doliolid bloom in a warm water filament using a video plankton recorder: Bloom development, fate, and effect on biogeochemical cycles and planktonic food webs. Limnology & Oceanography 60: 1763-1780.

Tebeau, C.M. and L.P. Madin (1994). Grazing rates for 3 life-history stages of the doliolid D. gegenbauri Uljanin (Tunicata, Thaliacea). Journal of Plankton Research 16: 1075-1081.