Research Program

Steve is working on deep-sea gelatinous zooplankton. He is doing research on bioluminescence, biodiversity, and ecology of deep-sea and open-ocean ctenophores, siphonophores, radiolarians, and medusae. In addition to assembling phylogenies for these groups, he is interested in cloning novel photoproteins and fluorescent proteins from these jellies.

Steve’s educational background includes a B.S. from Harvey Mudd College and a Ph.D. from the University of California, Santa Barbara.

He also holds an adjunct professorship in the Ecology and Evolutionary Biology department at the University of California, Santa Cruz.



  • Winnikoff, J.R., S.H.D. Haddock, I. Budin (2021) Depth- and temperature-specific fatty acid adaptations in ctenophores from extreme habitats. J. Exp. Biol. 224: jeb242800
  • Elor, A., Thang, T., Hughes, B.P., Crosby, A., Phung, A., Gonzalez, E., Katija, K., Haddock, S.H.D., Martin, E.J., Erwin, B.E., and Takayama, L. (2021). Catching Jellies in Immersive Virtual Reality: A Comparative Teleoperation Study of ROVs in Underwater Capture Tasks. Proc. 27th ACM Symposium on Virtual Reality Software and Technology. Article17:1-10.
  • Christianson, L.M., S.B. Johnson, D.T. Schultz, S.H.D. Haddock (2021) Hidden diversity of Ctenophora revealed by new mitochondrial COI primers and sequences. Molec. Ecol. Resources.
  • Schultz, D.T., W.R. Francis, J. McBroome, L.C. Chri
  • Damian-Serrano, A., S.H.D. Haddock, C.W. Dunn (2021) The evolutionary history of siphonophore tentilla: novelties, convergence, and integration. Integrative Organismal Biol. vol 3. pbab019
  • Meech, M.E., C.E. Mills, S.H.D. Haddock, R.W. Meech. (2021). Escape swimming in hydromedusae. J. Exp.
  • Stefanoudis, P. V., L.M. Biancani, S. Cambronero-Solano, M.R. Clark, J.T. Copley, E. Easton, F. Elmer, S.H.D. Haddock, S. Herrera, I.S. Iglesias, A.M. Quattrini, J. Sigwart, C. Yesson, A.G. Glover. (2021) Moving conferences online: lessons learned from an international virtual meeting. Proc. R. Soc. B. 20211769.
  • Damian-Serrano, A., S.H.D. Haddock, C.W. Dunn (2021). The evolution of siphonophore tentilla for specialized prey capture in the open ocean. Proc. Nat. Acad. Sci. 118 (8) e2005063118 (cover)
  • Puente-Tapia, F. A., R. Gasca, A. Schiariti, S.H.D. Haddock (2021) An updated checklist of ctenophores (Ctenophora: Nuda and Tentaculata) of Mexican seas. Regional Studies in Marine Science. 41:101555
  • Gibbons, M.J., S.H.D. Haddock, G.I. Matsumoto, C. Foster. (2021) Records of ctenophores from South Africa. PeerJ. 9:e10697
  • Miller, S.D., Haddock, S.H.D., Straka, W.C., Seaman, C.J., Combs, C.L., Wang, M., Shi, W., & Nam S.H. (2021). Honing in on bioluminescent milky seas from space. Sci Rep, 11, 15443.


  • Haddock, S.H.D. and C. Anela Choy (2020) Treasure and turmoil in the deep sea. New York Times.
  • Schultz, D.T., J.M Eizenga, R.B. Corbett-Detig, W.R. Francis, L.M. Christianson, S.H.D. Haddock. (2020). Conserved novel ORFs in the mitochondrial genome of the ctenophore Beroe forskalii. PeerJ. 8:e8356.
  • Drazen, J.C., C.R. Smith, K.M. Gjerde, S.H.D. Haddock, et al. (2020). Midwater ecosystems must be considered when evaluating environmental risks of deep-sea mining. Proc. Nat. Acad. Sci.
  • Bessho-Uehara, M., W.R. Francis, S.H.D. Haddock (2020). Biochemical characterization of diverse deep-sea anthozoan bioluminescence systems. Marine Biology 167: 114.
  • Bessho-Uehara, M., W. Huang, W.L. Patry, W.E. Browne, J.-K. Weng, S.H.D. Haddock (2020). Evidence for de novo Biosynthesis of the Luminous Substrate Coelenterazine in Ctenophores, iScience. doi: 10.1016/j.isci.2020.101859.
  • Martini, S., D.T. Schultz, L. Lundsten, S.H.D. Haddock (2020). Bioluminescence in an undescribed species of carnivorous sponge (Cladorhizidae) from the deep sea, Front. Mar. Sci.
  • Puente-Tapia, F.A., R. Gasca, A. Schiariti, S.H.D. Haddock (2020). An updated checklist of ctenophores (Ctenophora: Nuda and Tentaculata) of Mexican seas, Regional Studies in Marine Science. doi: 10.1016/j.rsma.2020.101555.
  • Leonardi, N.D., E.V. Thuesen, S.H.D. Haddock (2020). A sticky thicket of glue cells: A comparative morphometric analysis of colloblasts in 20 species of comb jelly (phylum Ctenophora), Ciencias Marinas, 46:4.


  • Winnikoff, J.R., W. R. Francis, E.V. Thuesen, and S.H.D. Haddock. (2019) Combing transcriptomes for secrets of deep-sea survival: Environmental diversity drives patterns of protein evolution. Integrative and Comparative Biology, pp 1-13. PDF.
  • Messié, M., I. Shulman, S. Martini, and S.H.D. Haddock. (2019) Using fluorescence and bioluminescence sensors to characterize auto- and heterotrophic plankton communities. Progress in Oceanography 171: 76-92.
  • Martini, S., Kuhnz, L., Mallefet, J., and S.H.D. Haddock. (2019) Distribution and quantification of bioluminescence as an ecological trait in the deep sea benthos. Scientific Reports 9: 14654.


  • Munro, C.,S. Siebert, F.. Zapata, M. Howison, A. Damian-Serrano, S. Church, F.E. Goetz, P,R, Pugh, S.H.D. Haddock, C.W. Dunn (2018) Improved phylogenetic resolution within Siphonophora (Cnidaria) with implications for trait evolution. Mol. Phylo. Evol. 127:823-833,
  • Pugh, P.R., C.W. Dunn, and S.H.D. Haddock (2018) A description of Tottonophyes enigmatica gen. nov., sp. nov. in a new family of calycophoran siphonophores (Hydrozoa), with notes on calycophoran canals. Zootaxa 4415: 452-472. PDF.
  • Winnikoff, J.W., T. M. Wilson, E.V. Thuesen, and S.H.D. Haddock (2017) Enzymes feel the squeeze: biochemical adaptation to pressure in the deep sea. The Biochemist. 39: 26-29. PDF.
  • Haddock, S.H.D., L.M. Christianson, W.R. Francis, S. Martini, C.W. Dunn, P.R. Pugh, C.E. Mills, K.J. Osborn, B.A. Seibel, C.A. Choy, C.E. Schnitzler, G.I. Matsumoto, M. Messié, D.T. Schultz, J.R. Winnikoff, M.L. Powers, R. Gasca, W.E. Browne, S. Johnsen, K.L. Schlining, S. von Thun, B.E. Erwin, J.F. Ryan, and E.V. Thuesen. 2017. Insights into the biodiversity, behavior, and bioluminescence of deep-sea organisms using molecular and maritime technology. Oceanography 30(4): 38–47, PDF.
  • Schultz, D.T., A.A. Kotlobay, R. Ziganshin, A. Bannikov, N.M. Markina, T.V. Chepurnyh, E.S. Shakhova, K. Palkina, S.H.D. Haddock, I.V. Yampolsky, Y. Oba (2018). Luciferase of the Japanese syllid polychaete Odontosyllis undecimdonta. Biochem. Biophys. Res. Comm. 502: 318–23.
  • Babonis, L., M.B. Debiasse, W.R. Francis, L.M. Christianson, S.H.D. Haddock, M.Q. Martindale, J.F. Ryan. (2018) Integrating embryonic development and evolutionary history to characterize tentacle-specific cell types in a ctenophore. Molecular Biol. Evol.


  • Hoving, H.J.T., S.L. Bush, S.H.D. Haddock, B.H. Robison (2017) Bathyal feasting: post-spawning squid as a source of carbon for deep-sea communities. Proc. Roy. Soc. B. 284: 20172096
    Choy CA, Haddock SHD, Robison BH. 2017 Deep pelagic food web structure as revealed by in situ feeding observations. Proc. R. Soc. B 284: 20172116.
  • Francis, W.R., Christianson, L.M., and S.H.D. Haddock (2017). Symplectin evolved from multiple duplications in bioluminescent squid. PeerJ. doi:10.7717/peerj.3633
  • Hoving, H.J.T. and S.H.D. Haddock (2017). The giant deep-sea octopus Haliphron atlanticus forages on gelatinous fauna. Nature Scientific Reports. doi:10.1038/srep44952.
    Martini, S. and S.H.D. Haddock (2017). Quantification of bioluminescence from the surface to the deep sea demonstrates its predominance as an ecological trait. Nature Scientific Reports. doi:10.1038/srep45750.


  • Gasca, R. and S.H.D. Haddock (2016). The rare deep-living hyperiid amphipod Megalanceoloides remipes (Barnard, 1932): complementary description and symbiosis. Zootaxa. 4178: 138. doi: 10.11646/zootaxa.4178.1.7.
  • Fossette, S., Katija, K., Goldbogen, J.A., Bograd, S., Patry W., Howard, M.J., Knowles, T., Haddock, S.H.D., Bedell, L., Hazen, E.L., Robison, B.H., Mooney, T.A., Shorter, K.A., Bastian, T., Gleiss, A.C. (2016) How to tag a jellyfish? A methodological review and guidelines to successful jellyfish tagging. J. Plankton Res. doi:10.1093/plankt/fbw073.
  • Francis, W.R., Powers, M.L., and S.H.D. Haddock (2016). Bioluminescence spectra from three deep‑sea polychaete worms. Mar. Biol. 163:255. doi:10.1007/s00227-016-3028-2
  • Francis, W.R., Christianson, L.M., Powers, M.L., Schnitzler, C.E., Haddock, S.H. (2016) Non-excitable fluorescent protein orthologs found in ctenophores. BMC Evol Biol 16, 167. doi:10.1186/s12862-016-0738-5.
  • Pugh, P.R. and S.H.D. Haddock (2016). A description of two new species of the genus Erenna (Siphonophora, Physonectae, Erennidae), with notes on recently collected specimens of other Erenna species. Zootaxa. 4189: 401-446. doi:10.11646/zootaxa.4189.3.1.


  • Dunn, C. W., Leys, S. P., & Haddock, S. H. D. (2015). The hidden biology of sponges and ctenophores. Trends in Ecology & Evolution, 30, 282–291. PDF. New Yorker article.
  • Francis, W. R., Shaner, N. C., Christianson, L. M., Powers, M. L., & Haddock, S. H. D. (2015). Occurrence of Isopenicillin-N-Synthase Homologs in Bioluminescent Ctenophores and Implications for Coelenterazine Biosynthesis. PLoS ONE, 10, e0128742. doi:10.1371/journal.pone.0128742. PDF.
  • Gasca, R., Hoover, R., & Haddock, S. H. D. (2015). New symbiotic associations of hyperiid amphipods (Peracarida) with gelatinous zooplankton in deep waters off California. Journal of the Marine Biological Association of the United Kingdom, 95, 503–511. doi:10.1017/S0025315414001416.
  • Gasca, R., Suárez-Morales, E., & Haddock, S. H. D. (2015). Sapphirina iris Dana, 1849 and S. Sinuicauda Brady, 1883 (Copepoda, Cyclopoida): predators of salps in Monterey Bay and the Gulf of California. Crustaceana, 88, 689–699. doi:10.1163/15685403-00003438.
  • Haddock, S. H. D., & Dunn, C. W. (2015). Fluorescent proteins function as a prey attractant: experimental evidence from the hydromedusa Olindias formosus and other marine organisms. Biology Open. doi:10.1242/bio.012138. Supplemental video.
  • Siebert, S., Goetz, F. E., Church, S. H., Bhattacharyya, P., Zapata, F., Haddock, S. H. D., & Dunn, C. W. (2015). Stem cells in Nanomia bijuga (Siphonophora), a colonial animal with localized growth zones. Evodevo, 6, 22. doi:10.1186/s13227-015-0018-2. PDF.
  • Zapata, F., Goetz, F. E., Smith, S. A., Howison, M., Siebert, S., Church, S. H., … Cartwright, P. (2015). Phylogenomic analyses support traditional relationships within Cnidaria. PLoS ONE, 10, e0139068. doi:10.1371/journal.pone.0139068.


  • Haddock, S.H.D (2014). Letter to the Bulletin editors: A plea for meaningful colors in ODV, Limnology and Oceanography Bulletin, 19:2. doi: 10.1002/lob.201019244. PDF.
  • Cazenave, F., Kecy, C., Risi, M., & Haddock, S. H. D. (2014). SeeStar: A low-cost, modular and open-source camera system for subsea observations (p. 7 pp.). Presented at the Proceedings of Marine Technology Society/Institute for Electrical and Electronics Engineers Oceans 2014. doi:10.1109/OCEANS.2014/7003077.
  • Francis, W. R., Powers, M. L., & Haddock, S. H. D. (2014). Characterization of an anthraquinone fluor from the bioluminescent, pelagic polychaeteTomopteris. Luminescence, 29, 1135–1140. doi:10.1002/bio.2671. PDF.
  • Global Invertebrate Genomics Alliance Community of Scientists, & (S. H.D. Haddock is primary member of consortium group). (2014). The Global Invertebrate Genomics Alliance (GIGA): Developing community resources to study diverse invertebrate genomes. Journal of Heredity, 105, 1–18. doi:10:1093/jhered/est084. PDF.
  • Wilson, G., Aruliah, D. A., Brown, C. T., Hong, N. P. C., Davis, M., Guy, R. T., … Wilson, P. (2014). Best practices for scientific computing. PLoS Biology,12, e1001745. doi:10.1371/journal.pbio.1001745. PDF.


  • Francis, W. R., Christianson, L. M., Kiko, R., Powers, M. L., Shaner, N. C., & Haddock, S. H. D. (2013). A comparison across non-model animals suggests an optimal sequencing depth for de novo transcriptome assembly. BMC Genomics, 14, 167. doi:10.1186/1471-2164-14-167. PDF.
  • Hurt, C., Haddock, S. H. D., & Browne, W. E. (2013). Molecular phylogenetic evidence for the reorganization of the Hyperiid amphipods, a diverse group of pelagic crustaceans. Molecular Phylogenetics and Evolution, 67, 28–37. doi:10.1016/j.ympev.2012.12.021.
  • Madin, L. P., Hamner, W. M., Haddock, S. H. D., & Matsumoto, G. I. (2013). Scuba diving in blue water: A window on ecology and evolution in the epipelagic ocean (In: Research and Discovery: The Revolution of Science through Scuba) (pp. pp. 71–82). Smithsonian Institution, Washington D.C. PDF.
  • Ryan, J. F., Pang, K., Schnitzler, C. E., Nguyen, A.-D., Moreland, R. T., Simmons, D. K., … Baxevanis, A. D. (2013). The genome of the ctenophore Mnemiopsis leidyi and its implications for cell type evolution. Science, 342, 1336–1344. doi:10.1126/science.1242592. PDF.
  • Siebert, S., Pugh, P. R., Haddock, S. H. D., & Dunn, C. W. (2013). Re-evaluation of characters in Apolemiidae (Siphonophora), with description of two new species from Monterey Bay, California. Zootaxa, 3702, 201–232. doi:10.11646/zootaxa.3702.3. PDF.
  • Thuesen, E. V., & Haddock, S. H. D. (2013). Archeterokrohnia docrickettsae (Chaetognatha: Phragmophora: Heterokrohniidae), a new species of deep-sea arrow worm from the Gulf of California. Zootaxa, 3717, 320–328. doi:10.11646/zootaxa.3717.3.2. PDF.


  • Condon, R. H., Duarte, C. M., Pitt, K. A., Robinson, K. L., Lucas, C. H., Sutherland, K. R., … Graham, W. M. (2012). Recurrent jellyfish blooms are a consequence of global oscillations. Proceedings of the National Academy of Sciences of the United States of America, 110, 1000–5. doi:10.1073/pnas.1210920110. PDF.
  • Condon, R. H., Graham, W. M., Duarte, C. M., Pitt, K. A., Lucas, C. H., Haddock, S. H. D., … Madin, L. P. (2012). Questioning the rise of gelatinous zooplankton in the world’s oceans. BioScience, 62, 160–169. doi:10.1525/bio.2012.62.2.9.
  • Johnsen, S., Frank, T. M., Haddock, S. H. D., Widder, E. A., & Messing, C. G. (2012). Light and vision in the deep-sea benthos: I. Bioluminescence at 500–1000 m depth in the Bahamian Islands. Journal of Experimental Biology, 215, 3335–3343. doi:10.1242/jeb.072009. PDF.
  • Powers, M. L., McDermott, A. G., Shaner, N., & Haddock, S. H. D. (2012). Expression and characterization of the calcium-activated photoprotein from the ctenophore Bathocyroe fosteri: Insights into light-sensitive photoproteins. Biochemical and Biophysical Research Communications. doi:10.1016/j.bbrc.2012.12.026. PDF.
  • Schnitzler, C. E., Pang, K., Powers, M. L., Reitzel, A. M., Ryan, J. F., Simmons, D., … Baxevanis, A. D. (2012). Genomic organization, evolution, and expression of photoprotein and opsin genes in Mnemiopsis leidyi: a new view of ctenophore photocytes. BMC Biology, 10. doi:10.1186/1741-7007-10-107. PDF.
  • Shulman, I., Penta, B., Moline, M. A., Haddock, S. H. D., Anderson, S., Oliver, M. J., & Sakalaukus, P. (2012). Can vertical migrations of dinoflagellates explain observed bioluminescence patterns during an upwelling event in Monterey Bay, California? Journal of Geophysical Research, 117, C01016. doi:10.1029/2011JC007480. PDF.


  • Osborn, K. J., Haddock, S. H. D., & Rouse, G. W. (2011). Swima (Annelida, Acrocirridae), holopelagic worms from the deep Pacific. Zoological Journal of the Linnean Society, 163, 663–678. doi:10.1111/j.1096-3642.2011.00727.x. PDF.
  • Shulman, I., Moline, M. A., Penta, B., Anderson, S., Oliver, M., & Haddock, S. H. D. (2011). Observed and modeled bio-optical, bioluminescent, and physical properties during a coastal upwelling event in Monterey Bay, California. Journal of Geophysical Research-Oceans, 116. doi:10.1029/2010JC006525. PDF.


  • Bayha, K. M., Dawson, M. N., Collins, A. G., Barbeitos, M. S., & Haddock, S. H. D. (2010). Evolutionary relationships among scyphozoan jellyfish families based on complete taxon sampling and phylogenetic analyses of 18S and 28S ribosomal DNA. Integrative and Comparative Biology, 50, 436–455. doi:10.1093/icb/icq074. PDF.
  • Haddock, S. H. D. (2010). A plea for meaningful colors in ODV. Limnology and Oceanography Bulletin, 19, 44–45. PDF.
  • Haddock, S. H. D., & Dunn, C. W. (2010). Practical Computing for Biologists. Sinauer Associates.
  • Haddock, S. H. D., Mastroianni, N., & Christianson, L. M. (2010). A photoactivatable green-fluorescent protein from the phylum Ctenophora. Proceedings of the Royal Society B, 277, 1155–1160. doi:10.1098/rspb.2009.1774. PDF. Retracted.
  • Haddock, S. H. D., Moline, M. A., & Case, J. F. (2010). Bioluminescence in the sea. Annual Review of Marine Science, 2, 443–493. doi:10.1146/annurev-marine-120308-081028. PDF.
  • Pugh, P. R., & Haddock, S. H. D. (2010). Three new species of resomiid siphonophores (Siphonophora, Physonectae). Journal of the Marine Biological Association of the United Kingdom, 90, 1119–1143. PDF.
  • Thuesen, E. V., Goetz, F. E., & Haddock, S. H. D. (2010). Bioluminescent organs of two deep-sea arrow worms, Eukrohnia fowleri and Caecosagitta macrocephala, with further observations on bioluminescence in chaetognaths. Biological Bulletin, 219, 100–111. PDF.


  • Gorokhova, E., Lehtiniemi, M., Viitasalo-Frösen, S., & Haddock, S. H. D. (2009). Molecular evidence for the occurrence of ctenophore Mertensia ovum in the northern Baltic Sea and implications for current distribution of the invasive species Mnemiopsis leidyi. Limnology and Oceanography, 54, 2025–2033. PDF.
  • Moline, M. A., Blackwell, S. M., Case, J. F., Haddock, S. H. D., Herren, C. M., Orrico, C. M., & Terrill, E. (2009). Bioluminescence to reveal structure and interaction of coastal planktonic communities. Deep-Sea Research Part II, 56, 232–245. doi:10.1016/j.dsr2.2008.08.002. PDF.
  • Osborn, K. J., Haddock, S. H. D., Pleijel, F., Madin, L. P., & Rouse, G. W. (2009). Deep-sea, swimming worms with luminescent “bombs.” Science, 325, 964. doi:10.1126/science.1172488. PDF.


  • Collins, A. G., Bentlage, B., Lindner, A., Lindsay, D., Haddock, S. H. D., Jarms, G., … Cartwright, P. (2008). Phylogenetics of Trachylina (Cnidaria: Hydrozoa) with new insights on the evolution of some problematical taxa. Journal of the Marine Biological Association of the United Kingdom, 88, 1673–1685. doi:10.1017/S0025315408001732. PDF.
  • Dunn, C. W., Hejnol, A., Matus, D. Q., Pang, K., Browne, W. E., Smith, S. A., … Giribet, G. (2008). Broad phylogenomic sampling improves resolution of the animal tree of life. Nature, 452, 745–9. doi:10.1038/nature06614. PDF.
  • Haddock, S. H. D. (2008). Reconsidering evidence for potential climate-related increases in jellyfish. Limnology and Oceanography, 53, 2759–2762. PDF.
  • Schnitzler, C. E., Keenan, R. J., McCord, R., Matysik, A., Christianson, L. M., & Haddock, S. H. D. (2008). Spectral diversity of fluorescent proteins from the anthozoan Corynactis californica. Marine Biotechnology, 10, 328–342. doi:10.1007/s10126-007-9072-7. PDF.
  • Staaf, D. J., Camarillo-Coop, S., Haddock, S. H. D., Nyack, A. C., Payne, J., Ramirez-Rojo, R., … Gilly, W. F. (2008). Natural egg mass deposition by the Humboldt squid (Dosidicus gigas) in the Gulf of California and characteristics of hatchlings and paralarvae. Journal of the Marine Biological Association of the United Kingdom, 88, 759–770. doi:10.1017/S0025315408001422. PDF.


  • Browne, W. E., Haddock, S. H. D., & Martindale, M. Q. (2007). Phylogenetic analysis of lineage relationships among hyperiid amphipods as revealed by examination of the mitochondrial gene, cytochrome oxidase I (COI). Integrative and Comparative Biology, 47, 815–830. doi:10.1093/icb/icm093. PDF.
  • Haddock, S. H. D. (2007). Comparative feeding behavior of planktonic ctenophores. Integrative and Comparative Biology, 47, 847–853. doi:10.1093/icb/icm088. PDF.
  • Mills, C. E., & Haddock, S. H. D. (2007). Key to the ctenophora. In J. T. Carlton (Ed.), Light and Smith’s Manual: Intertidal invertebrates of the central California coast (pp. 189–199). University of California Press.
  • Mills, C. E., Haddock, S. H. D., Dunn, C. W., & Pugh, P. R. (2007). Key to the siphonophora. In J. T. Carlton (Ed.), Light and Smith’s Manual: Intertidal invertebrates of the central California coast (pp. 150–166). University of California Press.
  • Sweeney, A. M., Haddock, S. H. D., & Johnsen, S. (2007). Comparative visual acuity of coleolid cephalopods. Integrative and Comparative Biology, 47, 808–814. PDF.


  • Collins, A. G., Bentlage, B., Matsumoto, G. I., Haddock, S. H. D., Osborn, K. J., & Schierwater, B. (2006). Solution to the phylogenetic enigma ofTetraplatia, a worm-shaped cnidarian. Biological Letters, 2, 120–124. doi:10.1098/rsbl.2005.0372. PDF.
  • Gasca, R., Suárez-Morales, E., & Haddock, S. H. D. (2006). Symbiotic associations between crustaceans and gelatinous zooplankton in deep and surface waters off California. Marine Biology, 151, 233–242. PDF.
  • Haddock, S. H. D. (2006). Luminous marine organisms. In S. Daunert & S. K. Deo (Eds.), Photoproteins in Bioanalysis (pp. 25–47). New York: Wiley-VCH.
  • Miller, S. D., Haddock, S. H. D., Elvidge, C. D., & Lee, T. F. (2006). Twenty thousand leagues over the seas: The first satellite perspective on bioluminescent “milky seas.” International Journal of Remote Sensing, 27, 5131–5143. doi:10.1080/01431160600554298.


  • Dunn, C. W., P. R. Pugh, & Haddock, S. H. D. (2005). Marrus claudanielis, a new species of deep-sea physonect siphonophore (Siphonophora, Physonectae). Bulletin of Marine Science, 76, 699–714. PDF.
  • Dunn, C. W., Pugh, P. R., & Haddock, S. H. D. (2005). Molecular Phylogenetics of the Siphonophora (Cnidaria), with implications for the evolution of functional specialization. Systematic Biology, 54, 916–935. PDF.
  • Haddock, S. H. D., & Dunn, C. W. (2005). The complex world of siphonophores. JMBA Global Marine Environment, 2, 24–25. PDF.
  • Haddock, S. H. D., Dunn, C. W., & Pugh, P. R. (2005). A reexamination of siphonophore terminology and morphology, applied to the description of two new prayine species with remarkable bio-optical properties. Journal of the Marine Biological Association of the United Kingdom, 85, 695–707. PDF.
  • Haddock, S. H. D., Dunn, C. W., Pugh, P. R., & Schnitzler, C. E. (2005). Bioluminescent and red-fluorescent lures in a deep-sea siphonophore. Science, 309, 263. doi:10.1126/science.1110441. PDF.
  • Haddock, S. H. D., & Heine, J. N. (2005). Scientific blue-water diving. California Sea Grant, La Jolla. PDF.
  • Herren, C. M., Haddock, S. H. D., Johnson, C., Moline, M. A., Orrico, C. M., & Case, J. F. (2005). A multi-platform bathyphotometer for fine-scale, coastal bioluminescence research. Limnology and Oceanography: Methods, 3, 247–262. PDF.
  • Miller, S. D., Haddock, S. H. D., Elvidge, C. D., & Lee, T. F. (2005). Detection of a bioluminescent milky sea from space. Proceedings of the National Academy of Sciences, 102, 14181–14184. doi:10.1073/pnas.0507253102. PDF.
  • Seibel, B. A., Robison, B. H., & Haddock, S. H. D. (2005a). First observations of brooding in a squid. Nature, 438, 929. doi:10.1038/438929a. PDF.
  • Seibel, B. A., Robison, B. H., & Haddock, S. H. D. (2005b). Post-spawning egg care by a squid. Nature, 438, 929.
  • Shulman, I., McGillicuddy Jr., D. J., Moline, M. A., Haddock, S. H. D., Kindle, J. C., Nechaev, D., & Phelps, M. W. (2005). Bioluminescence intensity modeling and sampling strategy optimization. Journal of Atmospheric and Oceanic Technology, 22, 1267–1281. PDF.
  • Widder, E. A., Robison, B. H., Reisenbichler, K. R., & Haddock, S. H. D. (2005). Using red light for in situ observations of deep-sea fishes. Deep-Sea Research Part I, 52, 2077–2085. doi:10.1016/j.dsr.2005.06.007. PDF.


  • Gasca, R., & Haddock, S. H. D. (2004). Associations between gelatinous zooplankton and hyperiid amphipods (Crustacea: Peracarida) in the Gulf of California. Hydrobiologia, 530/531, 529–535. PDF.
  • Haddock, S. H. D. (2004). A golden age of gelata: past and future research on planktonic ctenophores and cnidarians. Hydrobiologia, 530, 549–556. doi:10.1007/s10750-004-2653-9. PDF.


  • Robison, B. H., Reisenbichler, K. R., Hunt, J. C., & Haddock, S. H. D. (2003). Light Production by the Arm Tips of the Deep-Sea Cephalopod Vampyroteuthis infernalis. The Biological Bulletin, 205, 102–109. PDF.
  • Shulman, I., Haddock, S. H. D., McGillicuddy, J., Paduan, J. D., & Bissett, P. W. (2003). Numerical modeling of bioluminescence distributions in the coastal ocean. Journal of Atmospheric and Oceanic Technology, 20, 1060–1068. doi:10.1175/1468.1. PDF.

Other Publications

  • Haddock, S. H. D. (2011). Photographing your jellyfish. Make, 27, 91.
  • Haddock, S. H. D. (2013). The other Jellyfish. Catamaran Literary Reader, Fall issue, 25–29

Older Publications

  • Cairns, S. D., D. R. Calder, A. Brinckmann-Voss, C. B. Castro, D. G. Fautin, P. R. Pugh, … D. M. Opresko. (2002). Common and scientific names of aquatic invertebrates from the United States and Canada: Cnidaria and Ctenophora – Second Edition. Bethesda, Maryland: American Fisheries Society Special Publication #28.
  • Case, J. F., P. J. Herring, B. H. Robison, S. H.D. Haddock, L. J. Kricka, & P. E. Stanley. (2001). Bioluminescence and Chemiluminescence 2000. Singapore: World Scientific. PDF.
  • Haddock, S. H. D., Rivers, T. J., & Robison, B. H. (2001). Can coelenterates make coelenterazine? Dietary requirement for luciferin in cnidarian bioluminescence. Proceedings of the National Academy of Sciences of the United States of America, 98, 11148–11151. doi:10.1073/pnas.201329798. PDF.
  • Haddock, S. H. D., & Case, J. F. (1999). Bioluminescence spectra of shallow and deep-sea gelatinous zooplankton: ctenophores, medusae and siphonophores. Marine Biology, 133, 571–582. doi:10.1007/s002270050497. PDF.
  • Podar, M., Haddock, S. H. D., Sogin, M. L., & Harbison, G. R. (2001). A molecular phylogenetic framework for the phylum Ctenophora using 18S rRNA genes. Molecular Phylogenetics and Evolution, 21, 218–230. doi:10.1006/mpev.2001.1036. PDF.
  • Alldredge, A. L., Passow, U., & Haddock, S. H. D. (1998). The characteristics and transparent exopolymer particle (TEP) content of marine snow formed from thecate dinoflagellates. Journal of Plankton Research, 20, 393–406. doi:10.1093/plankt/20.3.393. PDF.
  • Haddock, S. H. D., Neilson, D. J., Widder, E. A., & Case, J. F. (1998). Feasibility of using in situ measurements of bioluminescence spectra to determine the vertical distribution of plankton (Pelagic biogeography ICoPB II: Proceedings of the 2nd international conference: final report of SCOR/IOC Working Group 93 “Pelagic Biogeography”: Noordwijkerhout, The Netherlands, 9 July-14 July 1995) (pp. 137–142). Paris, France: UNESCO. PDF.
  • White, G. B., Pfahnl, A., Haddock, S. H. D., Lamers, S., Greenberg, R. M., & Anderson, P. A. (1998). Structure of a putative sodium channel from the sea anemone Aiptasia pallida. Invert Neurosci, 3, 317–26. PDF.
  • Hsin, Y. L., & Haddock, S. H. D. (1997). The enclosing latticed sphere of Tuscaridium cygneum (Murray), a eurybathyal phaeodarian Radiolaria, from the North Pacific. Paleontological Research, 1, 144–149. PDF.
  • Mills, C. E., Pugh, P. R., Harbison, G. R., & Haddock, S. H. D. (1996). Medusae, siphonophores and ctenophores of the Alborán Sea, south western Mediterranean. Scientia Marina, 60, 145–163. PDF.
  • Haddock, S. H. D., & Case, J. F. (1995). Not All Ctenophores Are Bioluminescent: Pleurobrachia. The Biological Bulletin, 189, 356–362. PDF.
  • Case, J. F., Haddock, S. H. D., & Harper, R. D. (1994). The ecology of Bioluminescence. In Campbell, Kricka, & Stanley (Eds.), (pp. 115–122). Presented at the 8th International Symposium on Bioluminescence and Chemiluminescence, John Wiley and Sons. PDF.
  • Haddock, S. H. D., & Case, J. F. (1994). A bioluminescent chaetognath. Nature, 367, 225–226. PDF.
  • Herring, P. J., Widder, E. A., & Haddock, S. H. D. (1992). Correlation of bioluminescence emissions with ventral photophores in the mesopelagic squid Abralia veranyi (Cephalopoda: Enoploteuthidae). Marine Biology, 112, 293–298. doi:10.1007/BF00702474. PDF.
  • Haddock, S. H. D. (1989). Air pockets – a subtidal habitat for the intertidal marine Pulmonate limpet Trimusculus reticulatus. Veliger, 32, 403–405. PDF.


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