Our research focuses on how invertebrate animals develop, how they function, and how they have evolved. Specific projects often focus on the ecology and evolution of embryos and larval stages, especially those of annelids (like this scaleworm larva) and echinoderms, but also include a diversity of other topics in invertebrate biology. Look at our publications to get a better sense of our interests.

If interested in doing research in the lab, please contact me (the lab PI, Bruno Pernet)! I am always happy to speak with potential graduate students from around the U.S. and the world, as well as CSULB undergraduates. We can discuss project ideas you’ve come up with based on your coursework or field or lab observations, or I can suggest projects that I think are interesting. We are equipped to answer questions with descriptive and experimental techniques including larval culture, light and electron microscopy, video microscopy, basic DNA analyses, and field work; many more techniques are accessible by collaboration with other CSULB faculty, or faculty at nearby institutions.


Bruno Pernet. B.A. Biology, University of California Santa Cruz (1991); Ph.D. Zoology, University of Washington (1998). Faculty in the Department of Biological Sciences at California State University Long Beach since 2004.

Ariel Gilligan. Effects of salinity on metabolism and planktonic duration of larvae of the euryhaline annelid Ficopomatus enigmaticus.

Alex Mendelson. Do larvae of the annelid Ficopomatus enigmaticus fit the “founder vs. aggregator” model of settlement?
Keomonyroth Nuon. Do larvae of the annelid Ficopomatus enigmaticus fit the “founder vs. aggregator” model of settlement?

Peter Nilsson. Effects of natural assemblages of large particles on feeding and time to competence in echinoderm larvae.
Bridget Steiner. Constructional and functional differences between pluteus and dipleurula larval forms.
Alison Yee (M.S. 2019). Limits to the distribution of the invasive annelid Ficopomatus enigmaticus. PDF
David Lizárraga (M.S. 2017). Effects of large inedible particles on the feeding performance of echinoderm larvae. PDF
Caleb Jones (M.S. 2015). Evolutionary changes in development associated with a transition in larval nutritional mode in spiralians. PDF
Bruno Passarelli (M.S. 2010). The marine live bait trade in California: a pathway for introduction of non-indigenous species? PDF
Ellen Kosman (M.S. 2008). Patterns and consequences of variation in per-offspring investment in four bryozoan species. PDF

Sarah Anderson, Amanda Bell, Jeffrey Chang, Jimmy Chapman, Claire Cockroft, Andrea Danihel, Aimee Deconinck, Jenny Drechsler, Nichole Gatten, Truc Hoang, Jeanette Hofstee, Ali Krajewski, Valerie Langland, Angela Llaban, Veronica Madril, Christine March, Lynn McArthur, Amberle McKee, Helen Nguyen, Veronica Oria, Jessica Peria, Leah Sloan, Caitlin Sojka, Amber Von Tungeln, Marissa Velarde, Kelley Voss, David Wang


#Bolded names are CSULB student coauthors.
#Nilsson, P., and B. Pernet. 2022. Echinoid larvae can express food-conditioned morphological plasticity at ecologically relevant culture densities. Marine Ecology Progress Series DOI, PDF

#Nguyen, H., #T. Hoang, #D. Hawkins, B. Allen, and B. Pernet. 2021. Temporal variation in food limitation in larvae of the sand dollar Dendraster excentricus. Marine Ecology Progress Series 665: 127-143 DOI

Ziegler, A., #A.M. Gilligan, J.G. Dillon, and B. Pernet. 2020. Schizasterid heart urchins host microorganisms in a digestive symbiosis of Mesozoic origin. Frontiers in Microbiology 11:1697 DOI, PDF

Pernet, B. 2020. An oral brush of cilia in the feeding larvae of Micronephtys cornuta (Annelida, Nephtyidae). Invertebrate Biology 139, article e12287 (7 pp.) DOI

Pernet, B. 2020. Opposed bands of cilia in the nonfeeding larvae of the serpulid annelid Salmacina tribranchiata. Invertebrate Biology 139, article e12285 (8 pp.) DOI

Fork, S., B. Pernet, and K. Wasson. 2020. Establishment of an extensive breeding population of a marine pulmonate snail far poleward of its previously documented range. Marine Biodiversity Records 13:5 DOI, PDF

Appy, R.G., S. Goffredi, B. Pernet, and C. Latino. 2019. Experimental elucidation of the life cycle of Rhinebothrium urobatidium (Cestoda: Rhinebothriidea) from the round stingray (Urobatis halleri) to first and second intermediate hosts. Bulletin of the Southern California Academy of Sciences 118(3): 139-157 DOI

#Peria, J., and B. Pernet. 2019. Tolerance to salinity and thermal stress by larvae and adults of the serpulid annelid Ficopomatus enigmaticus. Invertebrate Biology 138(4), article e12271 (8 pp.) DOI

#Yee, A., J. Mackie, and B. Pernet. 2019. The distribution and genetic diversity of the non-indigenous annelid Ficopomatus enigmaticus in central and southern California. Aquatic Invasions 14(2):250-266 DOI [PDF available at that site]

#Passarelli, B., and B. Pernet. 2019. The marine live bait trade as a pathway for the introduction of non-indigenous species into California: patterns of importation and thermal tolerances of imported specimens. Management of Biological Invasions 10(1):80-95 DOI, PDF

Pernet, B., E. Silverman, and P. Valentich-Scott. 2019. The seashells of an iconic public artwork: diversity and provenance of the mollusks of the Watts Towers. Journal of Conservation and Museum Studies 17(1):1-11 DOI, PDF [Related file: Pictorial guide to the mollusks of Watts Towers PDF]

Pernet, B., B.L. Livingston, #C. Sojka, and #D. Lizárraga. 2017. Embryogenesis and larval development of the seastar Astropecten armatus. Invertebrate Biology 136(2):121-133 DOI

#Von Tungeln, A., and B. Pernet. 2017. Differential host use affects fecundity of the gastropod Crepidula onyx. Marine Ecology 38(2), article e12421 (6 pp.) DOI

#Lizárraga, D., #A. Danihel, and B. Pernet. 2017. Low concentrations of large inedible particles reduce feeding rates of echinoderm larvae. Marine Biology 164, article 102 (12 pp.) DOI

#Jones, C., T. Stankowich, and B. Pernet. 2016. Allocation of cytoplasm to macromeres in embryos of annelids and molluscs is positively correlated with egg size. Evolution & Development 18:156-170 DOI

Pernet, B., #M. Barton, K. Fitzhugh, L.H. Harris, #D. Lizárraga, #R. Ohl, and C.R. Whitcraft. 2016. Establishment of the reef-forming tubeworm Ficopomatus enigmaticus (Annelida: Serpulidae) in southern California. Bioinvasions Records 5:13-19 DOI, PDF

Pernet, B., L.H. Harris, and P. Schroeder. 2015. Development and larval feeding in the capitellid annelid Notomastus cf. tenuis. Biological Bulletin 228:25-38 DOI

#McKee, A., J. Voltzow, and B. Pernet. 2013. Substrate attributes determine gait in a terrestrial gastropod. Biological Bulletin 224:53-61 DOI

Pernet, B., A. Amiel, and E. Seaver. 2012. Effects of maternal investment on larvae and juveniles of the annelid Capitella teleta determined by experimental reduction of embryo energy content. Invertebrate Biology 131:82-95 DOI

#Hofstee, J., and B. Pernet. 2011. Evolutionary simplification of velar ciliation in the nonfeeding larvae of periwinkles (Littorina spp.). Biological Bulletin 221:239-242 DOI

Pernet, B., and R.R. Strathmann. 2011. Opposed ciliary bands in the feeding larvae of sabellariid annelids. Biological Bulletin 220:186-198 DOI

Burnaford, J., S.Y. Henderson, and B. Pernet. 2011. Assemblage shift following population collapse of a non-indigenous bivalve in an urban lagoon. Marine Biology 158:1915-1927 DOI

#Kosman, E., and B. Pernet. 2011. Intraspecific variation in larval size and its effects on juvenile lophophore size in four bryozoans. Marine Ecology Progress Series 429:67-73 DOI, PDF

Pernet, B., and D. McHugh. 2010. Evolutionary changes in the timing of gut morphogenesis in larvae of the marine annelid Streblospio benedicti. Evolution & Development 12:618-627 DOI

Milonas, L., B. Pernet, and B.L. Bingham. 2010. Light influences feeding and growth of echinoplutei. Marine Ecology Progress Series 404:69-78 DOI, PDF

#Sloan, L., #S. Anderson, and B. Pernet. 2010. Kilometer-scale spatial variation in prevalence of the rhizocephalan Lernaeodiscus porcellanae on the porcelain crab Petrolisthes cabrilloi. Journal of Crustacean Biology 30:159-166 DOI, PDF

Pernet, B., #A. Deconinck, and L. Haney. 2010. Molecular and morphological markers for distinguishing the sympatric intertidal ghost shrimps Neotrypaea californiensis and N. gigas in the Eastern Pacific. Journal of Crustacean Biology 30:323-331 DOI, PDF

#Kosman, E., and B. Pernet. 2009. Diel variation in the sizes of larvae of Bugula neritina in field populations. Biological Bulletin 216:85-93 DOI

Pernet, B., #A. Deconinck, #A. Llaban, and J. Archie. 2008. Evaluating risks associated with transport of the ghost shrimp Neotrypaea californiensis as live bait. Marine Biology 153:1127-1140 DOI

Allen, J. and B. Pernet. 2007. Intermediate modes of larval development: bridging the gap between planktotrophy and lecithotrophy. Evolution and Development 9:643-653 DOI

Pernet, B. 2007. Determinate growth and variable size at maturity in the marine gastropod Amphissa columbiana. American Malacological Bulletin 22:7-15 DOI

Pernet, B. and #L. McArthur. 2006. Feeding by larvae of two different developmental modes in the annelid Streblospio benedicti. Marine Biology 149:803-811 DOI

Pernet, B. 2004. The cryptic feeding house of an invertebrate larva. Science 306:1757 DOI

Pernet, B. and W.B. Jaeckle. 2004. Size and organic content of eggs of marine annelids, and the underestimation of egg energy content by dichromate oxidation. Biological Bulletin 207:67-71 DOI

Pernet, B. 2003. Persistent ancestral feeding structures in nonfeeding annelid larvae. Biological Bulletin 205:295-307 DOI

Pernet, B. 2001. Escape hatches for the clonal offspring of serpulid polychaetes. Biological Bulletin 200:107-117 DOI

Pernet, B. 2000. A scaleworm’s setal snorkel. Invertebrate Biology 119:147-151 DOI

Pernet, B. 2000. Reproduction and development of three symbiotic scaleworms (Polychaeta: Polynoidae). Invertebrate Biology 119:45-57 DOI

Kohn, A.J., M. Nishi, and B. Pernet. 1999. Snail spears and scimitars: a character analysis of Conus radular teeth. Journal of Molluscan Studies 65:461-481 DOI, PDF

Pernet, B. 1999. Gamete interactions and genetic differentiation among three sympatric polychaetes. Evolution 53:435-446 DOI, PDF

Miner, B., E. Sanford, R.R. Strathmann, B. Pernet, and R. Emlet. 1999. Functional and evolutionary implications of opposed bands, big mouths, and extensive oral ciliation in larval opheliids and echiurids. Biological Bulletin 197:14-25 DOI

Pernet, B. 1998. Benthic egg masses and larval development of Amblyosyllis speciosa (Polychaeta: Syllidae). Journal of the Marine Biological Association of the United Kingdom 78:1369-1372 DOI

Pernet, B. and A.J. Kohn. 1998. Size-related obligate and facultative parasitism in the marine gastropod Trichotropis cancellata. Biological Bulletin 195:349-356 DOI

Pernet, B. 1997. Development of the keyhole and growth rate in Diodora aspera (Gastropoda: Fissurellidae). The Veliger 40:77-83

Phillips, N.E. and B. Pernet. 1996. Capture of large particles by suspension feeding scaleworm larvae (Polychaeta: Polynoidae). Biological Bulletin 191:199-208 DOI

Hodin J., A. Heyland, A. Mercier, B. Pernet, D.L. Cohen, J.-F. Hamel, J.D. Allen, J.S. McAlister, M. Byrne, P. Cisternas, and S.B. George. 2019. Culturing echinoderm larvae through metamorphosis. Pp. 125-169 in K.R. Foltz and A. Hamdoun, eds., Methods in Cell Biology. Volume 150: Echinoderms Part A. Elsevier (peer-reviewed)

Pernet, B. 2019. Larval feeding: mechanisms, rates, and performance in nature. Pp. 87-102 in T.J. Carrier, A.M. Reitzel, and A. Heyland, eds., Evolutionary Ecology of Marine Invertebrate Larvae. Oxford University Press (peer-reviewed)

Reish, D.J. and B. Pernet. 2009. Life cycle cultures of annelids: a review. Pp. 47-62 in D.H. Shain, ed., Annelids as Model Systems in the Biological Sciences. Wiley-Blackwell (peer-reviewed)

Pernet, B., P.-Y. Qian, G.W. Rouse, C.M. Young, and K.J. Eckelbarger. 2002. Phylum Annelida: Polychaeta. Pp. 209-243 in Young, C.M., ed., Atlas of Marine Invertebrate Larvae. Academic Press, London (peer-reviewed)


I have taught courses in Invertebrate Biology, Marine Biology, and Estuarine Biology at the University of Washington, the University of Oregon, and their marine stations (the Friday Harbor Laboratories and the Oregon Institute of Marine Biology, respectively). At CSU Long Beach, I routinely teach two courses:

Biol 211, Introduction to Evolution and Diversity. The first of a three semester introductory sequence for biology majors. This course introduces evolution as a process responsible for extant biological diversity at the molecular, cellular, and organismal levels.

Biol 313, Invertebrate Zoology. This course covers some of the massive diversity of invertebrates, focusing mostly on marine animals because the oceans are where most animal diversity (except for insects) lives. The focus of the course is on how invertebrates are built and how they function, but we also cover such things as phylogeny, paleontology, development, and ecology. We spend most of our time looking at living invertebrates in the lab, but also take a few fieldtrips to see intertidal and subtidal representatives of many phyla.


  • 562.985.5378
  • CSU Long Beach, Department of Biological Sciences, 1250 Bellflower Blvd, Long Beach, CA 90840
  • Lab HSCI-114, Office HSCI-124 (where HSCI=Hall of Science)
  • By appointment
  • DM Me