Publications    *student co-author


34. Jensen K.R., Krug P.J., DuPont A., and M. Nishina. 2014. A review of taxonomy and phylogenetic relationships within Costasiella

(Mollusca, Heterobranchia, Sacoglossa), with description of a new species. Journal of Molluscan Studies, 80: 562-574.



33.  Krug, P.J., Vendetti, J.E., Retana*, J., Rodriguez*, A., Hirano, Y., and C.D. Trowbridge. 2013. Integrative species delimitation

in photosynthetic sea slugs reveals twenty candidate species in three nominal species studied for drug discovery, plastid symbiosis

or biological control. Molecular Phylogenetics and Evolution, 69: 1101-1119. PDF (doi: 10.1016/j.ympev.2013.07.009)


32.  Kocot, K.M.,  Halanych, K., and P.J. Krug. 2013. Phylogenomics supports Panpulmonata: Resolving key evolutionary steps in a

major radiation of gastropod molluscs. Molecular Phylogenetics and Evolution, 69: 764-771. PDF (doi: 10.1016/j.ympev.2013.07.001)

31.  Romero*, M.R., Phuong*, M., Bishop, C.D., and P.J. Krug. 2013. Differential function of nitric oxide signaling during habitat 
choice by two larval morphs of the sea slug Alderia willowi: mechanistic insight into evolutionary transitions in dispersal strategies. 
Journal of Experimental Biology, 216: 1114-1125. PDF (doi: 10.1242/jeb.080747)  
30.  Marshall, D. J., Krug, P.J., Kupriyanova, E., Byrne, M., and R. B. Emlet. 2012. The biogeography of marine invertebrate life 
histories. Annual Reviews of Ecology, Evolution and Systematics, 43: 97-114. (doi: 10.1146/annurev-ecolsys-102710-145004)
29.  Krug, P.J., Gordon*, D., and M. Romero*. 2012. Seasonal polyphenism in larval type: Rearing environment influences the 
development mode expressed by adults in the sea slug Alderia willowi. Integrative and Comparative Biology, 52: 161-172. 
PDF (doi: 10.1093/icb/ics059)
28.  Vendetti, J.E., Trowbridge, C.D., and P. J. Krug. 2012. Poecilogony and population genetic structure in Elysia pusilla 
(Heterobranchia: Sacoglossa), and reproductive data for five sacoglossans that express dimorphisms in larval development. 
Integrative and Comparative Biology, 52: 138-150. (doi: 10.1093/icb/ics077) PDF  Table_S1
27.  Krug, P.J., Asif*, J.H., Baeza*, I., Morley, M., Blom, W. and T. Gosliner. 2012. Molecular identification of two species of the 
carnivorous sea slug Philine, invaders of the U.S. west coast. Biological Invasions, 14: 2447-2459. PDF  Table_S1
(doi: 10.1007/s10530-012-0242-9) 
26.  Asif*, J.H. and P.J. Krug. 2012. Lineage distribution and barriers to gene flow among populations of a globally invasive marine 
mollusc. Biological Invasions, 14: 1431-1444. (doi: 10.1007/s10530-011-0169-6) PDF   supplemental_material
25.  Krug, P.J., Händeler, K., and J. Vendetti. 2011. Genes, morphology, development and photosynthetic ability support the 
resurrection of Elysia cornigera (Heterobranchia: Plakobranchoidea) as distinct from the solar-powered sea slug E. timida. 
Invertebrate Systematics, 25: 477–489. (doi: 10.1071/IS11026) PDF   Table_S1
24.  Krug, P. J. 2011. Patterns of speciation in marine gastropods: A review of the phylogenetic evidence for localized radiations 
in the sea. American Malacological Bulletin, 29: 169-186. (doi: 10.4003/006.029.0210) PDF

23.  Händeler, K., Grzymbowski, Y.P., Krug, P.J. and H. Wägele. 2009. Functional chloroplasts in metazoan cells – a unique

evolutionary strategy in animal life. Frontiers in Zoology, 6: 28 http://www.frontiersinzoology.com/content/6/1/28


22.  Allen*, R.M., Krug, P.J., and D.J. Marshall. 2009. Larval size is determined by extra-embryonic provisioning but not egg size

in a marine gastropod. Marine Ecology Progress Series, 389: 127-137. PDF


21.  Krug, P.J. 2009. Not my "type": Bet-hedging and dispersal dimorphisms in opisthobranch life histories. Biological Bulletin, 216:

355-372. PDF


20.  *Smolenksy, N., *Romero, M. R., and P.J. Krug. 2009. Evidence for costs of mating and self-fertilization in a simultaneous

hermaphrodite with hypodermic insemination, the opisthobranch Alderia willowi. Biological Bulletin, 216: 188-199. PDF


19.  Krug, P.J., Riffell, J.A., and R. K. Zimmer. 2009. Endogenous signaling pathways and chemical communication between sperm

and egg. Journal of Experimental Biology, 212: 1092-1100. PDF


18.  Krug, P.J., Morley, M.S., *Asif, J.H., Hellyar, L.L. and W. M. Blom. 2008. Molecular confirmation of species status for the rare

cephalaspidean Melanochlamys lorrainae (Rudman, 1968), and comparison with its sister species M. cylindrica Cheeseman, 1881.

Journal of Molluscan Studies, 74: 267-276. PDF


17.  Krug, P.J. 2007. Poecilogony and larval ecology in the gastropod genus Alderia. American Malacological Bulletin 23: 99-111. PDF


16.  Krug, P.J., *Ellingson, R.A., Burton, R.A. and Á. Valdés. 2007. A new poecilogonous species of sea slug (Opisthobranchia:

Sacoglossa) from California: Comparison with the planktotrophic congener Alderia modesta (Lovén, 1844). Journal of Molluscan

Studies 73: 29-38. PDF


15.  *Ellingson, R.A. and P.J. Krug. 2006. Evolution of poecilogony from planktotrophy: Cryptic speciation, phylogeography and larval

development in the gastropod genus Alderia. Evolution 60: 2293-2310. PDF


14.  Zimmer, R.K., Schar, D.W., Ferrer, R.P., Krug, P.J., Katz, L.B., and W.C. Michel. 2006. The scent of danger: Tetrodotoxin (TTX)

as an olfactory cue of predation risk. Ecological Monographs 76: 585-600. PDF


13.  *Botello, G. and P.J. Krug. 2006. Desperate larvae revisited: Age, energy and experience affect sensitivity to settlement cues in

larvae of the gastropod Alderia sp. Marine Ecology Progress Series 312: 149-159. PDF


12.  Krug, P.J. 2006. “Defense of benthic invertebrates against surface colonization by larvae: A chemical arms race.” Pp. 1-53 in:

Clare A. and Fusetani N. (eds) Marine Molecular Biotechnology, vol 2. Springer, Berlin. PDF


11.  Krug, P.J. and and R.K. Zimmer. 2004. Developmental dimorphism: Consequences for larval behavior and dispersal potential in a

marine gastropod. Biological Bulletin 207: 233-246. PDF


10.  Riffell, J.A., Krug, P.J., and R.K. Zimmer. 2004. The ecological and evolutionary consequences of sperm chemoattraction.

Proceedings of the National Academy of Sciences 101: 4501-4506. PDF


9.  Riffell, J.A., Krug, P.J., and R.K. Zimmer. 2002. Fertilization in the sea: The chemical identity of an abalone sperm attractant.

Journal of Experimental Biology 205: 1439-1450. PDF


8.  Krug, P.J. 2001. Bet-hedging dispersal strategy of a specialist marine herbivore: A settlement dimorphism among sibling larvae

of Alderia modesta. Marine Ecology Progress Series 213: 177-192. PDF


7.  Krug, P.J. and and R.K. Zimmer. 2000. Developmental dimorphism and expression of chemosensory-mediated behavior: Habitat

selection by a specialist marine herbivore. Journal of Experimental Biology 203: 1741-1754. PDF


6.  Krug, P.J. and and R.K. Zimmer. 2000. Larval settlement: chemical markers for tracing production, transport, and distribution of

a waterborne cue. Marine Ecology Progress Series 207: 283-296. PDF


5.  Krug, P.J., and A.E. Manzi, 1999. Waterborne and surface-associated carbohydrates as settlement cues for larvae of the specialist

marine herbivore Alderia modesta. Biological Bulletin 197: 94-103. PDF


4.  Krug, P.J., 1998. Poecilogony in an estuarine opisthobranch: Planktotrophy, lecithotrophy, and mixed clutches in a population of

the estuarine opisthobranch Alderia modesta. Marine Biology 132: 483-494. PDF


3.  Graziani, E.I., Andersen, R.J., Krug, P.J., and D.J. Faulkner. 1996. Stable isotope incorporation evidence for the

de novo biosynthesis of terpenoic acid glycerides by dorid nudibranchs. Tetrahedron 52: 6869-6878. PDF


2.  Krug, P.J., Boyd, K.G., and D.J. Faulkner. 1995. Isolation and synthesis of Tanyolides A and B, metabolites of the nudibranch

Sclerodoris tanya. Tetrahedron 51: 11063-11074.


1.  Krug, P.J., Gileski, A.Z., Code, R., Torjussen, A., and M.B. Schmid. 1994. Endpoint bias in large Tn-10 catalyzed inversions in

Salmonella typhimurium. Genetics 136: 747-756. PDF