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Research Interests

Ink is a dramatic predator defense used by a variety of unrelated taxa such as sea hares, cephalopods, and even pygmy sperm whales. Ink is multimodal, attacking predators visually (acting as a smoke screen) and chemically. These groups of inking organisms do not share a common inking  ancestor and their ink productions sites are not homologous. This gives us the unique opportunity to study a very specific chemical defense arising three different times throughout evolutionary history.




I am interested in the comparative analysis of the chemical and physical properties of ink from these three different animal groups. I also explore the antipredatory effects of ink through behavioral and neurophysiological experimentation.

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California sea hares (Aplysia californica) inking in a holding tank

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A bonnethead shark's (Sphyrna tiburo) behavioral reaction to common cuttlefish (Sepia officinalis) ink

Ink as a Chemical Defense

I am interested in how differently shaped cartilaginous fish noses sample the aquatic environment. Through a combination of histology, scanning electron microscopy, CT scanning, and computational fluid dynamics I investigated how nose morphology effects sensory structure distribution and water flow


Underside of a spotted ratfish (Hydrolagus colliei) showing the  Ammpullae of Lorenzini, nostrils, and mouth

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Schematic of a bonnethead shark olfactory lamellae. EC- excurrent canal, EN- excurrent naris, IC- incurrent canal, IN- incurrent naris, OB- olfactory bulb, OL- olfactory lamellae, ONL- olfactory nerve layer, R- raphe.


CT scan of a Pacific spiny dogfish (Squalus suckleyi)

Chondrichthyan Olfactory Morphology

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