Rachel is interested in molluscan shell form. Animals’ shells defend them from a variety of environmental dangers and predatory attacks, including a multitude of low-magnitude, repeated stresses, which could cumulatively cause lethal fatigue damage. She studies how shell features—from composition and microstructure to overall morphology—contribute to fatigue resistance, and then how mussels respond to accumulated shell damage. By developing new mechanical techniques to test shell fatigue resistance, she hopes to answer questions about the evolution of shell shape and the ecological interactions between hard-shelled molluscs, their predators, and the environment.

Rachel received her B.A. in Biology from Swarthmore College where she worked with Rachel Merz studying worm burrowing and distribution in the muddy intertidal zone. She then worked as the lab manager in Sheila Patek’s lab at Duke University examining the behavioral strategies that mantis shrimp use when hammering open hard-shelled prey

Lab Affiliations

All Publications

  • A single heat-stress bout induces rapid and prolonged heat acclimation in the California mussel, Mytilus californianus. Proceedings. Biological sciences Moyen, N. E., Crane, R. L., Somero, G. N., Denny, M. W. 2020; 287 (1940): 20202561


    Climate change is not only causing steady increases in average global temperatures but also increasing the frequency with which extreme heating events occur. These extreme events may be pivotal in determining the ability of organisms to persist in their current habitats. Thus, it is important to understand how quickly an organism's heat tolerance can be gained and lost relative to the frequency with which extreme heating events occur in the field. We show that the California mussel, Mytilus californianus-a sessile intertidal species that experiences extreme temperature fluctuations and cannot behaviourally thermoregulate-can quickly (in 24-48 h) acquire improved heat tolerance after exposure to a single sublethal heat-stress bout (2 h at 30 or 35°C) and then maintain this improved tolerance for up to three weeks without further exposure to elevated temperatures. This adaptive response improved survival rates by approximately 75% under extreme heat-stress bouts (2 h at 40°C). To interpret these laboratory findings in an ecological context, we evaluated 4 years of mussel body temperatures recorded in the field. The majority (approx. 64%) of consecutive heat-stress bouts were separated by 24-48 h, but several consecutive heat bouts were separated by as much as 22 days. Thus, the ability of M. californianus to maintain improved heat tolerance for up to three weeks after a single sublethal heat-stress bout significantly improves their probability of survival, as approximately 33% of consecutive heat events are separated by 3-22 days. As a sessile animal, mussels likely evolved the capability to rapidly gain and slowly lose heat tolerance to survive the intermittent, and often unpredictable, heat events in the intertidal zone. This adaptive strategy will likely prove beneficial under the extreme heat events predicted with climate change.

    View details for DOI 10.1098/rspb.2020.2561

    View details for PubMedID 33290677

  • Mechanical fatigue fractures bivalve shells. The Journal of experimental biology Crane, R. L., Denny, M. W. 2020; 223 (Pt 10)


    Mollusk shells protect against diverse environmental and predatory physical threats, from one-time impacts to chronic, low-magnitude stresses. The effectiveness of shells as armor is often quantified with a test of shell strength: increasing force is applied until catastrophic fracture. This test does not capture the potential role of fatigue, a process by which chronic or repeated, low-magnitude forces weaken and break a structure. We quantified the strength and fatigue resistance of California mussel (Mytilus californianus) shells. Shells were fatigue tested until catastrophic failure by either loading a valve repeatedly to a set force (cyclic) or loading a valve under constant force (static). Valves fatigued under both cyclic and static loading, i.e. subcritical forces broke valves when applied repeatedly or for long durations. Stronger and more fatigue-resistant valves tended to be more massive, relatively wider and the right-hand valve. Furthermore, after accounting for the valves' predicted strength, fatigue resistance curves for cyclic and static loading did not differ, suggesting that fatigue fracture of mussels is more dependent on force duration than number of cycles. Contextualizing fatigue resistance with the forces mussels typically experience clarifies the range of threats for which fatigue becomes relevant. Some predators could rely on fatigue, and episodic events like large wave impacts or failed predation attempts could weaken shells across long time scales. Quantifying shell fatigue resistance when considering the ecology of shelled organisms or the evolution of shell form offers a perspective that accounts for the accumulating damage of a lifetime of threats, large and small.

    View details for DOI 10.1242/jeb.220277

    View details for PubMedID 32461264

  • Smashing mantis shrimp strategically impact shells. The Journal of experimental biology Crane, R. L., Cox, S. M., Kisare, S. A., Patek, S. N. 2018; 221 (Pt 11)


    Many predators fracture strong mollusk shells, requiring specialized weaponry and behaviors. The current shell fracture paradigm is based on jaw- and claw-based predators that slowly apply forces (high impulse, low peak force). However, predators also strike shells with transient intense impacts (low impulse, high peak force). Toward the goal of incorporating impact fracture strategies into the prevailing paradigm, we measured how mantis shrimp (Neogonodactylus bredini) impact snail shells, tested whether they strike shells in different locations depending on prey shape (Nerita spp., Cenchritis muricatus, Cerithium spp.) and deployed a physical model (Ninjabot) to test the effectiveness of strike locations. We found that, contrary to their formidable reputation, mantis shrimp struck shells tens to hundreds of times while targeting distinct shell locations. They consistently struck the aperture of globular shells and changed from the aperture to the apex of high-spired shells. Ninjabot tests revealed that mantis shrimp avoid strike locations that cause little damage and that reaching the threshold for eating soft tissue is increasingly difficult as fracture progresses. Their ballistic strategy requires feed-forward control, relying on extensive pre-strike set-up, unlike jaw- and claw-based strategies that can use real-time neural feedback when crushing. However, alongside this pre-processing cost to impact fracture comes the ability to circumvent gape limits and thus process larger prey. In sum, mantis shrimp target specific shell regions, alter their strategy depending on shell shape, and present a model system for studying the physics and materials of impact fracture in the context of the rich evolutionary history of predator-prey interactions.

    View details for DOI 10.1242/jeb.176099

    View details for PubMedID 29903746

  • Mechanical properties of sediment determine burrowing success and influence distribution of two lugworm species JOURNAL OF EXPERIMENTAL BIOLOGY Crane, R. L., Merz, R. A. 2017; 220 (18): 3248–59


    We apply new perspectives on how organisms burrow by examining the association of in situ variation in sediment mechanical properties with burrowing ability and species distribution of two sympatric lugworms, Abarenicola pacifica and Abarenicola claparedi We quantified the sediment's resistance to penetration and its grain size distribution at sites inhabited by each species. Abarenicola pacifica individuals were found in significantly harder to penetrate, more heterogeneous sediments. We compared worm burrowing ability using reciprocal transplant experiments. Worms from firmer sediments, A. pacifica, were able to make successful steep burrows in sediments characteristic of either species. In contrast, A. claparedi individuals often failed to complete successful burrows in the firmer A. pacifica sediment. To examine how morphological differences could explain these patterns, we compared body wall musculature and measured how well individuals support their own bodies when draped over a cantilever. Lugworms from the firmer sediment had thicker body wall musculature and held their bodies more rigidly than did worms from softer sediments. Additionally, we observed subtle differences in the papillae on the proboscises' surfaces, which could affect worm-sediment interactions, but we found no differences in the chaetae of the two species. Abarenicola claparedi produced more mucus, which could be important in shoring up burrow walls in their shifting, sandy habitat. This study presents the first example of using field-based experiments to determine how sediment mechanical properties and worm burrowing ability could act to determine organismal distribution. Our findings have broader ecological implications because of the role of lugworms as ecosystem engineers.

    View details for DOI 10.1242/jeb.156760

    View details for Web of Science ID 000411199600012

    View details for PubMedID 28931717