What good is a clam?

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When I mention to people that I study bivalves, I can sometimes sense from their facial expressions that they are secretly asking “why?” While clams are perfectly content to keep doing what they’re doing without being thanked, I think it’s important to enumerate all of the ways they make our world more livable and functional.

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Various roles that freshwater mussels can play in their local food webs (Source: Vaughn and Hoellein, 2018)

Bivalves are ecosystem engineers. While they may seem rather stationary and not up to much at any particular time, they are actually always working to actively maintain their habitat. The majority of clams are filter-feeders, meaning that they use their gills to gather particles from the water column for food. Some of these particles are ingested as food and later pooped out. Some inedible particles are discarded immediately by the clam as “pseudofeces”. Both mechanisms serve as a bridge between the water column and the benthos (the sediment at the bottom). In this way, clams are engines that take carbon fixed by algae floating in the water and transfer that material to be stored in the sediment. Their bodies also act as nutrition to feed all sorts of animals higher on the food chain like sea stars, lobsters, seabirds, sea otters and humans that depend on bivalves as food. They are literally sucking up the primary productivity (algae) to be used by the rest of the food chain.

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The filtration rate of oysters. Graphic from The Nature Conservancy

Different clam species vary in their precise filtration rate (how fast they can inhale and exhale water, filtering the particles within), but it is prodigious. Some freshwater mussels, for example, can pick-through 1-2 liters of water per hour for every gram of their own flesh. Since these individual bivalves can weigh over 100 g, they are capable of picking the food out of an immense quantity of water. In doing so, bivalves help improve the clarity of the water column, allowing more sunlight to reach deeper into the water body (the photic zone), providing more energy for additional photosynthesis to occur. While there are examples where invasive bivalves such as zebra or quagga mussels take this phenomenon too far, in well-functioning ecosystems, the filtration activity of clams helps improve the productivity of the community.

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An oyster reef. Source: The Nature Conservancy

Bivalves help make sediment through their filtration of material from the water column, and they also engineer and manipulate the sediment directly. Some bivalves, like oysters, are able to make huge mounds of dirt that serve as habitat for all sorts of life, increasing the diversity of the community. They do so both by excreting sediment, and also by passively trapping it between the shells of neighboring oysters (“baffling”). By doing so, they reduce rates of coastal erosion and increase the biodiversity of wetlands. For this reason, New York and other communities plan to seed oyster reefs to help fight sea level rise and reduce the threat of storm surges like the one that occurred during Superstorm Sandy.

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Comparison of sediments without bioturbation by digging animals, and with. Notice how the non-bioturbated sediment is layered and darkened due to activity by anaerobic bacteria, while the well-oxygenated, mixed sediment is light all the way through. From Norkko and Shumway, 2011

Other “infaunal” bivalves (burrowers) help to aerate the sediment through their tunneling, bringing oxygen deep under the surface of the dirt. This mixing of the sediment (also called bioturbation) ensures that nutrition from deep under the sediment surface is again made available for other organisms. Some bivalves can bore into coral reefs or solid rock, creating burrows which serve as habitat for other animals and can free up minerals for use by the surrounding ecosystem. Helpful shipworms assist in eating wood, assisting in returning nutrients stored in that tissue to the ecosystem as well.

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Enormous grouping of giant clams in a lagoon in French Polynesia. From Gilbert et al., 2005

Bivalves of course are also famous for their shells, and this activity also provides habitat to sponges, snails, barnacles and many other encrusting organisms specially adapted to live on bivalve shells and found nowhere else. Giant clams are the most legendary “hypercalcifiers,” and in some regions like New Caledonia can rival reef-building corals in terms of biomass. In areas where soft-bottoms dominate, bivalves like hammer oysters, adapted to “rafting” on the quicksand-like surface of the soft sediment, can assist by providing a platform for other animals to take refuge. In the deep sea, bathymodiolid mussels and other chemosymbiotic bivalves can feed directly on the methane and sulfur emitted from hot vents or cold seeps with the help of symbiotic bacteria, creating dense reefs which provide food and habitat for all sorts of life. Even once the clams die, their shells can continue to serve as homes for other creatures.

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Crabs feeding on Bathymodiolus in the deep sea (NOAA)

The shells of clams provide great scientific value in understanding our world. Much like tree rings serve as a record of environment thousands of years into the past, growth rings in clam shells serve as a diary of the animal’s life. These rings can be yearly, lunar, tidal or even daily in rhythm, with each ring serving as a page in the diary. The chemistry of those “pages” can be analyzed to figure out the temperature the clam experienced, what it ate, whether it suffered from pollution, and even the frequency of storms! The study of rings in the hard parts of animals is called sclerochronology, and it’s what first drew me to study bivalves. I was so fascinated by the idea that our beaches are covered with high-resolution records of the ocean environment, waiting to be cut open and read.

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This giant clam shell recorded an interruption in the animal’s daily growth caused by a typhoon! From Komagoe et al., 2018

While they don’t owe us anything, clams provide a lot of value to humans as well, serving as a sustainable and productive source of food. Humans have been farming bivalves for thousands of years, as evidenced by “oyster gardens” and shell middens which can be found all over the world. Particularly in seasons when food is scarce on land, native peoples could survive by taking advantage of the wealth of the sea, and bivalves are one of the most plentiful and accessible marine food sources available. But they aren’t just the past of our food; they may be part of the future. Bivalves are one of the most sustainable sources of meat known, requiring very little additional food to farm and actively cleaning the environment in the process. Mussels grown out on a rope farm are an easy investment, growing quickly and with very little required energy expenditure. Someday, giant clams may provide the first carbon-neutral meat source, as they gain their food from symbiotic algae within their flesh. I have never eaten one, but I’ve heard they’re delicious.

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A shell midden in Argentina. Photo from Mikel Zubimendi, Wikipedia
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Mussels being farmed on ropes

Clams are heroes we didn’t know we needed and maybe don’t deserve. They ask for nothing from us, but provide vast services which we take for granted. So the next time you see an inconspicuous airhole in the sand, thank the clam that could be deep below for aerating the sediment. The shell of that long-dead mussel at your feet may have fed a sea star, and now is a home for barnacles and many other creatures. While that mussel was alive, it sucked in algae to improve water quality on our beaches. And the sand itself may contain countless fragments of even more ancient shells. Clams silently serve as an important cog in the vast machine that makes our oceans, rivers and lakes such amazing places to be. Thank you clams!

 

Mystery of the “spurting” mussels

If you’ve read any of my posts, you should realize by now that clams are pretty weird. Some catch live prey. Some have algae in their bodies that they “farm” for food. Some can bore into hard rock. Some sail the seas on rafts of kelp. Clams live in a competitive world and have had hundreds of millions of years of time to evolve to try out all sorts of weird, unlikely ways of life.

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U. crassus in a Slovenian river (Alexander Mrkvicka)

The thick shelled river mussel (Unio crassus) is known from many rivers and streams of Central Europe. As this is a very well-studied region of the world, many generations of academics have noted an unusual, seemingly inexplicable behavior undertaken by these mussels at certain times of year.

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U. crassus propped up on its foot (UCforLife)

Using its muscular foot, U. crassus pulls itself to the edges of the streams and rivers it lives in until it is partially exposed to air. It orients itself at a right angle with the surface of the stream with its siphons (two little snorkels coming out of the shell) facing out towards the water. Like all bivalves, U. crassus can act as a bellows by opening and closing its shell to pull in and push out water through those siphons. It has one siphon above the water and one below, and it proceeds to suck in water and spray it into the center of the stream using the power of its suction. The water can travel over a meter away and they continue this spurting about once a minute, sometimes for hours.

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Squirting water into the stream! (Vicentini 2005)

Needless to say, this is a very strange and unlikely behavior to observe in a mussel. It is exposing itself to potential dessication or suffocation from exposure to air. It is vulnerable to predation from terrestrial mammals and birds. There has to be a very powerful benefit from this behavior to outweigh those risks. And why squirt water into the air?

Some researchers proposed that the mussels were traveling to shore to harvest from the more plentiful food particles deposited there. But why would they face their siphons away from the shore then? Other workers suggested that it was a way to reduce heat stress through evaporation, though that also seems unlikely, considering the water is warmest in the shallows. The question persisted for decades in the minds of curious malacologists.

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Top-down view of spurting behavior (Vicentini 2005)

In 2005, Heinrich Vicentini of the Swiss Bureau for Inland Fisheries and Freshwater Ecology decided to try settle the question of why these mussels spurt. He observed several dozen of the mussels crawl to the edge of the water and diligently begin squirting into the streams. In the name of science, he put himself in the path of these squirts, caught the water and used a hand lens to observe that the squirted water was full of mussel larvae (glochidia).

Lifecyle of U. crassus (Rita Larje via UCforLife)

U. crassus falls in the order Unionida, a group of freshwater mussels distinguished by a very unusual method of reproduction. They are parasites! Because they can’t swim well enough to colonize upstream against the current, they need to rely on fish to hitch a ride. Some have evolved elaborate lures to convince fish to take a bite, then allowing them to release their larvae, which attach to the fish’s gills like binder clips and ride all the way upstream. Once they have reached their destination, they detach and grow up into more conventional burrowing mussels. It’s a weird, creepy and wonderfully brilliant strategy that enabled the mussels to invade the inland rivers which would otherwise be inaccessible to them.

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Loach (type of freshwater fish) gills with unionid larvae attached (UCforLife)

The mussels appear to be spurting out not only water, but their babies. They gain a couple of advantages from this. For one, their larvae can distribute further than would be possible from the bottom of the creek. Instead, they are released at the center of the surface of the stream, where they can be carried for a much longer distance by the current before they settle at the bottom. In addition, the splash of water on the surface may mimic the behavior of insects and other fish food falling in the water. A curious minnow might venture to investigate the source of the splash, where it would promptly breathe in a cloud of larvae that get stuck on its gills. A pretty rude surprise, but a brilliant trick to give the baby mussels the best chance of surviving.

So again, clams prove themselves to be far more clever and interesting than they might initially seem. U. crassus and other members of the Unionida are an ancient and globally distributed lineage which have evolved all sorts of weird and wonderful ways to maintain their river lifestyle. Unfortunately, rivers are some of the most widely damaged environments in the world. A majority of freshwater mussel species worldwide including U. crassus are endangered by habitat loss, overharvesting and pollution. But more research into their unusual biology can help us understand ways we can enhance their conservation, with the hope of providing more habitat for them to recover populations in the future. New projects in Sweden and other countries aim to recover habitat for their larvae to settle along 300 km of rivers, and research the fish species which their larvae prefer to hitch a ride on. With more work, we can hopefully ensure that the streams of Europe will harbor little mini super-soakers for millennia to come.

Weird Clam Profile: Pinna nobilis

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A fan mussel among the seagrass it calls home (Arnaud Abadie on Flickr)

The fan mussels (Pinna nobilis) are a species of enormous mussel which live in seagrass beds of the Mediterranean Sea. They can grow to nearly 4 feet long (though most are 1-2 feet in size at maturity), and live with most of their bodies protruding straight up out of the sediment, anchored down into the sand with long rootlike byssal threads which grow out of their rear hinge.

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They are really enormous (Marc Arenas Camps on WordPress)

These mussels grow up to 20 cm per year, almost entirely in the vertical direction. As they gain in mass, their bodies start to sink in the sand beneath them, so it is believed this extremely fast growth rate evolved in order to stay above the sediment. It also helps them to remain elevated above the seagrass around them, where they can access passing phytoplankton and organic particles in the current.

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A sea silk glove (Wikipedia)
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Close up view of the hairlike byssus. I definitely am feeling some beard envy here. (Wikipedia)

Because they are exposed to the current like a giant fan, they need a very strong anchor. So they create huge quantities of byssal threads which root them down in the sand. These byssal threads are known as as “sea silk” and communities around the Mediterranean have used the silk to sew clothing for thousands of years. The material is extremely fine but strong, and has historically been of immense value as a result. Sea silk or sea wool is mentioned in writings of the ancient Egyptians, Greeks and Romans.

Unfortunately, the fan mussels are considered critically endangered due to overharvesting, pollution, climate change and destruction of their native seagrass habitats. However, they are now protected and active conservation efforts are underway. When the cruise ship Costa Concordia ran aground off of Italy in 2012, a community of fan mussels were rescued from a seagrass bed next to the wreck and moved to another nearby site. I hope someday to study the fan mussels because I find them to be a truly charismatic bivalve with many interesting mysteries still waiting to be uncovered about their unique lifestyle.

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Huge pen shell I saw at the Hebrew University Museum in Jerusalem. My lens cap is only 6 cm to give you a sense of scale! The shells are fragile and easily break.

Lessons of the Condors

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Condor 606 was born in Big Sur and regularly flies back and forth between there and Pinnacles.

Visiting Pinnacles National Park the other day, we were lucky to spot California condors several times. Their wingspan can reach up to 3 m. Their graceful flight is a sight to behold as they ride the warm updrafts of between the pinnacles of rock in the park, with their primary feathers bending up like a conductor’s fingers. Condors are the only remaining members of the genus Gymnogyps, which once contained five species. Four are only known from fossil specimens and went extinct at the end of the Pleistocene (~12,000 years ago), but Gymnogyps once ranged across the Americas. As such, the California condor (Gymnogyps californianus) is a relic species; a survivor of a long but mostly extinct lineage.

By 1987, poaching, lead poisoning and habitat destruction had reduced the population to 27 individuals, of which 22 were captured and put into an emergency captive breeding program. In the thirty years since this project began, the population has increased to around 450 individuals. The program was expensive, painstaking and a massive undertaking. To this day, all captive-bred individuals are individually numbered and continually monitored. They even have a very adorable directory on the Pinnacles site where you can look each bird up by its number. We saw #606 and #463, and a couple others from farther away that we couldn’t read.

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Condor #463 was born at a breeding center in Idaho

 

California condors can be distinguished by the much more common turkey vultures by their underwing coloration, with large white patches at the front of the wings. I think that turkey vultures are fun to watch as they fly in circles over the highways searching for roadkill, but when you see a condor fly low over your head, it is awe-inspiring. Even the smaller juveniles have much larger, more pronounced heads than turkey vultures, and seem to fly even more effortlessly with a more gently curving V shape in their wings. We had stopped on the trail to discuss field markings for condors with another hiker when #463 soared over our heads, and we couldn’t help but jump for joy and hug each other at the privilege to see one of the famous condors ourselves.

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Turkey vulture just checking if we were feeling ok as we sat in the shade.

As an advocate for invertebrate conservation, I have been known to unfairly poke fun at the human tendency to focus on large, charismatic megafauna for conservation as opposed to smaller and less exciting species that may make up more of an ecosystem’s biomass, or represent a more important link in the local food chain. Pandas have used up billions of conservation dollars, yet they are kind of an evolutionary oddball with their poorly evolved guts that can barely digest their chosen bamboo food, and their infamous failure to successfully mate. Koalas have a similar story. We reformed tuna fishing not out of concern for the fish, but because of concern for dolphins getting mistakenly caught. We tend to put a lot of time and effort into conserving species that we consider cute, or cool, or awe-inspiring. The condors are important decomposers of carcasses, ranging over huge distances in their search for food and efficiently returning the nutrients of dead animals back to the ecosystem. But the public outcry motivating their rescue was greatly helped by the fact that these creatures are incredibly impressive megafauna. If the turkey vulture was critically endangered, it might not get the same funding supporting its conservation.

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This condor was very far away, gracefully circling the canyon.

But looking at the condors, my skepticism melted away and I was left with only gratitude that I was able to witness the grandeur of these beasts as they soared through the air; gratitude that I was able to see them myself and not only read about them in a book. I will never see a Steller’s Sea Cow, or a Great Auk, or a thylacine, or a dodo, because they disappeared before we realized what we were doing, and that extinction is a real and irreversible loss. Perhaps Gymnogyps won’t be around in 10,000 years. Their lineage originally evolved to feed on the giant carcasses from a collection of North American megafauna that is believed to have been hunted to death by humans (though that is a topic of endless debate). But if we hadn’t intervened to save the condors, we would have had to live with the guilt of knowing that we as a species committed the killing blow and did nothing to stop when we knew the reality of our crime. Instead we went to extreme lengths to save these unusual and majestic creatures because we feel empathy for them. As I watched condor 463 soar over my head, I felt relief, and pride, and hope. Success stories are important motivators for conservation, and I couldn’t help but think his wings were spelling out a “V” for victory as he flew away.

 

People (and frogs!) before cows

It was hard not to feel irritated as I opened the homepage of East Meadow Action, a grassroots group recently formed to defeat the development of Student Housing West (SHW), particularly regarding the Family Student Housing project on East Campus. The group has its heart in the right place. They believe that the field at the base of campus is a setting deserving of preservation, and are trying to prevent the building of new student housing to achieve this goal. I’m writing this to counter a number of points brought up on their site about the planning and design of SHW. I feel qualified to discuss this matter because I’ve been serving as a graduate student rep on the planning committee for the project since its early days last summer. From the start, I’ve been dedicating my efforts to keeping the project economical to ensure lower rent and as dense as possible, to prevent campus from sprawling into land needed by wildlife. I’m sorry to say that East Meadow Action’s efforts to defeat construction will harm both students and the environment if they succeed.

Student Housing West is an enormous planned development on the West side of campus designed to house ~2600 undergraduates, 200-220 graduate students, nearly 140 apartments for student families and a childcare facility. Well, it was initially planned to be on the west side of campus, beginning below Kresge College and continuing down the hill to the current site of Family Student Housing. That was the site university administrators provided for developers pitching their concepts for the project in an extended series of meetings last summer. We selected one developer, Capstone Development Partners, because of their proven track record of student housing construction and management. I personally voted for them because their plan had the highest goals for sustainability and the team made it clear that they would work to adapt to any contingencies which would surely arise during the future planning process.

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The red-legged frog. Apparently Californians ate them to near-extinction, no joke. From Wikipedia.

It turned out that their promised adaptability was put to the test when we were informed that nearly half the site would not be usable for construction. It turns out that a very cute and endangered species, the Red-Legged Frog, uses that corridor for their annual migration from the northerly forests to the grasslands near the Arboretum. Capstone suddenly had to work with half the space that they had initially anticipated for the 2800 students destined to live at West Campus. In addition, it meant there would be no housing prepared for current families living in Family Student Housing (FSH) when it was torn down to make room for the new development.

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Aerial view of the two sites. Source: UCSC CHES

This news could well have killed the project. SHW would not be built in time. Students would be left to fend for themselves and find housing in town on Craigslist. But members of the planning committee discussed another site for the new FSH, one which I have long personally considered a waste of space: the “meadow” near Hagar and Coolidge Drive. As an environmentalist and earth scientist, I bristle at the suggestion that the field at the base of campus is some kind of natural grassland. This space is heavily damaged; if it was left alone, it would return in a few decades to being redwood forest much like North Campus. But it hasn’t been left alone. It is instead a haven for domestic cows which graze and trample the space year-round. The animals, while endearing, continuously roam mowing every blade of grass to a stub, happily emitting methane on a prime piece of real estate.

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The cows declined to comment on their thoughts re: Family Student Housing.

Perhaps the current ongoing environmental review of the site will find that the land is needed by some sensitive species other than domestic cattle. If that turns out to be the case, then I will shut up and the project will probably need to restart at the drawing board. But some of the “alternative sites” that East Meadow Action assures us are available on campus (but never get specific about) seem far worse to me. Some possibilities include the forested land north of campus or the trailer park on the Northwest of campus. Both of these places seem much worse candidates to pave over to me in terms of ecological value (redwood and oak woodland) or their need for people (the trailer park is some of the most affordable housing on campus and much beloved by the students living there). I would much rather pave over cow pasture than a forest or someone’s home.

When I read through the website of East Meadow Action, I am struck that the centerpiece of their argument is the aesthetic value of lower campus as open space. I confess that I find this extremely irritating. It’s irritating to me because aside from being an advocate for conservation of sensitive species like the red-legged frog, I am also a student who has to get by living in this town, and I don’t have the luxury of worrying about aesthetics. When they mentioned Ranch View Terrace as an example of “building done responsibly”, my opinion was sealed. Take one look at the floor plans described for the single family homes of Ranch View Terrace, “a large housing complex set back from the road and mitigated by vegetation and topography.” How exactly are we going to house 2500 students in single family homes, artfully hidden behind trees? East Meadow Action is not motivated by environmentalism. They’re motivated by the same Not In My BackYard mentality that is choking the development of dense housing in town. We are in a housing crisis and their ocean views are a luxury that we can’t afford.

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East Meadow Action’s ideal for student housing.

I agreed to be SHW rep because I want to make sure future graduate students will have an affordable housing option on campus. I currently am “lucky” to spend 40% of my stipend on rent in town. When I first came to attend grad school here I had to deal with Craigslist and ended up paying over 60% of my income for the first two years I lived here. Students are pursuing extreme solutions. I know people considering living in the woods, or in unzoned bedrooms in town that are another source of tension with the community. It will only continue to get worse as rents rise in the coming years.

Right now, all we can do to remedy the issue in Santa Cruz is pass rent control ordinances and increase supply. SHW is the best project we have to increase supply and do so ensuring maximum density. I want UCSC to continue to be a forest campus, which necessitates not chopping down trees. I am trying to ensure that students have an affordable housing option on campus. I want student families to have guaranteed housing that comes online right when their previous outdated complex is torn down. The sentimental affection that some may feel for the aesthetics of a cow pasture strikes me as the privilege of a comfortable and vocal minority. I value the croaking of frogs and the laughter of children over the yammering of NIMBYs and, yes, even the mooing of cows.

Dan Killam

Fourth Year PhD Candidate, UCSC Earth and Planetary Sciences

University of Southern California ’12, BS Environmental Studies

Graduate Representative for Student Housing West

What is Conservation Paleobiology?

In undergrad, I felt like my school and internship were training me to be two different types of researcher. At USC, I was majoring in Environmental Studies with an emphasis in Biology. It was essentially two majors in one, with a year of biology, a year of chemistry, a year of organic chem, a year of physics, molecular biology, biochemistry, etc. On top of that, I took courses on international environmental policy and went to Belize to study Mayan environmental history. Meanwhile, I was working at Jet Propulsion Laboratory in Pasadena researching trends in historical rainfall data. I loved both sides of my studies, but felt like neither was exactly hitting the spot of what I would want to spend my career researching. I love marine biology but am not particularly interested in working constantly in the lab, looking for expression of heat shock protein related genes or pouring stuff from one tube into another. On the other hand, I was fascinated by the process of untangling the complex history of rainfall in California, but I yearned to relate this environmental history to the reaction of ecological communities, which was outside the scope of the project.

During my gap year post-USC, I thought long and hard about how I could reconcile these disparate interests. I read a lot, and researched a bunch of competing specialized sub-fields. I realized that paleobiology fit the bill for my interests extremely well. Paleobiologists are considered earth scientists because they take a macro view of the earth as a system through both time and space. They have to understand environmental history to be able to explain the occurrences of organisms over geologic time. I really liked the idea of being able to place modern-day changes in their geologic context. What changes are humans making that are truly unprecedented in the history of life on earth?

But it doesn’t have to be all zoomed out to million-year processes. A growing sub-field known as Conservation Paleobiology (CPB) is focused on quantifying and providing context of how communities operated before humans were around and before the agricultural and industrial revolutions, in order to understand the feasibility of restoration for these communities in this Anthropocene world. Sometimes, this means creating a baseline of environmental health: how did oysters grow and build their reefs before they were harvested and human pollution altered the chemistry of their habitats? I’m personally researching whether giant clams grow faster in the past , or are they reacting in unexpected ways to human pollution? It appears that at least in the Gulf of Aqaba, they may be growing faster in the present day. Such difficult and counterintuitive answers are common in this field.

Sometimes, CPB requires thinking beyond the idea of baselines entirely. We are realizing that ecosystems sometimes have no “delicate balance” as described by some in the environmental community. While ecosystems can be fragile and vulnerable to human influence, their “natural” state is one of change. The question is whether human influence paves over that prior ecological variability and leads to a state change in the normal succession of ecosystems, particularly if those natural ecosystems provide services that are important to human well-being. In a way, the application of paleobiology to conservation requires a system of values. It always sounds great to call for restoring an ecosystem to its prior state before humans. But if that restoration would require even more human intervention than the environmental harms which caused the original damage, is it worth it? These are the kinds of tricky questions I think are necessary to ask, and which conservation paleobiology is uniquely suited to answer.

At the Annual Geological Society of America meeting in Seattle this year, the Paleo Society held the first-ever Conservation Paleobiology session. The room was standing room only the whole time, investigating fossil and modern ecosystems from many possible angles. This field is brand new, and the principles behind it are still being set down, which is very exciting. It’s great to be involved with a field that is fresh, interdisciplinary, and growing rapidly. I look forward to sharing what my research and others find in the future.