And now for something completely non clam-related…a speculative essay about LOTR as an environmental allegory, drawing on my posts to social media last week that clearly struck a nerve.
I’m the first to admit that Sauron’s rule of Mordor was highly problematic. But frankly, I think those concerns, focusing largely on his foreign policy, have been discussed ad nauseam, and not enough attention has been paid to his legacy as a steward of the environment. Plentiful examples exist of Sauron’s all-seeing eye for conservation. One need look no further than the Dead Marshes. These scenic lands were a well-maintained string bog which represented one of the environmental gems of Mordor, and were maintained as a protected reserve where orcish patrols did not enter. The marshes were an innovative nature-based solution to the environmental challenge of the thousands of corpses left over after the Battle of Dagorlad. The Dark Lord in his wisdom allowed the wetlands to expand to mediate the damage left over and return those nutrients to the ecosystem.
The marshes hosted diverse populations of anaerobic bacteria which helped preserve the famed bog bodies of the area, leading to them being an important archaeological site. The microbes produced a small amount of natural methane, and while this reduced the ability of the marsh to sequester carbon from Mordorian industry, the resulting little candles were a major attraction for visiting ecotourists.
A nazgul conducting an aerial survey of the marshes
Some may argue that the marshes were not part of Mordor proper, which is true in the strict sense, but the marshes were clearly part of Sauron’s zone of influence, especially considering the extensive Ring Wraith aerial monitoring surveys he sent to survey the environmental health of the region and keep out interlopers of all sizes. Other critics pointed to a lack of crunchable birdses, but visitors approaching in early March are unlikely to observe many waterfowl, considering most would have migrated north by then to sites like Long Lake near the Lonely Mountain.
Sauron also contributed to forest reserves such as Mirkwood Old Growth Forest Biodiversity Reserve. While neurotoxins causing amnesia were present in some waterways, these were likely produced by harmful algal blooms resulting from Elvish untreated sewage. Mirkwood was home to a rich ecosystem supporting a vibrant population of endemic colonial spiders, Ungoliantus shelobii, which remain understudied. A small remnant spider population was also present in Cirith Ungol, supported by a locally administered feeding program. Unfortunately, that population was subject to poaching by an invading hobbit force during the War of the Ring.
Sauron’s detractors will point to environmental failures such as his alliance with Isengard, which reportedly resulted in some amount of deforestation. Saruman was previously an ally to Entish concerns, but necessities of the war effort resulted in previous protections being lifted, as is often the case in times of combat. While Sauron did attempt to encourage Saruman’s use of hydropower instead of biomass burning, Uruk-hai production was admittedly not as sustainable as it could have been. A full lifecycle carbon analysis was never conducted of their production processes, which were shrouded in secrecy, but they did represent a type of recycling.
Meanwhile, attempts to blame Sauron for the destruction of the Brown Lands have been undermined by a lack of primary sources, aside from Gondorian propaganda. The Entwives, the only known witnesses, did not respond to requests for comment for this report. Overall, it is clear that Sauron’s mixed legacy in foreign policy is improved somewhat when looking at his environmental record. One does not simply dismiss his environmental management ability.
You might be familiar with the concept of the present “biodiversity crisis“. There is an increasing consensus in the ecological research community that the current loss of species this planet is experiencing is not sustainable, in the sense that the loss of some species may precipitate the loss of more, in an accelerating spiral. The paleontological community has found that the pattern of species loss is unusual even at the scale of geological time, potentially placing us among the great extinctions in geologic history, or at least a notably bad extinction event. A less diverse biosphere means the loss of ecosystem services associated with all of the species we lose, and potentially a less resilient biosphere, stacking the deck against us as the climate continues to change and life is forced to adapt. Because our global civilization depends on the wealth of the biosphere for our own well-being, this is definitely very bad news for humanity (I usually tend to avoid rationalizing conservation based on ecosystems’ value to us, believing that we have a moral imperative to preserve the biosphere, and organisms have an inherent right to exist outside of their economic value, but that’s a topic for another blog).
We are only aware of the loss of species due to centuries of careful collecting, cataloging, categorization and curation undertaken by conservationists around the world, including indigenous communities, museum professionals, taxonomists, seed banks, herbaria, and other very highly specialized and educated people. I won’t refer to these biodiversity experts as “countless”, because they’re actually a pretty small group of folks entrusted with an almost incomprehensible responsibility: to quantify the biological wealth of our world. They figure out when baselines are shifting, and their work keeps us accountable as we seek to stop the current bleeding of biodiversity.
I am writing this post because biological collections are having a moment of attention, and it’s been a topic I have been thinking of for some time as an outsider. Duke University recently announced that they will be throwing out their herbarium, an archive of plant samples which is one of the leading such collections in the US. The herbarium supports a vibrant ecosystem of research on the classification of plants, and is an important archive of plant diversity. Duke University, which has an endowment of $11 billion, claiming to not have the resources to support this archive is an unacceptable dereliction of their duty to preserve and nurture knowledge. And sadly, this closure of such an important collection is not a one-off event. Worldwide, taxonomist and curator jobs are declining. These are the people who spend decades learning how to tell one species of snail from another based on their genitalia. They discover cryptic species in collections. They prevent collections from degrading due to improper preservation, and charge in to save samples from fires. They process loans and when someone like me is belated in returning samples, they write a polite email reminding me to send samples back. When these people leave science, their skills can’t be easily replaced. If collections are lost, they literally can’t be replaced.
I am a biogeochemist, and not a museum worker by any means, but so much of my work has relied on biological archives. But by my count, 3/4 of my ongoing projects have used biological collections in some way. I wanted to list out some of the ways that biological collections have enabled my research, because I don’t think I’m unusual. Biodiversity curators are the keystone species in a vast ecosystem of interconnected research that wouldn’t happen without the hard work of maintaining collections. Please do what you can to protect biodiversity collections, whether by pressuring your representatives, your alma maters, and through donations.
Projects that relied on curators:
Some of my most influential educational experiences relied on teaching collections, including at Cabrillo Marine Aquarium, USC, LA Museum of Natural History, UC Santa Cruz and elsewhere. Without collections to get my hands on in lab exercises and other educational opportunities, my skills in ecology, organism ID and more would be greatly diminished.
My second PhD chapter relied on samples of well-preserved Jurassic lithiotid bivalves which came from outcrops which mostly no longer exist, having been quarried out or already sampled. These were loaned by curators at the University of Padova and University of Verona Natural History Museums
My third and fourth PhD chapters, and one of my postdoctoral papers used shells of giant clams that were stored at the Hebrew University of Jerusalem Natural History Museum. These shells had been confiscated from poachers at the Egypt-Israel border and were loaned to me by curator Henk Mienis. I would rather these clams be still alive in the Red Sea, but at least we were able to use these for a series of papers about their ecology and physiology.
Before my PhD fieldwork in Israel, I visited the California Academy of Science collection to view shells of Red Sea giant clams and practice species ID using a taxonomic key. I would also note that one of the species I have studied, Tridacna squamosina, is named because of the work of museum curators at University of Vienna.
Another postdoctoral paper (in progress) on violet bittersweet clams from the Eastern Mediterranean made use of preserved samples collected during research cruises off the coast of Israel in the 1960s-1980s. These specimens, loaned by the Steinhardt Museum in Tel Aviv, represent some of the last observed live violet bittersweets seen off of Israel. They since have (likely) gone extinct in the region, probably due to sediment changes in the late 1980s following the construction of the Aswan Dam.
Yet another postdoctoral paper in progress resulted from study of the growth and chemistry of shells of wavy turban shells loaned from the Santa Barbara Museum of Natural History, which we were comparing to individuals that grew in the hot waters of the Biosphere 2 tropical reef ocean tank. I met the curator Vanessa Delnavaz at the Southern California Union of Malacologists 2021 meeting, which was hosted by the museum. Museums are important centers of scientific organization and networking in addition to the value of their collections!
So I hope all of these anecdotes help make clear that biological collections are absolutely vital to enable an entire universe of research, and that we often can’t possibly predict what collections are going to be useful for which scientific purposes until long after the samples were preserved. So any budgetary bean-counters (not talking about bean taxonomists) should think twice before closing any collections! This is a core responsibility of academic institutions and we cannot allow any of these collections to be lost. Fund your local biological curators! Without their hard work, we’d be flying blind in the current biodiversity crisis. They’re the heroes we need, and that our earth deserves.
Two giant clams near Eilat in the Northern Red Sea. To the left is the small giant clam, Tridacna maxima, and to the right is a mature individual of the rare endemic giant clam Tridacna squamosina, only found in the Northern Red Sea.
You are what you eat, and clams are too. We’re made of atoms, which come in “flavors” called isotopes, relating back to the mass of the atoms themselves (how many protons and neutrons they have). Nitrogen, for example, comes in two stable (non-radioactive) forms called nitrogen-14 and nitrogen-15. Much like scientists can track the composition of a person’s diet from the isotopes of their hair, researchers have used the isotopes of clams to figure out their diet.
Nitrogen isotopes provide us with a useful and detailed record of food webs. Plants and algae tend to have more of the light isotope of nitrogen in their tissues than the animals that eat them (primary consumers), and the animals that eat those animals have even higher nitrogen isotope values. We can measure the amount of “heavy” atoms of nitrogen with a unit called δ¹⁵N (“delta 15 N”). A carnivore at the top of the food chain will have a very high δ¹⁵N, while plants will be the lowest. Clams, typically being filter feeders, will usually have an intermediate value, since they’re eating a lot of phytoplankton (tiny microscopic floating algae) and zooplankton (animal plankton that eat other plankton).
But I study a special kind of clam, the giant clams, which have a cheat code enabling them to become giant: they have algae *inside* of their bodies. The algae make food using photosynthesis and share it with their hosts! In exchange, the clams provide the algae with a stable environment free of predators, plenty of fertilizer in the form of their own waste, and even channel extra light to help the symbionts grow faster. This partnership is called photosymbiosis, and is pretty rare in clams, though it is common in other animals like the corals that build the reefs where giant clams are found! Previous researchers have shown that giant clams have very low nitrogen isotopic values in their tissue, like a plant, because they get most of their nutrition from the algae, rather than filter feeding.
I am a sclerochronologist. That means I study the hard parts of animals, in this case the shells of bivalves. Like the rings of tree, bivalves make growth lines in their shells which can serve as a diary of their lives. Some of my past work has looked at using chemistry of the growth lines of giant clams to measure the temperatures they grow at, compare the growth of ancient and modern clams, and even look at how much the clams grow in a day! Today though, I’m talking about my most recent paper, which looks at how we can use the shells of giant clams as a food diary.
But when they’re babies, the symbiosis in giant clams is not yet fully developed. During this early period of their lives, giant clams actually get more of their nutrition from filter-feeding like a “normal” non-photosymbiotic clam, until they’ve had a chance to grow in surface area and become a living solar panel. Like all bivalves, the shells of giant clams are made of calcium carbonate, bound together by a protein scaffold we call the shell organic matrix. Proteins are made of amino acids, which contain nitrogen! If we can get the nitrogen out of the shell from the early part of the clam’s life, and compare it to the nitrogen at the end of the clam’s life, it might record the clam’s transition from filter feeding to its mature plant-like lifestyle! If our hypothesis holds, we should record a decrease through its life in the shell δ¹⁵N values.
A model I made of the clams’ nitrogen intake, with the left plot how they switch from filter feeding to getting most of their nitrogen from dissolved sources around 5-6 years of age. Because the nitrogen isotopes of those two sources are different, that manifests in the expected values from the clam’s body (the right plot)!
A map made by my talented partner, Dana Shultz!
So I gathered a team of talented collaborators and set out to test that hypothesis, using giant clam shells that I was able to get on loan from the Hebrew University of Jerusalem Museum. These shells had been confiscated from poachers at the Egypt-Israel border. While I would have rather known these clams were still alive in the waters of the Northern Red Sea, being able to use them for research to understand the biology of their species was the next best thing! I had originally planned on pursuing a postdoc undertaking this project with Rowan Martindale, a professor at UT Austin who has studied the nitrogen isotopes of photosymbiotic corals, but when I started up at Biosphere 2, we ended up continuing with the project anyway as a collaboration! We measured the nitrogen isotopes of the shell material in the lab of Christopher Junium, a professor at Syracuse University, who has developed an exquisitely sensitive method to measure the nitrogen from shell material by essentially burning the shell powder and then scrubbing out unwanted material to isolate the nitrogen, to measure the isotopes in a machine called a mass spectrometer. Katelyn Gray is a specialist in isotopes of biominerals and assisted with drilling out powder from the shells with a Dremel. Shibajyoti Das, now at NOAA, is a specialist in measuring the shell nitrogen isotopes of other bivalves and he was master at doing much of the mass spectrometer work, and assisting in interpretation. Adina Paytan is a professor at UC Santa Cruz. She first provided the funding and support for me to go to the Gulf of Aqaba and collect these shells as part of an NSF-funded student research expedition! She also provided environmental data which helped us to interpret what the clams were actually eating!
A figure showing the four shells we sampled from, with the sampling areas in each hinge area showing colored and matching with the corresponding isotope plot to the right (colored points). 3 of the 4 shells show declines in isotope values with age. Shaded ribbon behind the data shows the model output.
So what did our crack team of scientists find out? We found that three of the four tested giant clams did indeed measure a decline in nitrogen isotopes over the course of their lives. Their earliest growth lines in the hinge areas of their shells record elevated δ¹⁵N values similar to other filter-feeders from the region. But as they aged, their later growth lines show much lower δ¹⁵N values, more like photosymbiotic corals and plants from the region. So clams indeed recorded the transition in nutrition as they became solar-powered! This degree and directionality of change in nitrogen isotopes was much greater than has been observed in any other clams measured in this way, which made sense considering their unique physiology. The clams have another area of the shell, the outer shell layer, which is closer to the symbionts than the hinge area. In this outer shell area, we did not observe much of a consistent trend in nitrogen isotopes. It’s likely that the outer layer is highly influenced by the photosymbionts even at the earliest stages of life.
Growth lines in the hinge area of two of the shells lit from behind, with the drilled areas for this study visible as well. The outer shell layer is the opaque and was also sampled for this study.
There was one clam that differed from the others in showing low δ¹⁵N values through life in its hinge shell layer. To help explain these differences, I created an independent model of the clams’ internal chemistry based on their growth rate, which slows as they age, and also is faster in the summer. When the clams are young filter feeders, they get most of their nitrogen from plankton, debris and other material floating in the water column making up floating material we call Particulate Organic Matter (POM). Meanwhile, when they are in their photosynthetic life stage, they get most of their nitrogen from nitrate, which is essentially Miracle Gro for the symbionts. The model showed that the clams should record a flip from filter feeding to photosynthesis around 4-5 years of age, which was confirmed by three of the shells! But what about the one that didn’t show this trend? My colleague Adina had fortunately measured the isotopes of POM and nitrate in different seasons in the Gulf of Aqaba. We found that in summer, as expected, POM δ¹⁵N is lower than nitrate. In the winter, meanwhile, that relationship is flipped! So if a clam grew more in winter, it would not record the same transition as was seen by the other clams. We think the clam that was the exception to the rule might have been more of a winter grower.
The chaotic nutrient environment of the Northern Red Sea, showing how in different seasons, dissolved nitrate has higher or lower δ¹⁵N values than the Particulate Organic Matter that the clams filter-feed on.
But long story short, we were able to demonstrate for the first time that giant clams show nitrogen isotopic values in their shells in line with expectations from their diet. Other clams have been measured this way, but the fact that we were able to conduct these analyses at all is a testament to the sensitivity of the elemental analyzer in Chris’s lab. Giant clams have *very* low concentrations of organic matter in their shells, so the forward march of technology was a major factor enabling this study to be possible.
Why does it matter that we can measure the transition of the clams from filter-feeding to photosymbiotic in their shell records? Well, giant clams are not the only bivalves which have photosymbionts. There are other clams in the fossil record which have been proposed to have had symbioses with algae, but until now we’ve never had a definitive geochemical way to measure this in fossils. We hope that this approach can be applied to the organic material in fossil shells, which is often well preserved, to see if huge clams in the Cretaceous and Jurassic had a similar way of life to the modern giant clams! If we can demonstrate that was the case, we can see how such species responded to past intervals of climate change, which will help us understand how giant clams will fare in the warming, acidifying ocean of the present.
These results also help explain the lives of giant clams themselves. We hope this kind of data can be used to measure the symbiotic development of giant clams in different places, with different types of food and nitrogen available, where we’d have the potential to measure pollution. Interestingly, the time that the model shows the clams transitioning to photosynthetic maturity is right around the time that they reach reproductive maturity (5-10 years of age). We’d like to investigate whether the time of clam maturity is controlled by the development of their symbiosis, which itself might relate to nutrients in the clams’ environment. If clams can grow faster, then they can mature faster, and potentially reproduce sooner in life. Will giant clams be able to thrive in the presence of increased nitrate, which is a common pollutant in coral reef environments? Like all worthwhile research projects, we have dozens of new questions to pursue as a result of this work, so stay tuned for the next installment in this journey of clam knowledge!
If you use Mastodon (or another Fediverse service), you can now follow this blog from there. As you might know, Mastodon is one of my favorite places to communicate science, so this is a cool feature for me! You can follow me by searching “@dantheclamman.blog@dantheclamman.blog” from your Mastodon page. Or maybe you’re already viewing this on Mastodon, in which case, Hello World!
Figure 1 from our paper, showing a comparison of a scallop, its growth increments and where it came from in France, to a giant clam shell section (dyed blue to show its growth lines), and where it came from in the Northern Red Sea
In 2020, I got an interesting email in my inbox from another mollusk researcher! Niels de Winter had emailed me, who I was familiar with from his past work on big Cretaceous rudist bivalves and giant snails. Niels had seen my paper published that year on giant clam shell isotopes from the Gulf of Aqaba in the Northern Red Sea, and was interested in teaming up on a new study to compare the daily growth of giant clams with another bivalve that has daily growth: scallops! I was intrigued because I had similar work underway to study the shells of clams I was growing at Biosphere 2, but I didn’t have any plans to measure my collected wild clam shells that way. So this sounded like a win-win opportunity to work together on a study that neither of us could do alone! Plus, I liked his work and had cited it in the past.
The shells of bivalves are very useful as each produces a shell diary consisting of growth lines, similar to the rings of a tree. Giant clams keep a very detailed diary, with a new growth line forming every night, which previous research has suggested was due to the control that the symbiotic algae inside giant clams have on their host. When the algae conduct photosynthesis, they use CO₂ in the fluid the clam makes its shell from, which increases the pH and accelerates the formation of the shell mineral crystals! The symbionts also directly assist by pumping calcium and other raw materials for the clam to use! Niels had found such daily lines in an ancient rudist bivalve from over 66 million years ago, and proposed it as a sign that the rudists might have had similar algae! I used the daily lines to compare giant clam growth before and after humans arrived in the Red Sea, finding that the clams are growing faster!
But it turns out that giant clams aren’t the only bivalves that make daily lines. Some species of scallops do it too, but that’s a bit confusing, since scallops have no symbionts that could be producing this daily growth period! One way we could investigate this is by bombarding the shells with very tiny laser beams only 20 µm across: the width of a hair is a flawed unit of measurement but 20 microns is as narrow as the narrowest type of hair you can think of! The laser would carry across the cross sections of the shell in a line, literally burning away tiny bits of shell, with the resulting gases captured by a machine called a mass spectrometer, which can figure out the concentrations of elements in the gas.
So we’d basically create a very detailed wiggly graph, where the wiggles represent years, months, days and even tides, depending on how fast the clams and scallops grew! I’m happy to report the paper was published earlier this year, so I thought I’d switch it up a bit and have a conversation with Niels through this blog post. Let me open it up to Niels, who I decided to bring in for this post in a kind of conversation!
Niels, what did you expect to find heading into this experiment? For me, I figured the giant clams would have greater amplitude of variation on a daily basis than the scallops, due to the influence of the symbionts. Is this what you expected? More or less. To be honest, that is what I was hoping to find, because if the daily lines were so much stronger in photosymbiotic shells than in the non-photosymbiotic scallops, it would make it easier to recognize photosymbiosis by studying modern and fossil shells. Also, a finding like that would obviously support the hypothesis we had about the ancient rudist bivalve. However, I was a bit skeptical as to whether the reality would be so clear-cut.
I mailed samples from six juvenile giant clams to Niels for analysis. We went with juveniles for a couple reasons: they grow faster at this life stage than they do as adults: 2-5 centimeters per year for the species we were studying, which meant the greatest opportunity to record a very detailed record from their shells! Scallops also grow extremely quickly, up to 5 cm/year, and so we would be able to get a similar resolution for both types of bivalves, since each page in their diaries would be a similar width.
When I start a new study like this, I always like to “outsource” the expertise about the topic a bit. Our work in sclerochronology often involves bringing together several fields of research and interpreting the results of complex measurements like these requires input from several people who look at them from different viewpoints. I had just finished a research stay at the University of Mainz in 2019, where I worked with Bernd Schöne and Lukas Fröhlich. I know Lukas was working on scallops together with Julien Thébault, whose team collects them alive in the Bay of Brest and keeps a very detailed record of the circumstances the scallops grow at. To carry out the laser measurements, I needed geochemistry experts, and Lennart de Nooijer, Wim Boer and Gert-Jan Reichart came to mind because I was already working with them on other topics and they run a very good lab for these analyses at the Royal Netherlands Institute for Sea Research (NIOZ). This is how the team came together.
Niels conducted a series of laser transects across the clam shells. He used some sophisticated time series analysis approaches to try to quantify the different periodic cycles that appeared in the clam and scallop growth. This was a different approach to how other workers have gone about finding daily growth cycles in giant clams and scallops, where they have often started by zooming in to find the wiggles, and work backwards from there. Niels instead tried to agnostically dissemble the growth records across each clam shell using mathematical approaches, based on the idea that this would be how future workers have to go about identifying daily growth patterns in fossil clams, where we often don’t have a real “growth model” up front to work with. By growth model, I mean the way that we convert the geochemical observations, which are arranged by distance along the shell, into units of time, which requires us to know how fast the clams grew. For the scallops, the age model was made by counting daily “striae” they form on the outside of their shells. For the giant clams, I helped with this by counting tiny growth lines inside the shell made visible by applying a dye called Mutvei’s solution. Because the growth lines weren’t visible all the way through the shell, I used a von Bertalanffy model to bridge across and create a continuous estimate of how old the clams were at each point along their shells.
Niels found some interesting results! I personally expected that the daily variation in giant clams would dwarf what was seen in the scallops, because of the impact of the daily activity of the symbionts. But it turned out that while the clams had a more regular pattern of daily shell growth than the scallops, likely controlled by the symbionts, that was still a minority of the variance across the clams’ records. Yet again, these clams destroyed my hypothesis, but in an interesting way!
Niels, what were your expectations going into this, and how did the results confirm or go against your hypotheses? What challenges did you run into in the course of your analysis, and how did you end up addressing those challenges?
This was honestly one of the most difficult shell-datasets I have worked with so far. The laser technique we used measures the elemental composition of the shells in very high detail, but while this is ideal for funding daily rhythms, it is both a blessing and a curse! In a dataset like this it becomes quite hard to separate the signal we are interested in from the noise that occurs due to measurement uncertainty. I ended up using a technique called spectral analysis, which is often used to detect rhythmic changes in successions of rocks. I guess this is where my geology background was helpful. With this technique, we were able to “filter out” the variability in the records of shell composition that happened at the scale of days and tides and remove the noise and the longer timescale variations. It turns out that, when you do this, you have to remove a surprisingly large fraction of the data, which shows us that the influence of the daily cycle on the composition of both the scallops and the clams is not very large (at most 20%). We did find a larger contribution in the giant clams, as expected, but the difference was much smaller than anticipated. I also find it interesting that most of the variability was not rhythmic. This shows that there are likely processes at play that control the composition of shells on a daily basis which we do not understand yet.
We were measuring a suite of different elements across both bivalve species, including strontium, magnesium, manganese and barium. All of these were reported relative to calcium, the dominant metal ion in the shell material (they’re made of calcium carbonate). This is why we call them “trace” elements; each is integrated into the material of the shell due to a variety of causes, including the temperature, the composition of the seawater, the growth rate of the clams, and also simply due to chance.
Examples of the time series of trace elements from a scallop shell (to the left) and giant clam (to the right), showing the very intricate wiggles in trace element values on a on a tidal and daily basis in each bivalve
In the giant clams, the elements that varied most on a daily basis were strontium and barium. Prior workers had found strontium was the strongest in terms of daily variation, but barium was more unexpected! Normally, barium is thought of as a record of the activity of plankton in the environment, and since there is very little plankton to be found in the Red Sea, it was not expected to see that element vary on a daily basis. It could be that barium gets included in the shell more as a function of the growth rate of the animals. Meanwhile, the scallops (from the Bay of Brest in France) were measuring strong tidal variability in barium and strontium, which makes sense because that location has huge tides compared to the Red Sea. Tides happen on periods of ~12.4 and 24.8 hours. The scallops showed swings lining up with both, and the tidal variability might be the main explanation for how scallops form daily lines. Because the lunar day is so close to a solar day, they would be hard to tell apart from each other! Interestingly, the giant clams also showed some sign of a ~12 hour cycle. While the Red Sea has pretty tiny tides, I had noticed that some of the clams make 2 growth lines a day, and if some clams in the shallowest waters were exposed on a tidal basis, that could explain why they’d make 2 lines: one at low tide, and one at night! Even in places without tides, like the Biosphere 2 ocean, I’d noticed evidence of 12-hour patterns of activity in the clams. It’s so nice (and rare!) when one of my hypotheses is confirmed!
A nice schematic Niels put together showing all the environmental factors that influence the shells of scallops and giant clams, and how much different elements vary as a function of sunlight, tides and other more irregular events like storms. Mn stands for manganese, Ba for barium, Sr for strontium and Mg for magnesium.
Both the giant clams and scallops recorded large irregular swings in all of the studied elements, likely due to non-periodic disturbances. In the case of the scallops, these included storms and the floods of sediment from rivers. For the giant clams, these probably included algae blooms that affect the Red Sea, as well as potentially dust storms that also come every 1-2 years. Both giant clams and scallops have a lot of potential to measure paleo-weather, which is something that other researchers have observed as well!
Niels, where do you see this work heading next?
The recent work looking at very short-term changes in shells is very promising, I think. I agree that there might be a possibility to detect weather patterns in these shells, but that would require some more work into understanding how these animals respond to changes in their environment on an hourly scale and what that response does to their shell composition.
In the meantime, I was intrigued to find that we were not the only people looking for daily cycles in the chemistry of giant clam shells. I had the pleasure of reviewing this paper by Iris Arndt and her colleagues from the university of Frankfurt (Germany). Iris took a similar approach to detecting these daily cycles by using spectral analysis, but she a smart tool called a “wavelet analysis” to visualize the presence of daily rhythms in the shell, which I think was more successful than my approach. She even wrote a small piece of software which can be used to (almost) automatically detect the days and “date” the clam shell based on them. This is quite a step forward, and if I were to do a project like this again, I would certainly try our Iris’ method.
Interesting, too, is that the fossil giant clams studied by Iris showed the daily cycles in magnesium concentration instead of strontium and barium. This shows that the incorporation of trace metals into clam shells is still not fully understood. So one of the things to do, in my opinion, would be to try to see if we can use shells grown under controlled conditions to link the shell composition to short-term changes in the environment. This would require a complex experimental setup in which we simulate an artificial day and night rhythm or an artificial “storm”, but I think it can be done using the culture experiments we do at the NIOZ.
This study represented a unique opportunity to collaborate with my colleague Niels on a topic that interested both of us, which we wouldn’t have been able to pursue on our own. I enjoyed collaborating with him on this work and we have some ideas for further studies down the road, so stay tuned for the next co-clam-boration!
Every clam is a door into the sea. If the “door” of its shell is open, the clam may be happily breathing, or eating, or doing other weirder things. If the door is closed, it may be hiding from a predator, or preventing itself from drying out at low tide, or protecting itself from some other source of stress. It turns out that by monitoring the opening and closing of a clam’s shell valves, a field called valvometry, scientists can learn a lot about the clam’s physiology, its ecology and the environment around it.
Valvometry involves attaching waterproof sensors to each shell valve of the bivalve, to measure the distance between them and their movement. Researchers have used valvometers to figure out that bivalves can be disturbed by underwater pollution like oil spills, harmful algal blooms, and more unexpected sources such as noise and light pollution.
A great video from Tom Scott discussing a Polish program to monitor water quality with valvometry
Giant clams are a group of unusually large bivalves (some species reach up to 3 feet long!) native to coral reefs of the Indo-Pacific, from Australia to Israel. They grow to such large size with the help of symbiotic algae living in their flesh, the same kind that corals partner with the corals that build the reefs. The algae photosynthesize and share the sugars they make with their host clam, and the clam gives the algae nitrogen fertilizer and other nutrients, a safe home from predation and even helps channel light to the algae using reflective cells called iridophores.
A Tridacna derasa clam in the Biosphere ocean. It has deep green flesh, covered with yellow stripes of iridophores and a blue fringe at the edge.
Previous studies have used valvometers on giant clams, but I was always perplexed by how few studies there were: only two that I know of! One study on clams in New Caledonia figured out that the clams partially close every night and bask wide open during the day. The clams’ shell opening behavior and growth was found to become more erratic at temperatures above 27 °C, and when light levels become too great. Another study showed the clams start to clam up when exposed to UV light to protect themselves from a sort of sunburn, which is a real threat in the shallow reef waters they live in.
Two clams sitting next to each other in the B2 ocean. They often moved themselves to “snuggle” next to each other this way. Safety in numbers!
There is clearly a lot of information to pick up about how clams react to their environments, which can help us understand the health of the clams and also the corals around them. Coral reefs are under global stress from climate change, overfishing and pollution. Giant clams are some of the most prolific and widespread bivalve inhabitants of reefs, and represent an appealing potential biomonitor of reef conditions. Many giant clam species are threatened by the same stressors that influence the corals which build the reefs they live on, as well as overharvesting for food and their shells. For that reason, wild examples should clearly not be bothered by applying valvometric sensors. But giant clams are increasingly grown for the aquarium trade, resulting in a wealth of cultured specimens which could serve as sentinels of reef health, if they were fitted out with sensors. All of these motivators made me more and more curious of why we don’t have more literature monitoring the behavior of these clams with valve sensors.
I wondered if one of the limiting factors preventing the use of valvometry on giant clams is expense and ease of access. Giant clams live primarily in regions bordering developing countries in the Indo-Pacific, and almost all the professional aquaculture of clams for the reef trade happens in such countries, including places like Palau, Thailand, and New Caledonia. These countries are far removed from the places where most of the proprietary valvometric systems are manufactured. These systems can cost several thousand dollars even in Europe, never mind Palau, where arranging the import of electronics can be difficult.
When I started my postdoctoral fellowship at Biosphere 2 in 2020, I set out to grow two dozen smooth giant clams (Tridacna derasa, a species which can grow to about 2 feet long) in the controlled environment of the Biosphere 2 ocean, a 700,000 gallon (over 2.6 million liter) saltwater tank used to grow corals and tropical fish and kept at a stable year-round temperature of 25 °C. We suspended a series of LED lights intended to simulate the powerful light levels these clams experience in the wild (light is a lot brighter in the tropics than it is in Arizona!). The main focus of my project involved measuring the shell chemistry of the clams, to determine how their body chemistry changed as they grew from mostly getting their energy from filtering algae food from the water like other clams, to getting most of their energy from sunlight like a plant. But as a “side project” I set about measuring the behavior of the clams with custom-built valvometers based on open-source, inexpensive hardware that would be more accessible to researchers in the developing world. That work has since been published in PLoS One!
In our design, we used Hall effect sensors. Hall effect sensors generate a voltage when a change in magnetic field is detected. They are cheap, easily obtained for less than $1.50 apiece and are common in the electronics hobby trade. You might have encountered one in a home security system door/window sensor, where they help detect if a door is open or shut. We stuck a hall sensor soldered to a long copper cable to one valve of a clam, and a small magnet to the other valve. When the clam closed, we could measure exactly how closed it was. You can see why I started off by calling clams doors into the sea: we were literally measuring them that way!
Showing the sensor soldered to the three strands of the cable.
But here the first challenge of my project appeared. The off-the-shelf Hall sensors don’t come in waterproof form, and I learned quickly that the ocean really, really loves to break my gear. After dozens of failures, I settled on coating the sensors in waterproof grease, wrapping that in heat-shrink tubing and then sealing that inside of aquarium-grade silicone. During this process, a gifted technician at Biosphere 2 named Douglas Cline helped with iterating on the first prototypes. At a certain point I taught myself to solder so I could do my part to improve the sensors.
It was also hard to figure out how to attach the sensors and magnets to the clams in a durable way. Neither of the prior studies mentioned how they attached the sensors to giant clams, and I tried and failed with literally a dozen different ways before settling on “pool putty,” a two-part adhesive often used to seal leaks in pools that can cure underwater. I found the pool putty had trouble attaching to the clams’ shells on its own, so I combined it with a special kind of cyanoacrylate superglue called “frag glue,” often used to attach pieces of corals to growth stubs. I also had to find a way to attach it to the clams without stressing them out. I determined five minutes out of the water was enough time to get the sensors attached to the clams, after which they could be returned to the water to finish curing. While giant clams are adapted to spend extended periods out of the water in their natural intertidal environment, we wanted to make sure to minimize their stress however possible, to ensure they would show natural cycles of behavior in the data.
Figure from the paper showing: A) schematic of the sensor attached to the clam, linked to an Arduino microcontroller and Raspberry pi computer. B) A sensor attached to one of the clams
We were pleased to see the cyborg clams seemed to pay no mind to the sensors. Giant clams are adapted to encourage all sorts of other critters to live on their shells as a form of natural camouflage, and I think the clams interpreted the sensors as pieces of coral or anemones sticking to the side of their shell. Whatever the case, as long as we kept the cable pointing to the side away from the clams’ flesh, they opened five minutes after being returned to the water, and their behavior and growth rates were indistinguishable from the clams that didn’t have sensors attached.
One of what became many sunsets on the Biosphere 2 ocean shore troubleshooting the clam sensors! Pardon the chaos: mad scientist at work!
So how did we measure the voltages coming from the sensors? Our design featured an Arduino microcontroller, sort of like a smart circuit board which can measure the voltages coming back over the copper cables. Arduinos are very cheap, and we chose a $25 model. Even more importantly, Arduino has a huge library of plug-ins available to keep the exact time of each observation using a clock attachment, and the data can be uploaded to SD cards or an attached computer. For the attached computer, I used a Raspberry Pi computer, which are open-source Linux-based tiny computers that are very cheap! Or rather they were very cheap before the pandemic, but fortunately there a lot of open-source alternatives that can be obtained more cheaply. We logged the data on the Raspberry Pi as it rolled over from the Arduino, and I could watch the read-out on a monitor right on the Biosphere 2 beach. We set the Arduino to record every 5 seconds.
Sensors attached to four of the clams. Notice the one on top left has closed a bit, after sensing my presence! They have eyes so they were able to detect me 😀
We ran the sensors for three months. During that time, the baby giant clams grew almost an inch! What did the sensors record them doing? During the day, the clams basked wide-open, exposing as much of their tissue as possible to light (other than the times that I disturbed them by swimming above them, of course)! This schedule of opening aligned pretty closely with the times that maximum sunlight hit their part of the Biosphere 2 ocean: the mornings, because the clams were on the east side of the building. At this time of day, the clams want to expose their symbiotic algae to as much light as possible, so they can conduct photosynthesis and make sugars that the clams use as food!
A) Plot of the valvometry data. Points higher on the plot mean the clam was more closed, up to 100% closed. The clams proceeded by opening in the early morning and then closing in the early afternoon. The big red circles represent times that the clams closed briefly, with bigger circles representing a longer time spent closed. Most of these rapid closures happened at night. B. A plot of Photosynthetically active Radiation (the amount of light the clams had to use for photosynthesis). The highest values were in the mid-morning when the clam lights were running in combination with direct sunlight hitting them from above.
Around mid-afternoon, the clams started to close partially, to about half closed. Why might that be? My hypothesis is that this posture represents a kind of “defensive crouch” to protect themselves from predators, in this case fireworms that live in the Biosphere 2 Ocean and were constantly kicking the clams’ tires. Similar nighttime behavior was observed in wild clams in a previous study, but not in a study that took place in a small predator-free terrarium tank. By remaining partially closed, the clams are prepared to rapidly close completely if they feel a predator approaching. But they only expend that energy of staying in that posture if predators are around!
One of the fireworms that proved to be my nemesis and continually attacked the clams during the experiment
And approach the fireworms did. We observed frequent closures at night lasting anywhere from a few seconds to hours, likely partially related to the activity of the worms around the clams. But the clams were engaging in another activity at night: filter feeding! Giant clams really get to have their cake and eat it too, because during the day, they act like a plant, but at night, they eat other plants in the form of plankton that they filter feed out of the water using their gills! At regular intervals, the clams need to clear uneaten material from their gills in a process sometimes called “valve-clapping”. The clams yank their shell valves together rapidly to force water out, blowing out pseudofeces: unwanted material packaged with mucus. We measured this valve-clapping mostly at night. The clams are likely scheduling this activity for the night-time so they can prioritize staying open and filter feeding during the day!
Figure comparing how often clams closed per day to measures of how high plankton numbers were in the Biosphere 2 ocean (chlorophyll is a marker of phytoplankton while phycocyanin is a measure of cyanobacteria), and how high the light levels were. Peaks in closure activity often happened shortly after rises in algae.
We observed that the frequency of valve clapping aligned closely with the rises and falls of chlorophyll concentration in the Biosphere 2 ocean, which is a measure of how much plankton is in the water column. The clams would engage in a burst of valve clapping around 4 days on average after a bloom in chlorophyll, suggesting they were filtering out plankton after they had died and settled to the bottom where the clams could eat them. We also found that the clam’s filtering activity peaked at times of highest pH. This likely is due to the fact that higher pH means the algae around the clams are being more active, and pulling CO2 in from the water to use in photosynthesis, making the water less acidic. More photosynthesis means potentially more material for the clams to filter through! This data helps quantify how giant clams help filter the water in their native environments! Coral reefs depend on very clear transparent water to allow maximum sunlight to reach the corals, and the filtering activity of giant clams likely plays a big role in helping preserve those conditions!
So we found that by adding sensors to clams, we could record their ability to feed from the sun, their feeding on plankton around them and their avoidance of predators. How can this technique be used next? We hope that by using cheap off-the-shelf resources and open-source software, we can enable more sensors to be put on clams all over the world, such as places where giant clams are farmed in Palau, New Caledonia, Thailand, Taiwan, Malaysia and more! If we can collect data on clam activity from all these places, we can compare how their feeding patterns differ in places that have more or less plankton floating by, or have more or less sunlight available, or different predators that affect the clams’ behavior. This data would have importance to the clams’ conservation, as well as our understanding of the reef overall. In future years, I hope we can develop a global network of cyborg giant clams from the Red Sea to the Great Barrier Reef, so we can better understand how these oversized and conspicuous but still mysterious bivalve work their magic!
Abraclam Lincoln (photo source) next to a handful of Mercenaria mercenaria, a closely related kind of quahog (photo from Wikipedia), illustrating that Abraclam is indeed a chonker. To the right we have Arctica islandica, the famous long-lived clam, illustrating how hard it is to tell these clams apart without specialized training (photo from conchology.be).
Recently, a bit of an amusing clamfuffel arose when the Gulf Specimen Marine Lab, a research institute in Florida, began posting about a supposedly 214-year-old clam they named “Abraclam Lincoln”, in honor of potentially sharing a birth year with Honest Abe. The story went viral, and while some of their clammy claims turned out to hinge on flawed assumptions about how clams can be aged, it was still a worthwhile opportunity to communicate about the wonderful world of bivalves with people.
I was particularly impressed with how GSML went out of their way to correct the record as more info came in. Long story short, Abraclam is not the long-lived ocean quahog Arctica islandica (more on them below), but in fact a specimen of the southern quahog Mercenaria campechiensis, which lives along the Gulf coast and grows much more quickly than A. islandica. So the large clam they found was likely more like 40 years old, which is not too unusual for this species, rather than two centuries old. It also makes much more sense to find Mercenaria in Florida than Arctica. Additionally, external shell growth lines in Mercenaria are known to not be reliable for aging the species. The shell would have to be cut open to view internal lines to figure out how old Abraclam really is, which would require killing them. Fortunately, they released the clam rather than cut them in half for science.
Known range of Arctica islandica as a heatmap, compared to location of GSML on the Florida Panhandle. Source: OBIS
Slice through a Mercenaria campechiensis shell. The shell is lit from behind to show the annual growth lines. From Moss et al. 2021, written by fellow clam man David Moss, a friend of mine!
For what it’s worth, I still think Abraclam is an interesting specimen of M. campechiensis– they have a huge scar in their shell that would be interesting to learn more about (maybe they were hit by a dredge in their youth, healed the break and recovered to reach a ripe old age), and an unusual undulating margin at the edge of the shell that could be a deformity reported in the past literature for this species. Sometimes, scientists are wrong the first time, but we’re open about how we’re wrong, and everyone ends up learning more than we would have known otherwise.
So far so good. Then a bull had to wander into this delicate china shell with the entry of Stephen Colbert into the debate. I’ll let Stephen speak for himself, but needless to say, after his rant I feel a need to respond:
Before I dig into this clambake, Stephen, kudos to you for covering the whole story and not just the initial incorrect information. You addressed all the big inaccuracies, from the size not being tremendously out of the ordinary, to the incorrect species ID, to the incorrect age. Maybe it’s not so bad that people are getting their news from comedians rather than the news media these days! But while there were definitely some pearls of wisdom within your monologue, I have to point out some misconceptions here.
“The only thing more heartbreaking than the lies we were fed in this story…is growing up to be a clam expert!”
– Stephen Colbert
This is just plain false, since I’m not heartbroken, because clams are frickin’ awesome. Clams are way cooler than you or me, and that means by extension that the people who study them are pretty cool and interesting too (not really referring to myself. I’m just an eclamgelist along for the ride). So here are three facts about big old clams, and information about the clam experts who discovered these facts!
The ocean quahog lives to >500 years old!
Arctica islandica shell I saw on the beach in Massachusetts. This individual was likely several decades old when it died based on its fairly large size!
The ocean quahog Arctica islandica (which Abraclam was initially misidentified as) is tremendously long-lived, one of the longest-lived animals on earth! It has been confirmed to live to at least half a millenium! One individual caught off the coast of Iceland was aged to ~507 years by counting tiny growth lines in its shell via microscope, combined with radiocarbon dating. This clam was named Hafrún (meaning “ocean mystery” in Icelandic), but is sometimes called Ming due to it being born in the Ming dynasty of China. So put that in your Ming vase and smoke it Colbert!
Part of the shell of Hafrún, which was cut open to determine its age via internal growth lines. Source: Museum Wales
Several scientists worked on aging Ming, but Alan Wanamaker at Iowa State was a lead author on the original work. He uses growth lines in the shells of many clam species as records of climate change and is generally one of the nicest people you could have the opportunity to meet.
Tridacna gigas grows to over 4 feet long and hundreds of pounds!
Large specimens of giant clams that are around 3 feet long at the California Academy of Science collection.
The author sitting in a scale model of Tridacna gigas at the Monterey Bay Aquarium
Abraclam might be only 6 inches long (which is respectable as he/she is quite girthy; length isn’t everything, Stephen!), but there are other types of clams that are bigger than one Colbert in mass. The giant clam Tridacna gigas grows to over 4 feet long and weighs hundreds of pounds. They live on tropical coral reefs and use the power of the photosynthetic algae in their flesh to speed up their growth. So basically these clams are bigger and way more interesting than you, Stephen, since they get to go out and tan in the sun for lunch while you have to gobble down a slice of pizza.
Mei-Lin Neo at University of Singapore is considered the world’s leading expert on Tridacna, and has done more than almost anyone I know to describe all twelve currently known species of giant clams found around the world. She’s a tremendous advocate for giant clam conservation and gave an outstanding TED talk about them to boot. You should have her on your show to be honest.
Geoducks: they looked like that first!
A 6.5 pound geoduck and admiring Washington Department of Fish and Wildlife Volunteer (Source)
Speaking of girthy, long clams, I’d be remiss not to mention the geoduck, Panopea generosa. Pronounced “gooey-duck,” these clams looked like this long before any part of human anatomy existed, having been around in various forms since at least the Jurassic. They have a long siphon that they use like a snorkel when they dig deep in the mud, and they can live for almost 200 years.
Brian Black at the University of Arizona is an expert in using their shells as a record of climate change. He was part of a group that was able to stitch together the growth line records from multiple geoduck shells to make a continuous record of climate change going back to 1725. Seems appropriate to note that 1725 was the year that Casanova was born…a man who may have channeled some qualities of geoducks.
Local experts on Abraclam
I’d like to mention two of the experts who corrected the record about Abraclam Lincoln and provoked Stephen’s attack in the first place. Dr. Dan Marelli wrote an op-ed correcting the record on how Mercenaria clams are aged for the Tallahassee Democrat. He’s an expert scientific diver and has published papers on clams ranging from endangered scallops to invasive mussels. Scientific diving is crucial to understand clams in their native environment, and to assist in their conservation. If I had to choose who had more interesting stories at the bar, it’d be an easy decision to listen to the swashbuckling diver over the late-night TV host!
Dr. Edward Petuch at Florida Atlantic University reached out to GSML to make sure they knew the correct species ID for Abraclam. He is well-known for his work describing the change in ecology of mollusks in Florida and the Caribbean over the last several million years. GSML expressed interest in working with Dr. Petuch in the future, and I can confirm that I’ve had fruitful scientific collaborations start when other scientists have reached out to me about how I was totally, embarrassingly wrong. Being wrong in science is part of the job, and that’s why I’m glad this Abraclam story came out in the first place.
“So what does your son do? He’s a marine biologist. Does he work with dolphins? …I’m gonna say yes.”
-Stephen Colbert
To close out, I’d like to address Stephen’s assertion that my mom isn’t proud of me for being a clam expert. Stephen, I’ll have you know that my mom is the most enthusiastic patron of my clam science. She reached out to the local paper to anonymously tip them to interview me about my clam work, had me give a speech about clams at the local women’s group she’s a part of, and when I defended my PhD thesis, she made t-shirts to commemorate the occasion. I can confidently say I wouldn’t be Dan the Clam Man if it weren’t for her support. Thanks Mom!
A geoduck clam (also called “elephant clams”) next to the elephant-like Mastodon mascot
Over the last couple weeks, I’ve seen hundreds of academics, nerds and everyday people I know open new accounts on Mastodon, in a phenomenon that has been called the great #TwitterMigration. Mastodon is an open-source microblogging platform similar in format to Twitter, but running on thousands of servers interconnected with each other in an open network called the “Fediverse” (referring to the fact that these services are “federated” to each other). Many researchers are disillusioned with the current state of Twitter, which was purchased recently by an erratic, bigoted oligarch, and are registering their disapproval by seeking out other places to share their science.
Personally, I am not “migrating” per se, as I have been using Mastodon for over 4 years now. I don’t intend to close my Twitter account, because I think the site will survive the current damage being done to it, though I’ve stopped posting for the time being, while they work through the process of learning the hard way that hate speech can never be allowed on the platform. I wanted to write about my experience using Mastodon to communicate science and why I think it has a lot of advantages over Twitter for certain use cases, precisely through the ways it does not seek to be a direct Twitter replacement.
Posting how I want
First of all, Mastodon gives me way more freedom to post the way I want. It is a true “micro-blog” in the way Twitter can’t be, since I get enough characters (500 at scicomm.xyz, and more on some servers!) to allow me to post a real paragraph. I have never found Twitter’s 180 characters to be enough space to really tell a satisfying story. Some people get around this by posting threads, but I also have never enjoyed writing threads! Other than that time I went on a giant clam fact rant. Mastodon also supports threads if you’re into that, and also allows you to set the visibility on your subsequent posts, so that your replies to yourself don’t spam everyone’s feed.
In my four years writing #clamfacts on Mastodon, I’ve written short facts. Long facts. Silly facts. Meaningful facts. I just have a lot more freedom with the format. Mastodon was also much faster to enable accessibility features like image descriptions than Twitter was. So the facts I shared are more accessible to disabled folks. While Twitter now includes alt-text, it still feels like Mastodon’s alt-text is a more mature feature. As with subtitles and other accessibility solutions, these features end up improving usability for everyone. In the case of alt-text, it gives me tons of space to describe scientific diagrams for anyone who might need additional context.
Mastodon recently added the ability to edit posts, which has been very advantageous for me, as a typo-prone individual. For Twitter, that feature is still locked behind a subscription. But even before editing was available on Mastodon, there was an option to “Delete and redraft”, which I used frequently to re-post when I had forgotten an image description, or to fix a typo. Mastodon has long provided far more options to control who can see a post and how, which is why I felt more incentive to be creative there than on Twitter.
A small pond by design: Engagement and sustained connection over reach
Twitter sometimes feels like an RSS feed with comments. Particularly for the more popular accounts or viral posts, while you can reply, there is such a torrent of feedback on the other end that it is difficult for them to respond to everyone. For my niche specialized clamposting, I am not interested in going viral. I just want to engage with people and learn from them as much as I share knowledge with them.
Mastodon is very well designed for this. Your posts get shared across the federated network, but only as much as other people “boost” it (analogous to retweeting) or reply to it. There are favorites (similar to the like button), but those are mostly just a direct message that someone liked a post, and have no impact on whether it spreads larger over the network. So the main people seeing my posts are my direct followers. And because Mastodon is a reverse chronological feed, with no opaque algorithm determining whether or not to show someone something, I am more confident that a bot moderator isn’t going to misidentify my clam content as NSFW and hide it by default from people’s feeds.
Culturally, Mastodon is driven by following people, and making your feed for yourself, rather than having posts from people you don’t follow pushed to you by a computer. If you follow someone back, you’re more likely to make a lasting connection through time, rather than trusting some algorithm to figure out who you enjoy to see. This leads to more lasting, meaningful connections in my experience. Truly powerful scicomm never happens in one direction; it relies on exchange.
I think that Mastodon will stay like this in the future, even as it continues to grow by leaps and bounds. Rather than one giant, sometimes dangerous ocean like Twitter, it’s more of a collection of small ponds. My reach is restricted to my followers and their followers, and sometimes their followers’ followers. That produces much more meaningful, sustained connections.
Hosted and moderated by scientists, for scientists
My home since the start has been Scicomm.xyz, a server run by a scientist in the UK going by the username Quokka. Recently he recruited another scientist and me to be moderators on the server, and we’re looking to add more. But since the start even before I was moderating myself, I’ve felt more secure sharing science when it’s hosted and moderated by another scientist. Even before Twitter laid off moderation staff en masse, and before the site announced scientific misinformation is now fair game, Twitter was not a place run by scientists, for scientists. If someone replied to me with misinformation about the coronavirus or climate change, my recourse against them was limited, since the moderation staff there are not exactly experienced peer reviewers. On Mastodon, there have always been data-conscious nerds running things. And now, there are a constellationofscience–specificserverstochoosefrom!
Science, including scicomm, is always more at home in an open-source environment
The last point I’ll bring up is that science always works better in an open-source environment. Mastodon is available free and open-source on Github for anyone to download, alter and run themselves. I prefer to use such open-source solutions in my own scientific work, from including Rstudio, QGIS, ImageJ, Raspberry Pi, Arduino, Ubuntu, Inkscape, Firefox/Thunderbird and more. So hosting my science communication on an open platform feels like preaching what I practice, as opposed to allowing a for-profit company to own my scientific content.
For all the reasons above, I have been extremely pleased to see the wave of scientists, technologists and other interested people join Mastodon over the last few weeks. I feel Twitter will still have a place in my science communication once it has worked through its current drama. But in the meantime, I look forward to sharing my clam facts with all the people I can, in my little pond on Mastodon.
The tree of life is often portrayed as a neatly branching structure, with each division point cleanly delineated and separated from its neighbors. The truth is that the various twigs of the tree of life often overlap and become tangled in a process we call symbiosis. I’ve talked about symbiosis before on this blog, which falls along a spectrum of wholesomeness. At one end we have mutualism, a partnership where both organisms benefit and achieve more than the sum of their parts. The other extreme is parasitism, when one organism benefits at the expense of the other. Between the two, there is a broad gray area including commensalism, when one organism’s presence doesn’t necessarily cost or benefit the other in any way. The tree of life is crowded and unpruned, and so sometimes the twigs might wrap around each other quietly and without much fuss. We live on a small planet, and have had to get used to living in uncomfortable intimacy with all sorts of creatures, such as the mites that are living on your eyelashes right now.
But things start to get really weird and tangled when the tree of life loops over on itself twice, or three times, or more. “Three-way” symbioses are surprisingly common, and the more you look for them, the more you realize that the tree of life is more of a knot than anything else.
A view of the Heteropsammia coral from the side
A recent paper from researchers in Bremen (Germany) and Saudi Arabia looked at such a three-way symbiosis between a coral, a worm and bivalves found off of Tanzania in East Africa. The relationship between solitary corals (Heteropsammia cochlea and Heterocyathus aequicostatus), a sipunculan worm (Aspidosiphon muelleri muelleri) and the clam Jousseaumiella, is a complex triangle of dependencies that had previously been noticed by other researchers, but never investigated at great depth. The worm lives with multiple tiny clams attached, all inside of a small solitary coral the size of a dime (1 cm long). Is the coral a willing host for this crowded boarding house, or has it been parasitized? Does the worm gain anything from the clams? The researchers sought to find out.
Part of the reason I enjoyed reading this study so much was that it had to take a narrative structure to describe the evolutionary ménage à trois of its focus. So much of modern science has moved away from anecdote to hard data, and while there is plenty of that to find in the study, it turns out that a lot of the study of symbiosis is storytelling. We need to know the setting and the characters.
In this case, the main characters are small solitary corals living in the tropical reefs of the Indo-Pacific. We denote them as solitary to distinguish them from their giant colony-forming compatriots that construct the coral reefs currently threatened by climate change and pollution. But like those giant reef-builders, these solitary corals get much of their food from sunlight through a mutualistic partnership with algae called Symbiodinium. The algae provide the host with sugars and other photosynthetic products, and the hosts give them nutrients and a safe cozy home in their tissue.
You might be thinking, “Wait! Dan just said this was a three-way partnership between a coral, a worm and some clams. So this is actually a four-way partnership between corals, worms, clams and algae?” You’d be exactly right. And I’m happy to say that the plot of this sordid story is about to thicken even further.
The side of the coral. See the little pores?
The Aspidosiphon worm is found in a spiral-shaped burrow inside of the skeleton of the coral. It is a pretty cozy home, with walls made of calcium carbonate by the coral, with breathing holes in the sides to allow the worm to breathe and release waste. The researchers wanted to know more about the structure of the burrow. Was it dug out by the worm using acid or an abrasive motion, like some clams use to dig into coral? So the researchers essentially gave the coral a CT scan to see its 3D internal structure. Inside they found growth features suggesting the coral grew around the worm, as if intentionally providing it a home.
Cross-sectional CT scans of the coral skeleton. In figure D, you can see the silhouette of the chambers of the snail shell where the worm made its first home!
Even more crazily, they found evidence that the worm had first settled inside an empty snail shell, like a hermit crab! The coral probably settled on a snail shell as a larva, and grew to engulf the whole snail shell, leaving growing space for the worm inside, with windows and all! So to review, this is now a five-way symbiosis between a dead snail, a worm that moved into its empty shell, the coral (powered by algae) that grew around it and encased the snail shell within its skeleton, and we haven’t even gotten to the clams. How many creatures are hiding stacked in this trench coat? Please bear with me as I explain!
An SEM image of Jousseaumiella. These are less than 1 mm long! Pinhead sized!
What are the clams doing in this picture? Jousseaumiella is part of a family of clams called Galeommatidae, which we previously mentioned on this blog in the context of some bivalves found growing in the gills of unfortunate sand crabs. Many members of the Galeommatidae family are parasitic or commensal with other marine organisms. In this case, Jousseaumiella are tiny flat-bodied clams less than 1 mm long, found attached to the body of the worm, squeezed inside the burrow in the coral’s skeleton. It feeds on the worm’s waste and potentially food particles coming through the pores in the sides of the burrow. Not the most dignified existence, but a more mobile home means more opportunities to eat a varied diet similar to that that the worm and coral are seeking out, and the clam also gets protection from predation tucked inside the coral. It is unclear if it benefits the worm directly to have clams attached to it.
A time lapse of the coral+worm moving from the paper’s supplement! It would be handy to navigate to greener pastures, if it became too muddy in a certain place!
It is, however, clear how living inside a coral would be a pretty good deal for the worm, which gets a stable, protective suit of carbonate armor to protect it from predators, and grows to fit it as it gets larger. They are normally found inside of rocks, shells and other hard inanimate objects, but having a living home is a cool upgrade. What is the coral getting out of the deal? The researchers note that the corals are often found in the crevices between other large reef-building corals, in areas of the reef that receive high supplies of nutrients and turbidity (dirt that blocks out light). These sorts of environments aren’t necessarily friendly places for a coral to be, since they reduce the light and therefore the food that the coral can receive from photosynthesis. These crevices also have a lot of variability in other conditions like temperature and water flow. But because the coral has hitched a ride on the back of a worm, it can actually move in the sediment to react to changing conditions and avoid being buried by piles of sediment floating by! The worm can also act as a sort of anchor preventing the worm from sinking in the sediment underneath, which would be a big hazard for the small, stubby coral on its own. The coral seems to go to great pains to make its partner comfortable, not growing its skeleton to cover the pore windows to the outside. The researchers note that as coral reefs worldwide are subject to increasing human-made pollution and climate change, it would be interesting to research whether this complex three-(five?) way symbiosis provides the various participants with an advantage compared to other corals.
So like any good story, this symbiosis features complex, growing characters, a dynamic setting, and still plenty of mystery demanding a sequel! To that end, there are lots of other great three-way symbioses to investigate. Snails which farm fungus that parasitizes plants. Bryozoans living on snail shells that have a hermit crab inside. Gobies serving as lookouts at the entrances of burrows built by shrimp, with a crab freeloader along for good measure. Algae and bacteria teaming up to attack mussels. The list keeps going! I could see this becoming quite a franchise!
Richmond, CA from the air, showing the turbid waters of the SF Bay
Well folks, it finally happened. I found a permanent scientific job. On January 31st, I’ll be starting as an Environmental Scientist at the San Francisco Estuary Institute (SFEI), working on the Nutrient Management Strategy (NMS) program. NMS is a group trying to understand how nutrient supply in the San Francisco Bay works.
The SF Bay is an extremely nutrient-enriched environment (eutrophic) due to human pollution and natural factors, to the extent that if all other factors were equal, scientists would expect it to be a nasty green sludgy mess. Yet up to today, due to factors that are still debated, the SF Bay is in much better shape than it should be. It is not a dead zone, choked off by algal blooms and oxygen-starved in the way that other high-productivity regions such as parts of the Gulf of Mexico have become. Those factors may include the cloudiness (turbidity) of the Bay’s water limiting algae growth, naturally rapid tidal mixing with ocean water, and the influence of clams and other grazing animals keeping the populations of potentially harmful plankton suppressed.
However, there is also evidence that this resilience may be fading as water temperatures in the Bay increase and the ecology of the system changes with climate change. Oxygen levels are dropping and levels of harmful algae are rising, which endangers the health and livelihoods of millions of people in the SF Bay area who depend on a clean, ecologically functioning SF Bay. In my role at NMS, I will be assisting in processing and interpreting huge quantities of environmental data on temperature, dissolved oxygen, water flow, light levels, algae concentrations, and harmful algae toxins, to help figure out how the SF Bay works and how we can protect it. I will be assisting another scientist joining the team in deploying more sensors to monitor the Bay on a minute by minute basis, and also packaging the data to help create models which allow us to figure out the various moving parts that make it work.
In a way, this is oddly similar to the work I’ve done during my postdoc at Biosphere 2, where I’ve been growing giant clams in their 700,000 gallon ocean tank since May 2020. The clams are biological sensors have been recording the environment of the Biosphere 2 ocean through their shells and valve opening/closing activity, and I have had to decode their diaries through comparison with the environmental data we collect on light, pH, dissolved oxygen, chlorophyll and other measurements. The SF Bay is a site of enormously influential research which has been important to understand estuaries around the world, but it is still a mysterious body of water in many ways. NMS is trying to understand how all its complex pieces fit together, much like I’ve been doing at Biosphere 2, which is why I jumped at the opportunity to apply for the job.
I also am excited to get involved in this work because it’s immensely important for everyday people’s lives. The SF Bay provides millions of people with food, employment, recreation and overall well-being, and the science that NMS produces has real-world value for making policy and a concrete plan to keep the Bay healthy. It represents exactly the kind of science that I wanted to do since I first jumped into environmental biology as a 19-year-old at USC. At that time, I was interning at JPL studying historical trends in California rainfall data, so this new job represents a homecoming of sorts to California water science!
This job will be a bit of a change of pace from my present work as at first, because I’ll be part of a scientific team with a shared mission, unlike most of my prior research, where I came up with ideas, pitched them to my advisors and funders and then coordinated the projects to collect and analyze data. There will be more teamwork, and while academic publications will still be one of our products, we also will be writing reports for policymakers and stakeholders who are deciding on how to regulate nutrient levels in the Bay.
I also won’t be working with clams on an everyday basis! But as I mentioned before, clams do play a major role in the Bay in terms of filtering the water, and so it is likely we will need to understand the activities of the clams and other grazers to explain the trends in nutrients that we see. I didn’t start as a Clam Man, but my curiosity about clams meant that my attention kept being drawn to these enigmatic but influential creatures, and I expect that dynamic will continue. I am, and always will be, Dan the Clam Man.
I will continue to get my present clam projects out the door as publications, so there will be lots of clamsplaining in the future months as those get out the door. Regarding the Biosphere 2 clams, we still have four individuals of Tridacna derasa (the smooth giant clam) growing in the 700,000 gallon ocean tank, and intend to leave them as long-term research subjects and an exhibit for visitors to enjoy and learn about. We also have proposals in the work for new projects to expand on this work. I hope I can continue to visit in the coming decades and see our clams grow to be true giants, two feet in length! I also hope to acquire pet giant clams of my own, with names rather than specimen numbers, to be my friends rather than my research subjects.
I’ll be starting the new job remotely at the end of this month, to give myself time to tie off loose ends in Tucson, intending to move to the Bay Area by March. I will really miss Biosphere 2 and Tucson, but this isn’t the last they’ll see of me, because my collaborations with people here will continue into the future. I knew from the start as a postdoctoral researcher that my position would not be permanent, but it is still bittersweet to leave. I will miss hiking in the Sonoran desert, swimming in the Biosphere 2 ocean tank and also my advisor Diane Thompson and her lab here, full of people who have been a joy to work with.
But I am excited for this new chapter, because the postdoc life has been lately losing its luster for me. I’ve enjoyed being a postdoc for the freedom it entails, both in my research topics and the way I structure that work. But postdoc work is emotionally exhausting, as I have been a journeying academic contractor on “soft money”. My employment for the following year has always been contingent on the next grant coming through. Moving between different institutions on different continents has been a big weight on my family and my partner, who I miss greatly.
As a postdoc, while I’ve had fun and wouldn’t change anything about it, I have felt like a plane trying to take off in unfavorable weather. I could see the end of the runway approaching as my current funding ends in May, which was a scary feeling. I’m willing to hustle and fight for research funding, but not my basic income. Looking back, I have applied to around 45-50 academic positions (including postdocs) since finishing my PhD and got interviewed for less than ten percent of those, and received offers for two postdocs. When I got the offer from SFEI, which was itself a rigorous, multi-stage process over months, I cannot describe what a relief it was to clear out my “job applications” folder in my to-do list. This SFEI job will allow me to pursue marine science that helps the environment and people, in a more emotionally sustainable way.
I’m excited to start my next chapter and share with you all the discoveries our team makes about the SF Bay, while also continuing to clamsplain here on my own time. Keep an eye out for my Biosphere 2 studies, which will be rolling out over the next months as the data arrives!
Clamsplaining 
