The Mystery of the Giant Clams of the Red Sea and Indian Ocean

I have always been fascinated by scientific discoveries that are hanging right in front of our noses. Cryptic species are one such surprise. Sometimes, researchers using genetic sequencing are surprised to discover that a group of animals that all look the same from the outside are actually reproductively isolated from each other; separate twigs on the tree of life. This surprise has happened over and over in the history of natural science.

It turns out such puzzles are frequent among the giant clams. These unusual bivalves are specialists in coral reef environments, growing to large size with the help of symbiotic algae that create sugars through photosynthesis. Within the genus Tridacna there are ~10 accepted species which vary in size, shape, color and mode of life.

Tridacna squamosina (right) sitting next to the small giant clam T. maxima (left) on the Israeli Red Sea coast

I specialize in the three species known (so far) from the Red Sea, including the small giant clam Tridacna maxima and the fluted giant clam T. squamosa, which are both found worldwide, all the way from the Red Sea to down past the equator along the Great Barrier Reef. The third local species, T. squamosina is more unusual, so far being only known from the Red Sea (an endemic species). T. squamosina is an example of a cryptic species, having previously been assumed to be a local variant of T. squamosa. It looks pretty similar, with long scutes (flap-like appendages) protruding from its shell, thought to help stabilize it on the flat bottom of loose coral rubble. But unlike T. squamosa, T. squamosina lives exclusively at the top of the reef in the shallowest waters closest to the sun. It has a very angular, zig-zag pattern in its plications (the wavy shapes at the edge of the shell) and a characteristic pair of green stripes where the soft tissue meets the edges of the shell. The soft tissue is covered with warty protuberances.

Pictures of details of T. squamosina from Richter et al. 2008

It was only first described in detail in the early 2000s, when an international team of researchers figured out using genetic sequencing that it was a distinct species and named it T. costata. They noted that in their surveys all around the shores of the Red Sea, they only found 13 live specimens, making it an extremely rare and possibly endangered species. Fossil specimens on local reefs appeared to be much more common, suggesting it had a much larger population in the past. Then in 2011, another team at the Natural History Museum in Vienna discovered a shell of one had been forgotten in its collection for over 100 years. Rudolf Sturany, the researcher on the 1895 research cruise who had originally collected the clam, had called it T. squamosina.

The T. squamosina shell in the collection of the Museum of Natural History in Vienna (from Huber and Eschner, 2011)

In taxonomy (the science of naming and classifying organisms), the first team to name the species wins, so the name T. costata was synonymized (retired) in favor of the earlier name T. squamosina, which became the name of record. It must be annoying to spend so much time working to name a species and then discover you had been scooped over a century before! But such is science.

A mystery clam thought to be T. squamosina, later identified as T. elongatissima found off of Mozambique by iNaturalist user bewambay

The strange part was that there were some murmurs over the last few years that T. squamosina was not only found in the Red Sea, but also had been seen along the coast of Africa as far south as Kenya, Mozambique and Madagascar. Divers and snorkelers had taken pictures of a giant clam that did indeed look strangely like T. squamosina, with a zigzag shell opening and green stripes at the edge of its tissue. But some aspects of these individuals seemed off. In the Red Sea, T. squamosina lives freely, not embedded in the coral as these pictures showed, and the geometry of the angles of the shell seemed a bit different. It also would be difficult for T. squamosina to be connected in population from the Red Sea all the way South to Mozambique, as there are natural barriers which would prevent its planktonic larvae from riding currents to intermix between the two regions. When populations are separated by a barrier, the flow of genes between them is cut off and evolution begins to separate the populations from each other until they are separate species, a process called allopatric speciation.

A large specimen of T. elongatissima observed by iNaturalist user dawngoebbels off of Kenya

I figured that someday, researchers would collect tissue samples from these mystery clams to settle whether they were actually T. squamosina or something else. And this year, a team did just that, traveling along the coast of Mozambique, Madagascar, Kenya and other places, collecting samples of tissue to compare how all the different clams they saw were related in a family tree. They genetically sequenced these “clamples” and in the process, found that the mystery clams were a new cryptic species, which they called T. elongatissima!

Shells of T. elongatissima from the Fauvelot et al. 2020 paper
For comparison, a shell of T. squamosina collected off of Sinai, Egypt. You can see why they’re easy to mix up!

T. elongatissima closely resembles T. squamosina, and they are sister species on the bivalve family tree. It’s hard to tell them apart without training. Even a professional would probably mix some of them up if they were all placed sitting next to each other. The major differences appear to relate to shell shape, with T. elongatissima having a less symmetrical shell than T. squamosina, and a bigger opening at the rear hinge for a foot to poke through. The symmetrical shell and closing of the foot opening may represent changes that T. squamosina took on to adapt to be able to sit freely on the bottom, rather than embedding in the coral like T. elongatissima seems to prefer. If you’ve read this far, you may be thinking “Who cares? A clam’s a clam and these look practically the same. Aren’t you just splitting clams at this point?” At the end of the day, a species is a man-made concept; an organizing tool for use by us humans. Species are the characters in our reconstruction of the history of the world. What can we learn about the world by having identified this species T. elongatissima?

A giant clam family tree! Notice T. squamosina and T. elongatissima right next to each other.

The researchers behind the new paper discuss that based on statistical analyses of the genetic differences between the species, the most recent common ancestor for T. elongatissima and T. squamosina probably lived more than 1.4 million years ago! Some researchers have previously suggested that T. squamosina probably began its development as a separate species due to geographic isolation by low sea level, caused by repeated glaciations. With so much water trapped as ice on land during this period, the narrow Strait of Bab al Mandab, currently the gateway to the Red Sea, became a land barrier as sea level fell (kind of like opposite of the Bering Sea land bridge that formed allowing humans to migrate to the Americas). Ancestral clams trapped on the Northern end of this barrier were proposed to have evolved to become the rare T. squamosina.

This has occurred with a variety of species that became Red Sea endemics (meaning they are unique species that evolved in the Red Sea and are found nowhere else), including a unique crown of thorns starfish. The issue is that during this time of low sea level, the Red Sea went through periods where it was a rather unfriendly place for clams to live. All sorts of creatures went extinct in the period when the sea was repeatedly cut off, because the water became extremely salty, along with other unfriendly changes. So it’s unlikely T. squamosina would be present for us to see today if it only lived in the Red Sea throughout the entire length of time.

A map from Fauvelot et al. 2020 showing the distributions of different giant clams the researchers identified along the coasts of Africa and the Red Sea. Notice the bright red dots representing T. squamosina, only found in the Red Sea, while green dots represent T. elongatissima. Notice how the currents (arrows) seem to meet and then go offshore from Kenya. More on that in the next paragraph.

The researchers of this new paper propose that T. squamosina was more likely to have initially branched off due to the barrier of the Horn of Africa. The seas off of Kenya and Somalia harbor a meeting of southward and northward currents which then group and head offshore, away from the reefs that giant clam larvae are trying to get to. So any tiny floating planktonic clam larvae would experience a strong “headwind” preventing them from crossing that point. It would also mean that during times that the Red Sea was not a happy place to be a clam, T. squamosina may have found refuge on the coasts of places like Eritrea, Oman and possibly even as far as Pakistan. During times when sea levels rose and Red Sea conditions became friendlier, it recolonized the area.

As far as we know, the Red Sea is the only place T. squamosina is now found, but it may well be present elsewhere like Yemen or Oman. If T. squamosina was found in other regions, it would be tremendously important for its conservation. Right now, the species is thought to be extremely rare, with a very small native range. If it inhabited a broader area, that would mean more reservoirs of genetic diversity. This would reduce the odds that it will go extinct as reefs are put under stress from climate change, pollution and overharvesting. To survive as a species, it helps to not put all your eggs in one basket. If you’re only found in one small place, it increases the chances that a disaster (like climate change) will wipe you out.

The only way we will know for sure is to visit reefs in understudied places like Yemen, Oman, Pakistan, Eritrea and Somalia, to understand the richness of the giant clams present. These areas are understudied for various reasons: lack of research funding for non-Western researchers, lack of interest from the scientific community too focused on familiar places, and geopolitical situations that make it difficult to conduct research. But I hope someday to collaborate with people in these countries to better understand the giant clams present in such understudied regions of the globe. It is virtually certain that there are more species of giant clams, both alive and as fossils, waiting to be discovered.

Israel: Field Report!

So I’ve been living in Israel since the start of November after a whirlwind of defending my PhD, moving out of Santa Cruz forever and suddenly moving to another continent for a postdoc. I’ve been working on clams, taking samples, using an SEM and planning a new manuscript, but I have also been learning a lot about living in a country that is at once strangely familiar and completely foreign. I’m coming back to California tomorrow for a Holiday break, so I took an hour to reflect on what I’ve learned about this country so far. Here are some random things I’ve learned about Israel during my time here.

Israel is small


Israel is a tiny country by my Californian standards. You can take a bus along the entire length of it in less than seven hours. I am in Haifa on the farthest northern part of the Mediterranean coast, but in 2016, I lived down in Eilat for two months. Despite its small size, Israel has a variable climate depending on where you are. Up in Haifa, they have have a classic Mediterranean climate which reminds me of California in a lot of ways (think chaparral and coastal dunes, though a little more humid than I’m used to and with more thunderstorms). The Negev desert is in the South, which is intensely dry, hot and sometimes completely devoid of vegetation.

Happy naturalists!

For birders, I’ve noticed the North is dominated by hooded crows from Europe while the South is dominated by house crows, an Asian species. In general, because they’re at the nexus between Europe, Asia and Africa and the gradient between those ecoregions, Israel and the Middle East overall are very biodiverse. As a result, there is a vibrant culture of naturalists in this country who want to know about every aspect of every species. When I post something to my iNaturalist, within a few hours someone who is an expert on that taxon confirms or corrects me, without fail.

Delicious food+drink

This slideshow requires JavaScript.

Living here, I have been subsisting of a diet based largely on hummus, falafel, shawerma, tahini (try the dark kind!) and pita, with plenty of veggies thrown in. In fact, there are restaurants where they just serve you a bowl of hummus with pita and you have at it. Be prepared for the food coma. For beers, I guess Danish beer companies got a big foothold here early on because the defaults are Carlsberg, Tuborg, Heineken etc. The biggest native Israeli brewery is Gold Star, and they’re not bad! And there are a growing number of Israeli craft breweries. Overall I approve, though they sometimes experiment a bit too much and taste odd, and a lot of them don’t really seem to know what an IPA is.

Cultural diversity

Israel, as you may or may not have heard, is indeed a very complicated place. There is no doubt that the tensions are high between Israelis and Palestinians and Hezbollah and Iran, which is often in the news. But day to day on the ground, Israel is a very safe place and safer than what I’m used to in the States. The crime rate is much lower than almost everywhere in the US, and I can walk around Haifa at night without ever feeling at risk. That is more than I can say for Santa Cruz and many parts of LA.

Haifa is a special place. It is a uniquely diverse city with a significant Arab population. The student body at the University of Haifa is over 30% Arab in descent and I can walk through hearing four languages in one hallway. The main temple of the Baha’i faith is here in Haifa.

There are Israeli Jews of all sorts of backgrounds, including Ashkenazi, Sephardic, Yemeni, Ethiopian and more. There are secular Jews, Conservative Jews, Orthodox Jews and the Haredim (Ultra-Orthodox), along with myriad others I haven’t learned about yet. There also is a huge population throughout Israel of Russian Jews who came here following the collapse of the Soviet Union. Russian is spoken heavily here and is on the street signs. Like America, Israel is a hugely diverse place and I believe could be a great strength for their future growth and prosperity.

Language and cultural challenges

As a secular American, there were some parts of Israel that have proved challenging to adapt to. The biggest challenge for me by far is that Israel’s national language is Hebrew, which is a very challenging language to learn. I now know the numbers, some letters and some words, but there’s no way I’m going to be able to pick up conversational Hebrew during my time here. And as all foreigners know, not being able to read and write makes literally everything about “adulting” more difficult. I have had to learn never to assume that English is understood here. I speak slowly and use hand gestures. I do everything in person, never over the phone. Trying to do something logistical over the phone has not once worked. Seriously, don’t even entertain the notion of trying to do stuff in English over the phone here.

Instead, I recommend going to the place you need to go, ask the person for help with a dumb blank smile on your face, and people will try to help you do what you need to do, whether that is opening a bank account, getting your bus card, or signing a lease for your apartment. People here are generally very direct and no-nonsense in everyday business dealings, but they also have proven very generous and willing to help me, which is not something I can say about service in America. However, on the rare occasions when I’ve found someone who speaks English, is available, and is exactly the person who can help me with the task at hand, I feel as though I should drop to my knees and thank the God of Abraham for his mercy. Day to day life here is tough for a non-Hebrew speaker.

If there was one aspect of life in Israel which I will openly complain about, it is Shabbat (from sundown Friday to sundown Saturday). In most of the Western world, we take for granted that Saturday and Sunday are the days of rest. But here in Israel, it is Friday and Saturday, and Israel is very hardcore about their days of rest. On Shabbat, any eating establishment that wants to be Kosher has to be closed. Almost all public transport is shut down.

There are a small number of more secular, diverse cities, luckily including Haifa, where a couple buses stay running Friday night and Saturday. But on Saturday, if I realize I need groceries, my choice is to splurge on a cab or walk 25 minutes down to the nearest 24/7 market (basically a convenience store). There is a reason Shabbat is a big deal in Israel and I get it. There is no other country on Earth where Jews of all creeds and colors can know that they will get to observe Shabbat in the way it was practiced by their ancestors. But for me as a secular person without a car, it presents a lot of logistical challenges.


Here is a list of other things I found notable and unexpected about life in Israel

  • They really like malls. There are new malls opening everywhere and they are always full of people. As an American, I think of malls as very last century, but they’re still the main social place here for many people.
  • They don’t really use mops. Instead, they use giant squeegees to clean their floors. I still don’t get how to use one.
  • When you sign a lease, many landlords want twelve pre-signed checks. I thought this was very strange (where do they keep them?!) and then noticed an option in the ATM to save checks for “safe-keeping.” Weird.
  • Israel is a cell phone paradise. I can get an unlocked SIM card with 30 gb of data, unlimited voice and text for $21.50. This is absolutely insane. How is this possible?
  • In Israel, they charge tenants a bimonthly property tax. That is annoying!
  • People say Israel is super expensive and yes, prices on food and basics are somewhat high by standards of some US States. But coming from California, I have been so relieved. I now can afford to live in my own apartment and pay <25% of my income on rent rather than 50%. I can once again stay under $10-15 a day on food which wasn’t possible for me in California for the last couple years. So I have more discretionary income for fun stuff which has been refreshing.
  • As I’ve noticed in Europe as well, it’s fun paying with coins! They have 10, 5, 2, and 1 shekel coins and I find myself actually using them, unlike the useless pocket change in the states that I just save to trade in later. There are around 3.7 shekels to the dollar.
  • They have an excellent bus system (except Friday or Saturday 😉 ). Buses come by frequently in all parts of Haifa, are clean, and cost about $1.50 per ride (1/3 less than that if you set up your student access card). The bus card allows you to connect for free within a certain time period as well.
  • Most locks in Israel use keys on the inside and outside. I’m not sure if this is an Israel-specific thing, or just something the rest of the world has that the US doesn’t, but it was surprising to see that I’d have to use my key to lock my front door from inside.
  • They have smarter crosswalks in Haifa that generally bridge over a median, with two separate pedestrian lights. You may have to stop in the middle but it means less risk of someone doing a turn and hitting you, and that is good urban planning in my book. Eilat has done away with streetlights altogether, completely converting to roundabouts. I frickin love roundabouts.
  • Israel has a semi private system of healthcare, but with generally very high quality care and low cost.
  • In Israel, life is completely transformed by their mandatory military service. While I have been out of college for a few years, many Israelis are only just starting college as they enter their late 20s after being discharged. So the student body trends older at Israeli universities.
  • I thought I’d stick out as an American goy being here, but apparently I don’t. People keep asking me for directions in Hebrew and Russian and I just say “sorry, English only”, and they look at me with disappointment. I guess I can say I look Jewish!

In conclusion, I have enjoyed my time in Israel so far, and I have found myself just watching life going on around me with great curiosity, because it is a very interesting place full of constant unexpected moments. Let me know if any of you visit anytime soon 🙂