Mystery of the “spurting” mussels

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

U. crassus in a Slovenian river (Alexander Mrkvicka)

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

U. crassus propped up on its foot (UCforLife)

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

Squirting water into the stream! (Vicentini 2005)

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

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

Top-down view of spurting behavior (Vicentini 2005)

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

Lifecyle of U. crassus (Rita Larje via UCforLife)

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

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

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

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

When a clam gets an offer it can’t refuse

Tridacna maxima in Eilat, Israel

I study the giant clams, bivalves which can grow over three feet long and and are willingly “infected” by a symbiotic algae which they house in an altered stomach cavity. They provide their algae partners with nitrogen, a stable environment and even funnel light in their direction, and the algae happily share the fruit of their labor in the form of sugars. Imagine yourself swallowing algae, storing it in your gut and developing windows in your flesh to let light into your stomach. You’d never have to eat again. This is the growth hack that enables the giant clams to grow to unusual sizes. But it turns out that this lovely, beautiful partnership may not have started so peacefully. The algae may have made an offer the clam couldn’t refuse.

Top left: normal mussel. Top right: heavily infected L-shaped shell opening. Bottom: view of an algae-infected mussel, including close up of pearls. From Zuykov et al. 2018

A team from University of Quebec recently discussed what such a fresh infection looks like in mussels and it ain’t pretty. The mussels basically have their shells and bodies overgrown by parasitic Coccomyxa algae, leaving its flesh bright green and transforming its shell from the classic elongated, acute angled margin typical of Mytilus mussels into a strange L-shaped overhang. The more algae are present in the mussel, the more extreme this deformity becomes. The researchers propose that this is no accident, but that as they move in, the algae also manipulates the biochemical pathway that the mussel uses to create its shell.

Mussels, like all bivalves, create their shells by laying down calcium carbonate in layers at the outer edge of the shell. The calcium is sourced from salts in the water column and the carbon primarily comes from carbonate ions also available in the water. This reaction is easier when the pH of the clam’s internal fluid is higher (less acidic), and that is exactly what the algae may assist with. Algae like all plants take in carbon dioxide to use in photosynthesis, and in doing so they increase the pH of the mussel’s body fluid,

The authors note that the region of shell which experiences abnormal thickening in the infected mussels is also the most exposed to light. The Coccomyxa algae may be causing runaway calcification of shell in the regions that they infect, and even may be directly assisting with the calcification in an additional way through the action of an enzyme called carbonic anhydrase, which is used in both their photosynthesis and in shell production (I won’t get into the nitty gritty of that reaction here). But the calcification of the mussels does appear to be in overdrive, as infected mussels were also observed to make pearls!

The algae’s photosynthesis may be assisting the mussel’s shell formation, though overall these are still quite unhealthy organisms of lower weight than their uninfected brethren. Still, Coccomyxa is known to form symbioses with lichens and mosses, so it could be that with enough generations of collaboration and a bit of evolution, the harmful algal infection could become a much more mutually beneficial partnership. It’s not so far fetched to imagine that an ancestor of today’s giant clams got a bad case of gastritis and decided to make the best of a bad situation. Making a deal with their invaders, they became greater than the sum of their parts and evolved to be the giant hyper-calcifiers we know today.

Revenge of the Clams

Lampsilis showing off its convincing fish-like lure. Photo: Chris Barnhart, Missouri State.

Clams are traditionally the victims of the aquatic realm. With some exceptions, clams are generally not predatory in nature, preferring to passively filter feed. When they are attacked, their defenses center around their protective shell, or swimming away, or just living in a place that is difficult for predators to reach. They are picked at by crabs, crushed in the jaws of fish, and pried apart by sea stars. But some clams are sick of being the victims. They have big dreams and places to be. For these clams, the rest of the tree of life is a ticket to bigger and better things. These clams have evolved to live inside of other living things.

Pocketbook mussels, for example, have a unique problem. They like to live inland along streams but their microscopic larvae would not be able to swim against the current to get upstream. The mussels have adapted a clever and evil strategy to solve this problem: they hitch a ride in the gills of fish. The mother mussel develops a lure that resembles a small fish, complete with a little fake eyespot, and invitingly wiggles it to attract the attention of a passing fish. When the foolish fish falls for the trick and bites the mussel’s lure, it explodes into a cloud of larvae which then flap up to attach to the gill tissue of the fish like little binder clips. They then encyst themselves in that tissue and feed on the fish’s blood, all the while hopefully hitching a ride further upstream, where they release and settle down to a more traditional clammy life of filter-feeding stuck in the sediment.

Very tiny Mytilus edulis living in the gills of a crab (Poulter et al, 2017)
The tiny 2.5 mm long Mimichlamys varia, living on the leg of a crab (Albano and Favero, 2011)



Clams live in the gills of all sorts of organisms. Because they broadcast spawn, any passing animal may breathe in clam larvae which find the gills a perfectly hospitable place to settle. Sure, it’s a bit cramped, but it’s safe, well oxygenated by definition and there is plenty of food available. They also may just settle on the bodies of other organisms. Most of these gill-dwelling clams are commensal: that means that their impact on the host organism is fairly neutral. They may cause some localized necrosis in the spot they’re living, but they’re mostly sucking up food particles which the host doesn’t really care about. In addition, in crabs and other arthropods, these clams will get shed off periodically when the crab molts away its exoskeleton, so they don’t build up too heavily.

Top: Kurtiella attached among the eggs of the mole crab. Bottom: aberrant Kurtiella living within the tissue of the crab (Bhaduri et al, 2018)

While being a parasite is often denigrated as taking the easy way out, it is actually quite challenging to pursue this unusual lifestyle. Parasitism has evolved a couple hundred times in 15 different phyla, but it is rare to find some organism midway in the process of becoming a true parasite. One team of researchers just published their observations of a commensal clam, Kurtiella pedroana, which may be flirting with true parasitism. These tiny clams normally live in the gill chambers of sand crabs on the Pacific coasts of the Americas. They attach their anchoring byssal threads to the insides of the chambers and live a comfortable life until the crabs molt, when they are shed away. The crabs mostly are unaffected by their presence, but the researchers noticed that some of the clams had actually burrowed into the gill tissue itself. This is an interesting development, because the clams would not be able to filter feed in such a location, so they must have been feeding on the crab’s hemocoel (internal blood). These unusual parasitic individuals are currently a “dead end” as they haven’t figured out how to get back out to reproduce, but if they ever do, they could potentially pass on this trait and become a new type of parasitic clam species. The researchers have potentially observed a rare example of an animal turning to the dark parasitic side of life, with some living in a neutral commensal way and other innovative individuals seeking a bit more out of their non-consensual relationship with their host crabs.  Considering the irritation that other bivalves suffer at the claws of pesky parasitic crabs, this seems a particularly sweet revenge.





Local thief spotted in my backyard


The brown-headed cowbird (Molothrus ater) is a notorious thief. This is merely a young thief which I observed in my yard; it’s still in training, but when it is an adult it will look more like this:

Source: Wikipedia

It will search out a nest of an unsupecting bird, perhaps a sparrow or other songbird, and it will sneakily lay an egg when the parents are away. The egg will hatch and the other birds will raise it as their own. This practice is known as brood parasitism. The unsuspecting sap that raises the young cowbird will unfortunately feed its own young proportionately less and its fitness will suffer as a result.

Sometimes, the victim figures out that one of the eggs isn’t its own and disposes of it. If the cowbird sometimes returns to the scene of the crime and if it discovers its egg is missing, it may destroy the nest in a darkly Darwinian form of payback called “mafia behavior.” Sometimes, a cowbird egg is an offer you can’t refuse.

When a clam has a stowaway

My mussel contained a tiny half-eaten crab! - Imgur
Source: jeredjeya on Reddit

Bivalves put a lot of energy into their shells. These hardened, hinged sheaths of carbonate are an effective defense against many predators looking to get at the squishy clam’s body encased inside. Parasitic pea crabs have evolved to free-ride on the bivalves’ hard work.


(video courtesy Dana Shultz)

Pea crabs are small (pea-sized), very specialized parasites which live in the mantle cavity of many bivalve groups including oysters, mussels, clams and more. The mantle is the wall encasing the soft body of the bivalve, and the cavity is the space between this soft gooey tissue and the shell itself.

For a pea crab, there is no better place to be than this tiny, claustrophobic space. In fact, they can’t live anywhere else, though some species have been found in other unusual places such as inside the anuses and respiratory tracts of sea cucumbers (link SFW, fortunately, unless you’re a sea cucumber). In a bivalve host, the crab is protected from predation by the shell, and the bivalve provides a constant buffet of food as it sucks in suspended particles with its gills. The crab steals some of this food from itself before the bivalve can digest it.

As you might imagine, having a crab living in you taking your food and pinching at your gills is not an ideal arrangement for the bivalve. Pea crabs damage their hosts’ gills with their constant picking, and bivalves infected with crabs suffer slower growth than uninfected individuals, particularly for those unlucky enough to play host to the larger female pea crabs. At a certain point, the males will sneak out of their hosts and find a bivalve with a female crab inside. At this point, they mate inside the host’s shell, adding great insult to injury. The female releases her larvae, which swim out to infect new hapless bivalves and start the cycle over again.

Aww, she’s expecting! The papers refer to this as being “with berry” which I find amusing for some reason. (photo from Dana Shultz)


You might think that commercial oysters with crab parasites would be thrown out, but to the contrary, finding a pea crab or its close relative the oyster crab with your meal is a cause for celebration in some areas, such as the Cheasapeake Bay. The crabs are eaten whole and often raw, and are said to have a texture akin to shrimp, with notes of sweetness and umami. Personally I prefer surprises in my Kinder eggs rather than in my shellfish, but to each their own.

A pea crab serves as a nice side dish for this lunching sea otter. Source: Brocken Inaglory on Wikipedia