A hinged shell does not a clam make (QUIZ)

Bivalves are so named for their two hard shell valves made of carbonate, linked by a soft ligament acting as a hinge. They use a strong adductor muscle to close their shell, and the relaxation of the muscle allows the springy ligament to reopen (you might be familiar with adductor muscles as the edible tasty part of a scallop). In deference to the bivalves, laptops and flip-phones are called “clamshell” designs. That satisfying snap into place when you spring the ligamen… I mean, hinge of a flip phone is an example of human design imitating the ingenuity of evolution. But it turns out that plenty of other members of the tree of life have also stumbled upon the durable idea of a hinged two-valve shell. On the other hand, plenty of bivalves have given up on the classic clamshell look. In fact, the ancestor of all bivalves had a one-part shell, and the hinge evolved later.

Test your knowledge by trying to identify which which pictures are bivalves and which aren’t. Answers and picture sources at the bottom!

A. brachiopod-semenov.jpg

B.IMG_20150808_120740529.jpg

C.

D.

E.

ostracod_waiwhetumontage_b_650.jpg

F.

G.

naticarius_plicatella_3_01_.jpg

H.

0fbc20dc-8bf8-4719-b6fd-06050caf85e3.png

I.

tn_burrowed_jpg.jpg

J.

K.

3630165727_a312f06fbc_o.jpg

L.

kuphus.jpg

ANSWERS

A. This is not even a mollusk, never mind a clam! It’s a different benthic (bottom-dwelling) invertebrate called a brachiopod, which make up their own phylum. To put that in perspective, brachiopods are as far from bivalves on the tree of life as you are (you’re in phylum Chordata)! Yet they evolved a similar look through a process called convergent evolution. If environmental needs are the same, organisms may come to the same solution multiple times. Much like wings for flight evolved independently in insects, birds and bats, bivalves and brachiopods both evolved a hinged shell as a form of protection from predation.

B. These are indeed bivalves: rock scallops commonly found off the coast of California (picture by me of an exhibit at Monterey Bay Aquarium). So similar to the brachiopods in their ridged, hinged shells. Like picture A, these guys specialize in living on hard, rocky nearshore bottoms. Some cultures do apparently eat brachiopods (I have not), but I have little doubt that rock scallops are tastier.

C. These are also bivalves: windowpane oysters. Also known as capiz shell, they are commonly used for decoration and art due to their beautiful, thin semi-transparent shells. A large industry harvests them off the shores of the Philippines, where they unfortunately are growing scarce due to overexploitation.

D. These are not bivalves! They are crustaceans called clam shrimp. They have little legs poking out of a hard hinged shell, and have been found in some of the harshest environments on earth, where they wait in extended hibernation, sometimes years, between bouts of rainfall.

E. Not a bivalve. These are another kind of crustacean called ostracods. Like clam shrimp, ostracods live in a hinged shell and swim around with the help of tiny legs, filter-feeding in the water column. Ostracods are everywhere in the oceans and in freshwater, but have undergone an extreme process of miniaturization from their ancestral form, and are now represent some of the smallest complex multicellular life known.

F. These are fossil ostracods. You can see why they are sometimes mistaken for bivalves! The givaway is that one valve is overhanging the other. Most bivalves have symmetry between the two halves of their shell, but ostracods and brachiopods do not.

G. This is a snail, so it’s a close molluskan cousin of bivalves. Some snails feature a hinged lid at their shell opening called an operculum. This operculum can be closed to protect from predators and also seal in water to help land snails from drying out between rains.

H. This is a bizarre bivalve called a rudist. They were common during the time of the dinosaurs but went extinct during the same extinction, 66 million years ago. While they come in many bizarre shapes, this elevator form (or as I prefer to call them, trash-can form) would have been stuck in the sediment with its small lid poking out at the surface. They could open and close the lid to filter-feed.

I. This is a giant clam, Tridacna crocea! Its shell is hidden, embedded in the coral that has grown to surround it on all sides. Only the mantle (soft “lips”) are exposed, and are brilliantly colored by the symbiotic algae in its tissue. It harvests the sugars made by the algae for food. Despite being embedded in the coral, the clam does have enough room to close and pull back its mantle if a predator approaches.

J. This is a one of the weirdest modern bivalves, called a hammer oyster. These guys are found in the tropics, and the hammerhead part of their shell is actually their hinge, extended at both sides. The hinge provides the surface area needed to “snow-shoe” on top of the soft sandy bottom where they live. Other bivalves sometimes take advantage and live on the oyster like a raft!

K. This is a different brachiopod. Notice the lack of symmetry between the valves which gives it away.

L. This is by far the weirdest modern bivalve, a shipworm. These guys live buried deep within wood and are the number-one killer of wooden ships. They secrete a long tube of carbonate and have largely given up the hinged lifestyle, looking more like worms than mollusks.

Fossil-Fueled Life

mussels-hires (1).jpg
Thick aggregates of Bathymodiolus mussels on a cold seep site off of Nantucket. Source: NOAA

Humans aren’t the only users of fossil fuels. In many parts of the ocean, natural gas (methane) is constantly bubbling out of the sediment. These areas are known as cold seeps and are often a marker of productive fossil fuel reservoirs in the crust underneath. The name cold seep is somewhat of a misnomer (they are often slightly warmer than surrounding waters), but they are indeed much cooler than the more famous hydrothermal vents, which form due to geothermal activity. They are often found in shallower waters than hydrothermal vents, which generally occur in the deepest regions of the ocean where the earth’s crust is rifting. As with hydrothermal vents, cold seeps provide a unique opportunity for ecosystems to arise which are based on chemical energy, rather than solar-powered photosynthesis like the rest of the biosphere. Unusual life forms harness the chemical energy of the methane and sulfide emitted at these seeps, and many can also be found at the more famous hydrothermal vents.

redcrab-600.jpg
Crabs feeding on a bed of Bathymodiolus. Source: NOAA

Bathymodiolus mussels, vesicomyid clams, and other bivalves thrive at these seep environments, and have evolved to partner with bacteria in their gills and stomachs which can directly consume and produce energy from the reaction of methane and sulfate continuously being sourced from the seep underneath, creating bicarbonate and sulfide as products. This form of metabolism, chemosynthesis, is distinct from the familiar photosynthesis/respiration that most life-forms at the surface use to create energy. The partnership between a multicellular animal and chemosynthetic bacteria is called chemosymbiosis. These reactions are of course being utilized by all sorts of microbes not partnering with bivalves, but the bacteria that take on metazoan hosts have the advantage of a stable environment and a constant flow of fresh seep gas brought in by the bivalves’ gills. The bivalves feed on the products of the microbe’s hard work. As they are some of the only inhabitants that can tolerate the oxygen-free, toxic environment of seeps, they form dense thick shell beds wherever a seep appears.

p0379hz2.jpg
A bed of tubeworms and a deep-living octopus in the Arctic. Source: MBARI via BBC

Other chemosymbiotic groups use the hydrogen sulfide byproducts from seeps, which they oxidize into elemental sulfur. The most prominent of these are the vestimentiferan tubeworms, which lack mouth or anus and are totally dependent on the activity of the bacteria that they house in a modified digestive tract. Some of these worms have been found in seeps in extreme environments, such as the truly cold cold seeps of the Arctic. They are less spectacular in appearance than their bright red counterparts from hydrothermal vents, but are believed to live for an extremely long time, depending on the consistency of the seep that they inhabit.

 

Hesiocaeca_methanicola_noaa.jpg
Hesiocaeca methanicola, one species of the poorly understood “ice worms.” Source: NOAA via Wikipedia

At the extreme pressures and low temperatures of the deep ocean, methane can freeze in combination with water molecules, forming structures called clathrates. Much of the deep ocean floor is dusted with deposits of clathrates. Microbes which feed on clathrates are believed to be a food source for grazing polychaete “ice worms.” These unusual organisms can survive up to 96 hours without a whiff of oxygen, an unbelievable feat for a moving, multicellular organism.

Imaginative artist representation of a probe visiting a vent under the ice of Europa. Source

Cold seep environments are perhaps merely one specific manifestation of a vast, poorly understood collection of biota which do not depend on the sun for their energy. We do not yet understand many of the deep ecosystems which may be present within the earth’s crust, in the deep ocean and trapped under polar ice. NASA studies cold seep and hydrothermal environments as the best analog for the conditions life could experience in the frozen oceans of Europa and Enceladus. It is eerie to think that such alien ecosystems  exist merely a few kilometers off of our familiar shores, and were using fossil fuel energy far before humans figured out how to combust it.

NOAA Ocean Explorer: Okeanos Explorer: Northeast U.S. Canyons 20
Methane bubbling out of the seafloor off of the Virginia coast. Source: NOAA

 

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.

IMG_2161
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.

Sea_otter_with_shells_2.jpg
A pea crab serves as a nice side dish for this lunching sea otter. Source: Brocken Inaglory on Wikipedia

 

The Many Homes of Hermit Crabs

IMGP3600
My boy Harry, a purple pincher (Coenobita clypeatus) inhabiting a tapestry turban snail (Turbo petholatus) shell. These seem to be his favorite kind, even though they do not come from his native Caribbean.

Hermit crabs (superfamily Paguroidea) are most famous for using snail shells as their home, having evolved a soft, spiral abdomen to be able to use them for protection. But they are more flexible about their choice of abode than you might expect.

tumblr_o85rmrh6Yw1sq04bjo1_1280.jpg
This crab was likely preserved buried alive in sediment. Note how it uses its claw as a protective trap door sealing the opening of the ammonite shell. Source: Jagt et al, 2006

Different groups of shelled organisms have risen and fallen in abundance through geological time. During the time of the dinosaurs, ammonites (relatives of modern squid and octopus) were among the most common marine organisms, and hermit crabs were there to recycle their shells when they died.

Capture
Each tiny pore (zooid) in this bryozoan contained a tiny tentacled organism. Together they grew in a shape that made for a nice hermit crab house (image 5 shows a cross section where the crab’s abdomen would fit). Source: Taylor and Schindler 2004

Mollusks aren’t the only contractors for hermit crabs. Some hermits utilize the skeletons of colonial organisms like bryozoans as a home. Bryozoans are filter-feeding colonial animals made up of thousands of tiny tentacled organisms living in the pores of a shared skeleton. The extinct bryozoan Hippoporidra lived in symbiotic partnership with hermit crabs, growing around a gastropod shell to attract a hermit crab partner. This was an example of mutualism: by providing a home for a crab, the bryozoan would be transported to new environments with plentiful food particles to eat, and also would be protected from their arch-enemy, nudibranchs (sea slugs). Some modern day hermits, such as Manucomplanus varians of the Gulf of California, have evolved very similar partnerships with live staghorn corals.

IMG_20160410_202354.jpg
Manucomplanus varians at Monterey Bay Aquarium

Not all hermit crabs live in hard houses. Some deep sea forms partner with anemones, with the stinging tentacles serving as an effective defense.

Source: Okeanos Explorer

The recently discovered green-eyed hermit crab, which also lives in deep water, lives in a glued-together mass of sand created by tiny anemones, which continue to grow the structure to fit the crab as it increases in size.

145304_web.jpg
The green-eyed hermit crab was found over 200 m deep off of South Africa. Source: Lannes Landschoff via Eurekalert 

Unfortunately, hermits adapted for gastropod shells are unable to find adequate homes in some areas, due to overharvesting of shells for the tourist trade as well as an excess of plastic trash. These crabs make do with whatever items that they can find. Plastic is not an ideal home material for hermits. Bottlecaps and narrow tubes do not allow the crab to fully retract for protection and leach chemicals which may harm the crab. The crabs also nibble on their shells as a source of calcium, which is obviously not possible with plastic.

image.jpg
Coenobita purpureus, a land hermit crab on Okinawa. Source: Shawn Miller

But hermits continue to impress me with their flexibility and ingenuity in their search for homes. For a hermit crab, home is where the abdomen is.

 

 

Why do worms surface after a rain?

6918874796_10bee09b82_o.jpg
Source: Eva the Weaver on Flickr

Biologists have long wondered at the capabilities of earthworms. When he wasn’t crafting foundational theories of modern biology, Charles Darwin discovered that earthworms assist plant growth by aerating and turning over soil. Previously derided as pests, his work helped to shed light on the dark, dirty, essential role of these silent tillers of the earth. Earthworms spend most of their lives converting organic matter into compost. They play an important role in nutrient cycling, as the castings that they leave behind are rich in important nutrients like phosphorus and calcium.

Our sunny, dry surface world is not safe for these subterranean beasts. Earthworms are not at home in daylight and are usually only there by necessity. One of the most common times to observe them is during and after a rainstorm, particularly at night. As a child, I often wondered why earthworms seemed so determined to drown themselves in puddles. Researchers have still not reached consensus on the causes for this behavior. Darwin supposed that the sicker worms were driven to the surface by flooded soil. Some propose that they are driven from the saturated soil by low oxygen content, where many drown in puddles as they attempt to find non-waterlogged soil. Other researchers are skeptical of this angle, as worms require moisture to breathe through their skin, and many species can survive immersion in water for extended periods. They suggest that the worms are using the wet conditions brought by rains to aid migration to new patches of soil, or even to mate. The reality, as often is the case, is likely something of a combination of all of the above.

One team found that different earthworm species have different tolerance levels for low oxygen conditions. The worms that can survive extended periods of immersion in water are the same species that often remains under the ground following a rainstorm. These worms have lower oxygen consumption. The worms with higher rates of oxygen consumption leave the ground when dissolved oxygen levels are too low to sustain respiration. They do so preferentially at night, partially because this is a period when they are more active, and thus respiring more oxygen, and also because at night they are less exposed to predation by birds. The early bird (at least before sunrise) really does tend to get the worm.

Some have suggested that the rhythm of raindrops stimulates the emergence of earthworms, and that the same mechanism allows fishermen (and animals) to use “worm fiddling” or charming to gather thousands of worms at a time for bait. Worm fiddling can take many forms, but one of the most common techniques observed involves rubbing a long piece of metal against a post driven into the ground. The vibrations induce thousands of worms to rise up out of their burrows.

Ken Catania, a biologist at Vanderbilt University, rejected the rain rhythm idea and proposed that worm fiddling actually works by imitating the noise of a burrowing mole, one of the most ravenous predators of worms. Rather than fleeing from the simulated rain, the worms are fleeing the jaws of their greatest enemy, only to be trapped at the surface by a different adversary. Like a whale driven to beach itself by noise pollution or the threat of a predator, surfaced earthworms deserve our sympathy, particularly if they’re destined to become our fishing bait.