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It has taken me months to gather all the photos and videos I needed for this post. I could blame it on the stress of teaching online for the first time, the COVID-19 pandemic itself, or residual malaise from the dumpster fire that was 2020. But really, it's the animal's fault.

In this case the animal is the orange cup coral, Balanophyllia elegans. I've written about this beast before, and lately I've been paying more attention to the corals that we have in the lab. In many ways it is the typical anthozoan—its life cycle consists of only a polyp stage (i.e., no medusa), it is benthic, and its body is vaguely anemone-like. Like the reef-building corals of the tropics, Balanophyllia is a scleractinian coral. This means that it secretes a calcareous base, or exoskeleton, upon which sits the living tissue of the polyp. But unlike the reef-building corals of the tropics, Balanophyllia is solitary, which means that it does not clone or form colonies. Nor does it contain photosynthetic zooxanthellae in its cells, as the reef-builders do. This means that Balanophyllia is a strict carnivore: unable to rely on photosynthetic symbionts to do the hard work of fixing carbon, Balanophyllia has to catch its own prey.

Balanophyllia has separate sexes and reproduces sexually. Males release sperm that are ingested by the female, and she then broods larvae for several months. What is strange about this is that, when you consider the anatomy of Balanophyllia (or any cnidarian polyp, for that matter) the question that comes to mind is, "Where are the larvae brooded?" So let's take a look at the basic cnidarian body plan.

Adult cup coral (Balanophyllia elegans)
© Allison J. Gong

The professor who taught my undergraduate course in invertebrate zoology described a cnidarian's body as a baggie inside a baggie, with a layer of jelly between the two baggies. The inner and outer baggies represent the tissue layers that developmental biologists refer to as endoderm and ectoderm, respectively. The jelly between the two tissue layers is called mesoglea. For the sake of this discussion, it's the endoderm that interests us. In a cnidarian, the endoderm is also called gastrodermis, because it is, literally, the skin of the digestive system. The cnidarian gut is a cavity that opens to the outside via a single opening. We could call that opening either a mouth or an anus, since both food and wastes pass through it, but for politeness' sake we call it a mouth. The gut itself does double duty as both the digestive system and the circulatory system, so its formal name is gastrovascular cavity (GVC).

It's easy to imagine the GVC as being essentially a tubular vase, but it's more complex than that. The GVC extends into each of the tentacles, so the tentacles are actually hollow. The main cavity of the GVC, in the stalk of the polyp, is partitioned by thin layers of endoderm. The partitions are called septa, and serve to increase the surface area of the endoderm for digestion. Think of a large office building, divided into small cubicles by movable partial walls—there's much more total wall space for hanging things such as calendars than there would be in the big room with only four walls. In anthozoans, gonad tissue is associated with the septa.

Now back to how Balanophyllia broods its larvae. As we've seen before, Balanophyllia's planula larva is an orange-reddish wormlike blob about 2 mm long, which is ciliated all over. It doesn't feed, but survives on energy reserves supplied by the mother in the egg; it may also be able to take up dissolved organic material directly from the seawater. After brooding larvae for some period of time in their GVC, Balanophyllia females release larvae. Or rather, the larvae ooze out of their mother's mouth and crawl around to settle and metamorphose nearby.

The planulation season, when larvae show up amongst the corals we keep in the lab, begins in the fall and runs through winter into the spring. Sometimes the larvae metamorphose right away—one day there are no larvae and the next day there are baby corals. Other larvae squirm around for days or weeks, not getting any smaller but not metamorphosing, either. This past season (Fall 2020-Spring 2021) we saw both extremes: planulae that settled and metamorphosed right away, and others that I collected weeks ago and are still worming around.

Orange cup coral (Balanophyllia elegans) with a planula larva
2020-12-12
© Allison J. Gong

I peered into a bunch of corals, hoping to see what the larvae are doing inside the GVC, with no luck. Then I decided to try some time-lapse video under the dissecting scope, and had a bit of success with that. In the video below you'll see about two hours of action compressed into about 1 minute. Watch for a small red ball squirming around inside the GVC. And remember that the GVC extends into the tentacles, so the larva can wiggle its way in there, too.

All of which makes me wonder if the pathway from mother's GVC to the scary world outside travels in only one direction, or if the larvae can retreat back into the safety of the mom's digestive system. How strange is it, that the safer location might be inside the mother's gut!

Eventually, whether it be a matter of hours or weeks, the planula larva settles (i.e., sticks to a surface and stops crawling) and metamorphoses (i.e., makes the anatomical and physiological changes into the juvenile body). There doesn't seem to be a clear connection between surface characteristics and whether or not larvae will settle there. You might think that they'd choose to settle nearby or maybe actually on the parent, but that's not always the case. At some point, however, the larva will have to either settle and metamorphose, or die. When they metamorphose into juveniles, they reveal other aspects of their nature as scleractinian corals.

From what I've seen, the larva begins the settlement process by sticking to a chosen spot and becoming a more circular (i.e., less elongated) blob. On the other hand, I've seen perfectly round red blobs that look for all the world as though they might be pushing out tentacles, change their minds and go vermiform again. But once they stick permanently and begin to metamorphose by forming the polyp's body, they have to continue.

Newly settled orange cup coral (Balanophyllia elegans)
2021-01-29
© Allison J. Gong

The juvenile in the photo above has not yet pushed out any tentacles, but it does show signs of its scleractinian ancestry. The Scleractinia (stony corals) and Actiniaria (sea anemones) are both members of a group called the Hexacorallia. The Hexacorallia and Octocorallia are in turn grouped together as members of the class Anthozoa. As you might infer from the name 'Hexacorallia', animals in this group of a body symmetry based on the number six. And you can see in the photo above that the juvenile coral's body is divided into 12 wedges. When the juvenile begins growing tentacles and a more polyp-like body, the hexamerous symmetry becomes even more evident:

Young orange cup coral (Balanophyllia elegans)
2021-02-22
© Allison J. Gong

The young coral begins by forming six primary tentacles, which establishes the visible hexamerous symmetry. Then it grows the six shorter secondary tentacles. The bumps on the tentacles are cnidocyte batteries, clusters of stinging cells, indicating that the animal can already capture and kill prey. Good thing, too, because if it hasn't already then it soon will deplete the energy stores its mother partitioned into its egg.

By this stage in its development the coral will remain where it is for the rest of its life. As it grows it will deposit CaCO3 and grow taller, but the living tissue will always be restricted to a layer sitting on top of the calcareous base. Once the base grows more than a few millimeters tall, there is no living tissue in contact with the surface. Thus there is no mechanism for the animal to move to another location, or even to re-attach itself if it gets broken off its surface. There are many animals that are likewise unable to move once their larvae have attached and metamorphosed. The decision of where to put down roots becomes quite stark for them, as a bad one can result in a very short life indeed. The selective pressures that enforce a good decision are quite clear, and are important factors in the distribution patterns we observe in the intertidal.

1

For several weeks now I've been raising another batch of bat star (Patiria miniata) larvae, from a fortuitous spawning that occurred in early January. Since this is rather old hat by now I'm not diligently taking photos or drawing the larvae as often as I would have years ago when this kind of undertaking was new to me. But I still change the water twice a week and look at them on Fridays, and I still have the set-up that attaches my old phone to the microscope so I can take pictures of them.

Last Friday it occurred to me that: (A) my gizmo holds the camera steady over the microscope, so I can take pictures at multiple focal planes within objects under the scope; and (B) I have software that will stitch those many snapshots into a single image. Neat!

So I made this:

Bipinnaria larva of the sea star Patiria miniata, age 44 days
44-day-old larva of the bat star, Patiria miniata
2021-02-19
© Allison J. Gong

This larval stage is called a bipinnaria or a brachiolaria. From top (anterior end) to bottom (posterior end) the larva is about 1 mm long. It swims with the anterior end in front. In some sea stars the bipinnaria grows long arms, at which point we call it a brachiolaria ('brachio' = 'arm' in Greek). Bat stars don't grow long arms, so the distinction between bipinnaria and brachiolaria is much fuzzier.

I took 11 photos of this larva, each one focused on a different horizontal plane, and did a focus merge in my photo processing software. Crossed my fingers as the software did its magic, and then peeked at the result. It worked! When looking through the microscope I have to focus up and down through the body to get an idea of its three-dimensional structure. But if the animal holds still long enough, I can do the focus merge thing and get images like this one.

And that slight halo that you see around the exterior surfaces of the larva? That is not an artifact of the photo taking or processing. That halo is due to the cilia that cover the body. There is a ciliated band, which you can see as the dark gold ribbon that snakes along the lobes of the body, and the other body surfaces are ciliated as well. The ciliated band is what the larva uses to swim through the water. Each photo freezes the ciliary action at the moment it was shot, but stitching several photos together causes the cilia to blur into that pale halo.

Nifty!

Intact shells are a limited resource in the rocky intertidal. Snails, of course, build and live in their shells for the duration of their lives. A snail's body is attached to its shell, so until it dies it is the sole proprietor of the shell. Once the snail dies, though, its shell goes on the market to whoever manages to claim it. Empty shells tend not to remain on the market for long.

Hermit crabs also live inside snail shells. They are the ones that compete for empty shells when they do become available. Here in California, at least, the hermit crabs can't kill snails for their shells; they have to wait for a snail to die. And once a shell comes on the market, it will have a taker even if it's not the ideal size for the crab. It's not at all uncommon to see hermit crabs that can fit only their abdomen into the shell, leaving the head and legs exposed and vulnerable. On the other end of the spectrum, many hermit crabs are so small that they can pull into the shell and not be seen by an inquisitive tidepool visitor. Anybody taking a snail shell home as a souvenir—where such takes are allowed, of course—must be certain that there is no tiny hermit crab hiding deep in the depths.

Hermit crab in black turban snail shell
Hermit crab (Pagurus samuelis) in shell of turban snail (Tegula funebralis) at Point Piños
2015-05-09
© Allison J. Gong

From a hermit crab's perspective, the best shell is one that is big enough to retreat into but light enough to be carried around. Snail shells come in a variety of shapes and corresponding internal volumes. Turban snails, with their roughly spherical shape, have a large interior space and are coveted by larger hermit crabs. For example, the grainy hand hermit crab (Pagurus granosimanus) seems to really like both black and brown turban snail shells.

Original inhabitant and builder of the shell:

Brown turban snail partially withdrawn into shell
Brown turban snail (Tegula brunnea) at Pistachio Beach
2021-02-09
© Allison J. Gong

And opportunistic second inhabitant of the same type of shell:

Grainy hand hermit crab in turban snail shell
Grainy hand hermit crab (Pagurus granosimanus) in brown turban snail (Tegula funebralis) shell
2018-06-01
© Allison J. Gong

Other snails are not even remotely spherical. Olivella biplicata, for example, is shaped like the pit of an olive. Unlike Tegula, of which both intertidal species are found in rocky areas, O. biplicata burrows in sand. Note the shape and habitat of this olive snail:

Olive snail
Olive snail (Olivella biplicata) burrowing through sand at Whaler's Cove
2019-11-24
© Allison J. Gong

These olive snails have a smaller internal volume, and thus tend to house smaller hermit crabs. Young individuals of P. granosimanus can be found in olive snail shells, but they quickly outgrow the cramped quarters and need to find a larger home. Smaller hermits such as Pagurus hirsutiusculus, though, are often found in olive shells.

Any hermit crab that finds itself robbed of its snail shell has a short life expectancy. The front end of the hermit resembles the front end of any crab, with the familiar armored legs, claws, eyestalks, and antennae. But the abdomen is soft and unarmored, covered by only a thin cuticle. The abdomen is coiled to follow the coiling of the snail shell, which allows the crab's body to curl around the columella, the central axis around which the shell spirals. In this way the crab can hang onto its snail shell and resist a tug by a would-be predator. A strong enough tug, though, will rip the crab's front end (head + thorax) away from its abdomen. So if you ever find yourself with a hermit crab in hand, do not be tempted to remove it from its shell by yanking it out!

The next time you encounter gastropod shells in the tidepools and want to know whether the inhabitant is a snail or a hermit crab, watch to see how it moves. Hermit crabs scuttle, as crabs do, while snails glide along very slowly. You would also notice a difference as you pick up the shell: snails stick to the rock with their foot, which you will feel as a suction. Hermit crabs don't stick at all, so if the shell comes away easily it likely houses a crab instead of a snail. See? Easy peasy lemon squeezy!

Sometimes even a well-known site can present a surprise. Here's an example. Yesterday I went up to Davenport to scope things out and see how the algae were doing. This is the time of year that they start growing back after the winter senescence. I also took my nature journal along, hoping to find a spot to sit and draw for a while.

The first thing I noticed was the amount of sand on the beach. Strong winter storms usually carve sand off the beaches, making them steeper. And during the calmer months of summer the beaches are flatter and less steep. Yesterday the beach was very thick and flat. It makes trudging across the sand in hip boots much easier!

The accumulation of sand meant that I could walk around the first point. Unless the tide is extremely low, such as we see around the solstices, the water is too deep for that. But yesterday I walked around it, and it wasn't until I got to the other side that it occurred to me that: (1) hey, I walked around the point; and (2) I could do that only because there was so much sand. See, a thick beach with a lot of sand makes a mediocre low tide feel lower because the water isn't as deep as it would be if the beach were thinner. When the tide isn't low enough for me to walk around the point, I have to clamber down a cliff. The cliff height varies depending on how much sand has built up, obviously, but is about head height for me. Getting down usually involves scooting on my butt and hoping my feet land on something that isn't slippery. As with most climbing, up is easier and less scary than down.

It's hard to imagine the amount of sand there was yesterday. Look at this picture.

Flat rock area and sandy area
North of Davenport Landing Beach
2021-02-08
© Allison J. Gong

See how the rocks in the foreground end? Usually that's the edge of the cliff. Yesterday I could have just taken a tiny step off the top of the cliff onto sand. That's over 1.5 meters of sand in that one spot! If the couple in the background were visiting this area for the first time, they'd have no idea of the conditions that made it so easy for them to get out onto the reef.

There was a lot of sand in the channels between rocks, too.

Sand between rocks in the intertidal
Intertidal area north of Davenport Landing Beach
2021-02-08
© Allison J. Gong

Normally those channels are deeper. You can see that some anemones were able to reach to the surface of the sand, but many more are buried, along with any other critters and algae unfortunate enough to be attached to the lower vertical surfaces. And while some of them will either suffocate or be scoured off as the sand washes away, many will survive and be ready to get on with life.

The second surprise of the day was a bright orange object. What I could see of it was about as big as my thumb, and at first I thought it was a nudibranch. Then when I crept closer for a better look, what popped into my head was "snailfish". Which was an odd thing, because I'd never seen a snailfish before. But something about the creature's posture looked somehow familiar.

Orange fish with large head and tail wrapped around the body
Tidepool snailfish (Liparis florae) at Davenport Landing
2021-02-08
© Allison J. Gong

Fortunately I had the presence of mind to take photos before trying to draw this little fish, because this is all I had time to get:

When I spooked the critter it took off really fast, confirming that it was no nudibranch. It was, indeed, a snailfish! It came to rest in a small hole in a rock, from where it looked out at me.

Tidepool snailfish (Liparis florae) at Davenport Landing
2021-02-08
© Allison J. Gong

The snailfishes are a very poorly studied group. As a group they are related to the sculpins. There are snailfishes throughout the northern temperate and polar regions, from the intertidal to the deep sea. iNaturalist shows 43 observations of L. florae, eight of which are in California. Before yesterday, none had been recorded at Davenport Landing.

Map of northeast Pacific coast, showing sighting of tidepool snailfish recoreded in iNaturalist
Observations of tidepool snailfish (Liparis florae) recorded in iNaturalist
2021-02-09
© iNaturalist

So there you have it, a snailfish! We don't know much about any of the snailfish species, even the intertidal ones. They apparently have pelvic fins modified to from a sucker, similar to the clingfishes, but I didn't have a chance to examine this specimen closely enough to confirm that. I don't know why they are called snailfishes, either. They're not snail-shaped at all.

Now, about that thing up there where I said "snailfish" came to mind even though I'd never seen one before. That happens quite a bit—a name will jump into my head before I've had a chance to think about it. Sometimes I'm wrong, but often I'm right. I know I hadn't seen a live snailfish before, but obviously I'd seen photos of them or I wouldn't have been able to recognize this orange creature as being one. It's fascinating how the brain forms search images, isn't it?

According to my notes at the lab, the last time I spawned urchins was December of 2016, making it four years ago. It has always been something I enjoyed doing, but I didn't have a reason to until now.

When the coronavirus pandemic began almost a year ago now, access to all facilities at the marine lab was restricted to a group of people deemed essential. In my case, "essential" had to do with the fact that I keep animals alive. There were many hoops to jump through and inane questions to answer—for example, "What will happen if you don't go in to check on water and food?" and "How many animals will die if you do not have access to the lab, and how much effort [i.e., $$$] would it take to replace them?"—but in the end someone higher up in the food chain exercised some common sense and decided to let me have continuous access to the lab. So I've been at the lab pretty much every day, to check on things and make sure that air and water are flowing.

So over the summer we were running sort of bare-bones operations at the lab. There were many fewer people looking after everyday things. The autoclave broke and wasn't fixed until September. One of the casualties of this less-than-normal vigilance was one of the cultures in the phytoplankton lab. Our Rhodomonas flasks had been contaminated since late 2019, and we were struggling to rescue them. I tried so hard to keep them going ahead of the contamination, but ultimately failed. As of this writing all of the old Rhodomonas cultures have died.

In October, after the autoclave had been repaired, I decided to take action and replace our inevitably doomed Rhodomonas cultures. I found a company that sells small aliquots of many marine microalgae and ordered a strain of Rhodomonas that was isolated in Pacific Grove. May as well see if a local strain of algae works as a food for local larvae, right? The new Rhodomonas cultures seem to be growing well and it's time to see of urchin larvae will eat and thrive on it.

Equipment and glassware used to spawn sea urchins

About a month ago I collected 10 urchins to spawn. Yesterday was their lucky day! Purple sea urchins (Strongylocentrotus purpuratus) are broadcast spawners, and spawning is both inducible and synchronous. We can take advantage of the inducibility to make them spawn when we want, as long as they have ripe gonads. The difficulty is that we can't tell by looking whether or not an urchin is gravid, so all we can do is try to induce them and then hope for the best.

As I've written before, we induce spawning in sea urchins by injecting them with a solution of potassium chloride (KCl). KCl is a salt solution that causes an urchin's gonopores to open and release gametes if the gonads are ripe. I shot up 10 urchins yesterday, and eight of them spawned. An 80% spawning rate isn't bad, but only two of the eight were female and neither of them had a lot of eggs to give.

Since the gonopores are located on the aboral (top) of the urchin, the easiest way to collect eggs is to invert the animal on a beaker of seawater, like so:

Female sea urchin (Strongylocentrotus purpuratus) spawning
2021-01-12
© Allison J. Gong

In nature the eggs, which are a pale orange color, would be whisked away by currents to be (hopefully) fertilized in the water column. In the lab we can collect the eggs in the beaker, as follows:

This is much less damaging to the animal than trying to pipet eggs off the top of the urchin.

We try to collect sperm and keep it dry, so there is no putting males upside-down on beakers of water. Instead we pipet up the sperm and keep it dry in dishes on ice. When it's time to fertilize the eggs we dilute the sperm with filtered seawater and add a small amount to the eggs.

One of my favorite things ever is watching fertilization take place in real time, under the microscope. It truly is one of nature's most amazing phenomena. It is a great thrill to watch the creation of new beings.

In the video you see eggs being bombarded with sperm, probably at much higher concentrations than they would encounter in the wild. It is common knowledge that it takes only one sperm to fertilize an egg, but what would happen if two sperm penetrated an egg at the same time? I've written about polyspermy and the fast and slow blocks thereto, in case you'd like to refresh your memory about what is happening in the video.

A successfully fertilized egg is easily recognized by its fertilization envelope, which is the slow block to polyspermy.

Zygotes of the purple sea urchin (Strongylocentrotus purpuratus)
2021-01-12
© Allison J. Gong

After fertilization, the next step to watch for is the first cleavage division, which occurs about two hours later.

2-cell embryos of the purple sea urchin (Strongylocentrotus purpuratus)
2021-01-12
© Allison J. Gong

Aren't they pretty?

Over the next day or so the cleavage divisions continue, resulting in the stage that hatches out of the fertilization envelope. This stage is a blastula, which is a hollow ball of ciliated cells. The hollow space inside is called the blastocoel, and it is here that the larval gut will soon develop.

Blastula of the purple sea urchin (Strongylocentrotus purpuratus)
2021-01-12
© Allison J. Gong

It's easier to see the 3-dimensional structure of the blastula by watching it spin around.

As the blastula rotates under the coverslip, you can see the ciliary currents that would propel it through the water. You also see some objects that look like sperm and are, in fact, dead sperm, getting caught up in the currents.

The blastula is the same size as the egg. The embryo can't begin to grow until it eats, which won't happen until it has a gut. Over the next few days an invagination will begin at a certain location on the blastula which is called the blastopore; this invagination will eventually form the first larval gut. At that point I will have to start feeding them and calling them larvae.

And just to remind you of our humble beginnings, we begin life in much the same way as sea urchins. That blastopore, or initial opening to the larval gut, is the anus. The mouth doesn't exist until the invagination breaks through to the opposite end of the embryo. So yes, like the sea urchin, you had an anus before you had a mouth!

4

On the penultimate day of 2020 I met up with my goddaughter, Katherine, and her family up at Pigeon Point to have two adventures. The first one was to find a marble that had been hidden a part of a game. We got skunked on that one, although the marble was found after we left and the hider had sent an additional clue. The second adventure was an excursion to the tidepools. I've had a lackadaisical attitude towards the afternoon low tides this winter, not feeling enthusiastic about heading out with all of the people and the wind and having to fight darkness. But the invitation to join the marble hunt, on a day with a decent low tide, meant that I could spend a good deal of quality time with Katherine.

It is not unusual for a promising low tide to be cancelled out by a big swell. It happens, especially during winter's combination of afternoon lows and occasional storms. The swell yesterday was pretty big.

Here's the view to the north, from Pigeon Point:

Looking north from Pigeon Point
View to the north from Pigeon Point
2020-12-30
©Allison J. Gong

All that whitewash breaking over the rocks is not good for tidepooling, especially with small kids in tow.

This is how things looked to the south of the point:

View to the south from Pigeon Point
2020-12-30
©Allison J. Gong

This is Whaler's Cove, a sandy beach that lies on the leeward side of the point itself. See how the water is much calmer? It's amazing how different the two sides of the point are, in terms of hydrography, wind, and biota. The south side is much easier to get to, especially for newbies or people who are less steady on their feet. Being sheltered from the brunt of the prevailing southbound current means that the biological diversity is, shall we say, a bit subdued when compared to what we see on the north side of the point.

I first took Katherine tidepooling when her sister, Lizzie, was an infant riding in her mom's backpack. Katherine was about four at the time. Her mom and I were suprised at how much she remembered. She recognized the anemones right away, even the closed up cloning anemones (Anthopleura elegantissima) on the high rocks. She remembered to avoid stepping on them—that's my girl!

She wasn't all that keen on touching the anemones, though, even after we told her it feels like touching tape.

Giant green anemone in tidepool
Giant green anemone (Anthopleura xanthogrammica)
2020-12-30
©Allison J. Gong

She did like the sea stars, too. Purple is my favorite color and I think hers, too, so the purple and orange ochre stars were a hit. It was nice to see two large healthy ones.

I had some actual collecting to do, so it was a work trip for me. Late December is not the best time to collect algae, but I wanted to bring some edible seaweeds back to the lab to feed animals. We haven't had any kelp brought in since the late summer, and urchins are very hungry. They will eat intertidal seaweeds, though, and when I go out to the tidepools I bring back what I can. It will be a couple of months until we see the algae growing towards their summer lushness, but even a few handfuls of sea lettuce will be welcome to hungry mouths.

Bright green sea lettuce growing with red algae
Sea lettuce (Ulva sp.)
2020-12-30
©Allison J. Gong

Katherine and I walked up the beach for a little way to study one of the several large-ish crab corpses on the sand. This one was a molt rather than an actual corpse.

Rock crab molt on sand
Rock crab (Romaleon antennarium) molt
2020-12-30
©Allison J. Gong

Katherine found the missing leg a little way off, and we discussed why we call these limbs legs instead of arms. "They use their claws to pinch things, like hands," she said. Not wanting to get into a discussion of serial homology and crustacean evolution with a 6-year-old, I told her that calling the claws "hands" isn't a bad idea, since they are used a lot like the way we use our hands. But, I continued, the crab walks on its other limbs like we walk with our legs, so can we call those legs? She was happy to agree with that. I can tell I will have to be careful about how I explain things to her, so that she doesn't come up with some wonky ideas about how evolution works.

In the meantime, Lizzie, the little sister, was having a grand old time. She flooded her little boots without a complaint and, after her mom emptied the water from them, squelched happily along with soggy socks. That girl may very well grow up to be a marine biologist!

Once the sun went behind the cliff it started getting cold. With one child already wet we decided to head back. On our way up the beach we saw this thing, which I pointed out to Katherine:

"What is it?" she asked. When I asked what she thought it was she cocked her head to one said and said, "It looks like a rock." Then I told her to touch it, which she didn't want to do. So I picked it up and turned it over, to show her the underside:

Gumboot chiton (Cryptochiton stelleri)
2020-12-30
©Allison J. Gong

These big gumboot chitons do look more interesting from this side, because you can at least see that they are probably some kind of animal. Katherine had seen some smaller chitons on the rocks, so she had some idea of what a chiton is, but these are so big that they don't look anything like the ones we showed her earlier. Plus, with their shell plates being covered with a tough piece of skin and invisible, there are no outward signs that this bizarre thing is indeed a chiton. Katherine was not impressed.

At this time of year, when the sun decides to go down it goes down fast. But as we were walking back across the rocks the tide was at its lowest, so there was more terrain to explore. Then it was back up the stairs to the cars, where we could get warm and dry.

Beach and lighthouse at Pigeon Point

Oh, and Katherine and her mom and sister were able to find the hidden marble! They also hid one of their own for someone else to find.

At the end of August I got to play animal wrangler for a film production. Back in the late winter I had been contacted by an intern at KQED in San Francisco, who wanted to shoot some time-lapse footage of anemones dividing. We went out and collected anemones, I got them set up in tanks at the marine lab, and then COVID19 hit and everything went on shut-down. The intern finished her internship remotely and went on to her next position, and in the meantime the anemones stubbornly refused to divide.

The KQED lead videographer for the Deep Look video series, Josh Cassidy, who would had recorded the anemones dividing if they had divided and if the marine lab were not closed, asked me over the summer if we could somehow arrange to meet up to film something else. He had heard of some research that showed the emergent property of sea stars bouncing as they walk along on their many tube feet. Is there any way, he asked, that he could film some of the stars at the lab?

Well, filming at the lab was out of the question. Only essential personnel are allowed in the buildings, and there was no way I could sneak in Josh and all of his gear. We discussed options such as meeting up at a beach but I decided that I needed more control of the site to keep things safe for the animals. We ended up borrowing some friends' back yard for the day, which worked out pretty well. They have a covered pavilion, which was ideal because of course it turned out to be hot the day we filmed. I had several bags of frozen seawater to keep things cool-ish, two coolers for the movie stars themselves, a battery-operated air pump, and 30 gallons of seawater on hand.

Filming for production purposes takes a really long time. Even for a short film, we were working most of the day. Because of course most of the stars were uncooperative. They don't have anything even remotely resembling a brain, but damn if they can't bugger things up. I was feeling kind of bad that my animals were being such troublemakers; Josh, fortunately, was much more patient with them.

And here's the film! You'll see my right hand for about 1.5 seconds.

I didn't realize this at the time, but Josh also writes an article for each episode of Deep Look, for the KQED website. For this episode the article describes the research into the biomechanics of sea star bouncing. I'm quoted at the end.

So watch this short film. I hope it helps put a little bounce in your step.

5

On the afternoon of July 31, 2020 the world of invertebrate biology and marine ecology in California lost a giant in our field. Professor Emeritus John S. Pearse died after battling cancer and the aftereffects of a stroke.

John Pearse in the intertidal at Soquel Point
2017-05-28
© Allison J. Gong

John was one of the very first people I met when I came to UC Santa Cruz. Before we moved here, my husband and I came and met with John, who was not my official faculty sponsor but agreed to show us around so we could check out different areas for a place to live. In fact, I had applied to the department to do my graduate work in John's lab, but because he was considering retirement the department wouldn't let him take on a new Ph.D. student. But when we needed some help getting acquainted with Santa Cruz, John and his wife, Vicki Buchsbaum Pearse, graciously let us stay at their house and spent a day driving us around town and showing us eateries as well as potential neighborhoods.

By happenstance we ended up living down the hill from John and Vicki. We had met their blue duck, Lily, and I used to fill spaghetti sauce jars with snails from our tiny yard and trudge up the hill to feed them to her. She gobbled them up like they were her favorite treat.

As one of the regional experts in invertebrate biology, John was on all of my graduate committees. There were always a half-dozen or so of us grad students working with invertebrates, and we all tended to hang out together. John was one of the things we shared in common. And even if he wasn't technically on one's committee, he would always be available for consultation or advice as needed.

When John retired, he didn't leave the campus. He remained a presence at the marine lab, and still did field work. He started incorporating young students in his long-term intertidal monitoring research, which morphed into the LiMPETS project. The combination of working with students while producing robust scientific data was the perfect distillation of John's legacy. He said this about LiMPETS:

This is one of the best things I could ever do to enhance science education and conservation of our spectacular coastline. Working with teachers and their students is a wonderful and fulfilling experience.

John S. Pearse, Professor Emeritus
UC Santa Cruz

The last time I saw John was in the summer of 2019, during his annual Critter Count. He started these Critter Counts back in the 1970s, monitoring biota at two intertidal sites in Santa Cruz. These sites have since been incorporated into the LiMPETS program. I'm sure it made John smile whenever he thought of generation after generation of schoolkids traipsing down to the intertidal with their quadrats and transect lines, counting organisms the way he had for so many years.

When I started teaching my Ecology class, John suggested that I take the students out to Davenport Landing to monitor at the LiMPETS site there. That is another of his long-term sites, and he was worried about losing information if it were not sampled at least once a year. My students have done LiMPETS monitoring three years now, and John accompanied us on at least two of those visits. I tried to impress upon the students that having John Pearse himself come out with us was a Big Deal, but am not sure I was able to convince them of how fortunate they were. I bet there are a lot of marine biologists in California who would dearly love to go tidepooling with John. And now no one else will.

John Pearse and Todd Newberry, the other professor who gets the blame for how I think about biology, taught an Intertidal Biology class. I came along on many of the field trips the last year they taught it. I remember getting up before dawn to drive down to Carmel, park in the posh neighborhood streets, and walk down to meet John and Todd in the intertidal. I remember slogging through the sticky mud at Elkhorn Slough, digging for Urechis and hearing John shout "It's a goddamned brachiopod!" from across the flat. I remember bringing phoronid worms back to the lab, looking at them under the scope, and watching blood flow into and out of their tentacles. I remember John taking an undergraduate, Jen, and me out to Franklin Point, and showing me my very first staurozoans. That was probably around 1996, and I'm still in love with those animals.

I'm no John Pearse or Todd Newberry, but I'm a small part of their giant legacy in this part of the world. I strive to instill in my students the joy and intellectual pleasure in studying the natural world that I inherited from John and Todd. Partly to honor them, but mostly because it suits my own inclinations, I'm on a one-woman crusade to bring natural history back into modern science and science education.

I've spent the last two mornings in the intertidal at two of the LiMPETS sites, as part of a personal tribute to John. I thought there would be no greater way to memorialize John than by spending some quality time in the intertidal, where he trained so many young minds. I was thinking of him as I took photos, and thought he would be pleased if I shared them.

Natural Bridges—4 August 2020

Shore crab (Pachygrapsus crassipes)
2020-08-04
© Allison J. Gong

And because, like me, John had a special affinity for the anemones:

Sunburst anemone (Anthopleura sola)
2020-08-04
© Allison J. Gong
Sunburst anemone (Anthopleura sola)
2020-08-04
© Allison J. Gong

And he would have loved this. What is going on here? How did this pattern come to be?

Anemones (Anthopleura elegantissima and possibly A. sola)
2020-08-04
© Allison J. Gong

And look at this, three species of Anthopleura in one tidepool! Can you identify them?

Tidepool at Natural Bridges
2020-08-04
© Allison J. Gong

Davenport Landing—5 August 2020

It was windy and drizzly this morning. I ran into a friend, Rani, and her family out on the flats; they were leaving as I arrived. I hadn't seen her since before the COVID-19 lockdown began back in March. She was also visiting the tidepools to honor John Pearse. We chatted from a distance and exchanged virtual hugs before heading our separate ways.

It felt like a John Pearse kind of morning. I recorded the video clip I needed for class, collected some algae and mussels for a video shoot tomorrow, and took a few photos.

A typical intertidal assemblage (sea stars, sea anemones, and algae) at Daveport Landing
2020-08-05
© Allison J. Gong

And even though I'm not very good at finding nudibranchs, even I couldn't miss this one. It was almost 4 cm long!

Nudibranch (Triopha maculata) at Davenport Landing
2020-08-05
© Allison J. Gong

The ultimate prize for any tidepool explorer is always an octopus. When I take newbies into the field that's what they always want to see. I have to explain that while octopuses are undoubtedly there and common, they are very difficult to find. You can't be looking for them, unless you really like being frustrated.

But John must have been with me in spirit this morning, because I found this:

Red octopus (Octopus rubescens) at Davenport Landing
2020-08-05
© Allison J. Gong

It was just a small one, with the mantle about as long as my thumb. I found it because I spotted something strange poking out from a piece of algae. It was the arm curled with the suckers facing outward. I touched it, and the arm retracted. It didn't seem to like how I tasted.

And lastly, for me this is the epitome of John Pearse's legacy: Working in the intertidal, showing students how to identify owl limpets. I hope they never forget what it was like to learn from the man who with his wife, literally wrote the book about invertebrates and founded LiMPETS.

John Pearse in the intertidal with my students
2016-04-29
© Allison J. Gong

RIP, John S. Pearse. You left behind some enormous shoes to fill and a legacy that will stretch down through generations. I count myself lucky to have spent time with you in the field and in the lab. While I will miss you sorely, it is my privilege to pass on your lessons. Thank you for all you have taught me.

Every summer, like clockwork, my big female whelk lays eggs. She is one of a pair of Kellett's whelks (Kellettia kellettii) that I inherited from a labmate many years ago now. True whelks of the family Buccinidae are predatory or scavenging snails, and can get pretty big. The female, the larger of the two I have, is almost the length of my hand; her mate is a little bit smaller.

Many marine snails (e.g., abalones, limpets, and turban snails) are broadcast spawners, spewing large numbers of gametes into the ocean and hoping for the best. These spawners have high fecundity, but very few, if any, of the thousands of eggs shed will survive to adulthood. We say that in these species, parental investment in offspring extends only as far as gamete production. Fertilization and larval development occur in the water column, and embryos and larvae are left to fend for themselves.

The whelks, on the other hand, are more involved parents. They maximize the probability of fertilization by copulating, and the female produces yolky eggs that provide energy for the developing embryos and larvae. Rather than throw her eggs to the outside world and hoping for the best, the female whelk deposits dozens of egg capsules, each of which contains a few hundred fertilized eggs.

Over a period of about three weeks I shot several time-lapse video clips of the mama whelk laying eggs. Due to the pandemic we need to work in shifts at the lab. Fortunately I have the morning shift, which means I can start as early as I want as long as I leave before 11:00 when the next person comes in. Each 2.5-hr stint at the lab yielded about 30 seconds of video, not all of which was interesting; even in time-lapse, whelks operate at a snail's pace. Still, I was surprised at how active the female could be while she was apparently doing nothing.

The freshly deposited capsules are a creamy white color, as are the embryos inside them. As the embryos and then larvae grow, they get darker. Each of the fertilized eggs develops through the first molluscan larval stage, called a trochophore larva, within its own egg membrane. The embryo, and then the trochophore, survives on energy reserves provided by the mother snail when she produced the egg. These larvae don't hatch from their egg membrane until they've reached the veliger stage.

Pumpkin seed-shaped egg capsule of the whelk Kellettia kellettii, 13 mm tall.
Egg capsule of Kellettia kellettii
2020-06-20
© Allison J. Gong
Veliger larva of Kellettia kellettii
2020-07-25
© Allison J. Gong

The veliger larva gets its name from a lobed ciliated structure called a velum. Gastropods and bivalves have veliger larvae. As you might expect, the bivalve veliger has two shells, and the gastropod veliger has a coiled snail shell. These Kellettia veligers have dark opaque patches on the foot and some of the internal organs. That coloration is what you see in the photo of the egg capsule. You can see below which of the egg capsules are the oldest, right?

Mated pair of Kellettia kellettii and their egg capsules
2020-07-25
© Allison J. Gong

By the time the veligers emerge from the egg capsule, they have burned through almost all of the energy packaged in the yolk of the egg. They need to begin feeding very soon. The current generated by the beating cilia on the velum both propels the larva through the water and brings food particles to the larva's mouth. The velum can be pulled into the shell, and, as in any snail the opening to the shell can be shut by a little operculum on the veliger's foot. As is the case with most bodies, the veliger is slightly negatively buoyant, so as soon as it withdraws into the shell it begins to sink. However, once the velum pops back out the larva can swim rapidly.

Watch how the veliger swims. You can also see the heart beat!

So now the egg capsules are being emptied as the larvae emerge. I'm not keeping the veligers, so they are making their way through the drainage system back out to the ocean. As of now there are no iNaturalist observations of Kellettia kellettii in the northern half of Monterey Bay, so it appears that for whatever reason the whelks have not been able to establish viable populations here. Or it might be that the whelks are here but there aren't enough SCUBA divers in the water to see them.

These little veligers will be very lucky if any of them happen to encounter a subtidal habitat where they can take up residence as juvenile whelks. Even for animals that show a relatively high degree of parental care, the chances of any individual larva surviving to adulthood are exceedingly small. However, for the reproductive strategy of Kellettia to have evolved and persisted, there must be a payoff. In this case, the reward is an equal or greater reproductive success compared to snails that simply broadcast thousands of unprotected eggs into the water. Some gastropods such as the slipper shell Crepidula adunca, take parental care even further than Kellettia; in this species the mother broods her young under her shell until they've become tiny miniatures of herself, then she pushes them out to face the world and find a turban snail to live on. Crepidula adunca does not have a swimming larval stage at all. The fact that we see a variety of strategies—many eggs with little care, fewer eggs with more care, and brooding—indicates that there's more than one way to be successful.

Biology is a field of science with very few absolutes. For every rule that we teach, there seems to be at least one exception. I imagine this is very frustrating for students who want to know that Something = Something every single time. It certainly is easier to remember a few rules that apply to everything, than to keep track of all the cases when they don't.

Take, for example, the tube feet of sea stars. Among the generalities that we teach are: (1) sea star tube feet are used for locomotion and feeding; and (2) sea star tube feet are used to stick firmly to rocks and to pry open mussel shells. And we can show many examples of stars clinging to vertical and overhanging surfaces.

Purple sea star sticking to vertical rock surface in the intertidal
Ochre star (Pisaster ochraceus) at Franklin Point
2015-07-15
© Allison J. Gong
Purple sea star sticking to granite boulder in the intertidal
Ochre star (Pisaster ochraceus) at Mitchell's Cove
2019-06-03
© Allison J. Gong

Sometimes we can even find Pisaster doing both at the same time:

Ochre star (Pisaster ochraceus) wrapped around mussel(s) (Mytilus californianus) while clinging to an overhanging surface at Natural Bridges
2020-07-06
© Allison J. Gong

A photo like the one above is merely a snapshot of an event that lasts for hours. What's going on in there? Chances are it's a life-or-lunch battle, with the star trying to pry open the mussel just enough to slide its stomach between the shells while the bivalve is holding its shells clamped shut for dear life.

Each of these behaviors-—sticking to rocks and prying open mussels—is possible because Pisaster ochraceus has suckered tube feet. The tube foot itself has a flattened surface that squishes out a tiny dab of sticky adhesive glue. Together, the tube feet can adhere quite strongly to hard surfaces. I know from experience that it is impossible to pry an ochre star off a rock after it has had a chance to hang on, unless you're willing to damage several dozen tube feet. The tube feet will grow back, but there's no point in causing harm to the animal.

Arm tip of a sea star in a tidepool, showing suckered tube feet sticking to a rock
Tube feet at the arm tip of Pisaster ochraceus
2019-06-18
© Allison J. Gong

So that's the general story we teach in school. For most students, that's the entire story. However, it's always the exceptions, the deviations from the norm, that are the most interesting.

Not every sea star clings to rocks in the intertidal. There are several species that are equally at home on both rocks and sand. And among the rock-clingers, not all are as strong as Pisaster ochraceus. The ochre star's sucker-shaped tube feet are an example of the relationship between form and function: the tube feet's morphology provides the surface area for adhesion that allows the animal to feed and locomote over hard surfaces.

Spiny sand star (Astropecten armatus) at the Seymour Marine Discovery Center
2020-07-15
© Allison J. Gong

As you might expect, sea stars that don't cling to rocks and pry open mussels may not have sucker-shaped tube feet. The spiny sand star, Astropecten armatus, has pointed tube feet! It's hard to see exactly what the tube feet look like in the photo, but here's a video:

See how the tube feet on the underside of that arm end in points rather than suckers? If we revisit the notion of form and function, what questions come to mind when you look at the morphology of the tube feet? And given Astropecten's common name and its habitat, can you think of how it can survive and get around without the sticking power of Pisaster's tube feet?

Observation of Astropecten in its natural habitat would show that it spends a lot of time buried in the sand. It somehow has to get below the surface of the sand, where it feeds on olive snails or other animals that live buried there. How can it do that? Would the generalized sea star sucker-shaped tube feet that we teach to students be useful for burrowing? We can also think about it in a more familiar context: If you had to dig a hole in the ground, would you reach for a plunger? Clearly you wouldn't. You'd use a shovel, or a spade.

Astropecten's pointed tube feet are perfect for punching down between sand grains, enabling the star to work its way down into the sand. The sand star has hundreds of tiny spades at its disposal to use for digging. Circular structures shaped like miniature horse hooves wouldn't be very good at this job, nor would pointed tube feet be very good at sticking to rocks. This animal doesn't obey the "rules" of sea star biology, but form and function, as always, go together.

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