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2

Sometimes things just work out, through no fault of my own. In terms of good minus tides occurring in daylight hours, this weekend's tides are the best we will have all season. Today (Saturday 29 May) is the third of five intertidal excursions I have planned. This morning I went up to Pistachio Beach to collect some things for the Seymour Center. I always feel a teensy bit apprehensive agreeing to collect for anybody but myself, because it is quite likely that I will get skunked and not be able to bring back what is needed. So usually I just agree to keep my eyes open for things that are on the wish list and make no promises.

The current wish list for the Seymour Center includes fishes. I've already brought them some sculpins and a clingfish, but small pricklebacks are also welcome. Pistachio is a popular place for people who fish for large pricklebacks. Apparently they (the pricklebacks) put up a good fight and make tasty eating. The usual way of fishing for them is poke-poling. I am not entirely sure how that works, but it involves a long pole and baited hooks. I think the idea is to lure a prickleback out from its hiding place at low tide, when it is sort of stranded away from open water. Adults get up to 70-80 cm long, and are as big around as my forearm.

Unlike the fishermen, I was fishing for young pricklebacks, hoping to find some that were about the length of my hand. Possessing the ideal set of characteristics for avoiding capture—a long eel-like body, small head, slimy coating, and the ability to augur really quickly into even the tiniest crack amongst the cobbles—these small fish led me on a merry chase for quite a while. However, the advantages that I have over even a wily prickleback are an enlarged cerebral cortex, opposable thumbs, and the dexterity to use both a dip net and a zip-loc baggie. When all was said and done I had two appropriately sized pricklebacks in my baggie, and two others had gotten away from me. Oh, and I did also bag another clingfish!

Having had that bit of success and not wanting to press my luck, I started poking around just for the hell of it, without any clear objective in mind. As I've said before, what we gain from a super low tide like this (-1.6 ft) is not only access to more real estate in the low intertidal, but more time to spend there before the tide returns. I took lots of photos, which I will present in chronological order. These will give you an idea of what it was like out there this morning.

Even the hike across the beach yielded something nice—this small stand of Postelsia palmaeformis, the sea palm. These poor junior kelps will be taking a beating with these spring tides rushing up and down. That's the price they pay for living out there on those exposed rocky points.

Group of 6 sea palms on the beach
06:53 Postelsia palmaeformis
2021-05-29
© Allison J. Gong

The leather star Dermasterias imbricata isn't one of the most common stars in the intertidal around here. It was one of the species that was hit pretty hard by the most recent outbreak of Sea Star Wasting Syndrome. We see one every so often, but they are nowhere as abundant as the ochre stars or bat stars.

07:10 Dermasterias imbricata
2021-05-29
© Allison J. Gong

Pistachio Beach isn't the best place for large anemones, but of course there are some. This is one of the few big Anthopleura anemones that I saw today. There are many of the small cloning anemones, A. elegantissima, in the high intertidal, as well as the moonglow anemones, A. artemisia, in the mid and low sandy areas.

07:12 Anthopleura xanthogrammica
2021-05-29
© Allison J. Gong

I was so pleased to see my favorite red alga doing really well in the low zone! It is so pretty.

Red seaweed
07:29 Erythrophyllum delesserioides
2021-05-29
© Allison J. Gong

And at the same time I accidentally discovered a pretty big rock crab, which was tucked under a rock. For its species, this one was pretty calm and didn't come at me with big claws up. It could be that this crab is a male, and is clasping a female beneath him. I didn't check.

Dorsal view of a rock crab
07:29 Romaleon antennarium
2021-05-29
© Allison J. Gong

One of the things I found while turning over rocks to look for fish is this purple urchin:

Sea urchin with purple and green coloration
08:02 Strongylocentrotus purpuratus
2021-05-29
© Allison J. Gong

And a bit later, a nice healthy group of Dictyoneurum californicum. As these thalli age, they will develop longitudinal splits at the base of the blades. Right now they are young and crispy.

Blades of a brown seaweed with a waffle-like texture
08:15 Dictyoneurum californicum
2021-05-29
© Allison J. Gong

And who can resist such an exuberantly decorated limpet? Certainly not I! Reminds me of the fancy hats that ladies used to wear for Easter. Or Beach Blanket Babylon.

Limpet heavily fouled with encrusting and upright coralline algae
08:28 Limpet, probably Lottia sp.
2021-05-29
© Allison J. Gong

Chitons, the overlooked molluscs that reach peak abundance and diversity in the intertidal, can be very common along the coast. Species composition varies from site to site, though. Here at Pistachio Beach, the two species of Tonicella are very common. I found several of them on the undersides of rocks. This one is T. lokii.

Chiton with dark wavy lines on the shell plates and alternating pink and beige patches on the girdle
08:52 Tonicella lokii
2021-05-29
© Allison J. Gong

After two hours of catching fish and looking around, I was getting cold. Time to head back up and out. That took an additional half-hour or so, because I kept getting distracted by the algae. For example, look at how beautiful this Fucus is. And note the swollen tips, which mean this thallus is getting sexy. 'Tis the season, after all.

Olive-green seaweed with wide dichotomous branches and swollen branch tips
09:15 Fucus distichus
2021-05-29
© Allison J. Gong

One of the other rockweeds, Pelvetiopsis limitata, was also very thick and abundant.

Olive-green seaweed with narrow dichotomous branches
09:19 Pelvetiopsis limitata
2021-05-29
© Allison J. Gong

The rockweeds share the high intertidal with a few species of red algae. The most common reds in this zone are the two (or however many there are) species of Mastocarpus, and Endocladia muricata.

Reddish-brown seaweed with wavy blades, covered with tiny bumps
09:21 Mastocarpus papillatus
2021-05-29
© Allison J. Gong

I always want to stop and look around in the high zone on my way down. Because when I walk past sights like this, it's hard not to stay and study more closely. Then I remember that I can take as much time as I want in the high zone on the way out. This morning I took lots of photos of these reds and rockweeds.

How many different types of seaweed can you see?

09:24 High intertidal algal assemblage
2021-05-29
© Allison J. Gong

So there you have it, my morning summarized in about a dozen photos. I hope your Saturday was as enjoyable as mine was!

1

As we speed towards the summer solstice the days continue to get longer. The early morning low tides are much easier to get up for, as the sky is lightening by 05:30. Even so, when traveling an hour to get to the site, it's nice when the low is later than that. This past Saturday the low wasn't until 08:00. My parents were in Monterey for the weekend, so I decided it would be a good day to work the tide at the southern end of Monterey Bay, and then visit my parents. The Monterey Peninsula has some of the most spectacular tidepooling terrain in the region, and if I lived closer you can bet I'd know those sites better. Not that there is anything at all wrong with the sites on my end of the Bay and up the coast. But sometimes it's good to get out of one's comfort zone and explore the less well known.

Rocks and tidepools
Rocky intertidal at Asilomar State Beach
2021-05-15
© Allison J. Gong

So explore we did. It was cold and windy. The tide wasn't all that low and the swell was up, so we didn't get beyond the mid-tidal zone. My hip boots have deteriorated to the point that I have pinprick leaks at the seam where the boot part meets the leg part. Usually the tiny leaks don't bother me, but when the water is cold I definitely feel the trickles. What all this means is that I didn't get down into the low zone, which is fine. Biodiversity is highest in the mid zone anyway. The mediocrity of the low tide meant that I had to keep an eye out for sneaker swells, so less heads-down poking around and more scanning from above and then zooming in on individual items of interest.

One thing we noticed right away is that groups of Tegula funebralis, the black turban snail, were clumped together above the waterline of the high pools.

I'm trying to decide whether or not this is noteworthy. The pattern did catch my eye, but that might be only because it's unusual (although not particularly interesting). It was a cold and drizzly morning, so the snails didn't have to worry about desiccation. Was the clumping together benefiting the snails in any significant way? Hard to say.

The T. funebralis were also clumping together in the water! Here's a large clump of Tegula shells in a pool.

Clump of black turban snails in a tidepool
Black turban snails (Tegula funebralis) and one hermit crab (Pagurus samuelis)
2021-05-15
© Allison J. Gong

Almost all of these are snails, but can you see the one that is a hermit crab?

Poor Tegula funebralis. It is so common that it is invisible and vastly underappreciated. I find them quite charming, though. There's something about a grazing snail's slow way of life that is very soothing. Not that you might not fall asleep waiting for them to do something interesting, but it is good to slow down to the pace of nature. Anyway, Tegula is one of my favorite animals, precisely because it is so unassuming and ignored. One of delightful things about Tegula funebralis is when it plays host to Crepidula adunca. I've written about the biology of C. adunca before and don't want to rehash that here. I just wanted to show off my favorite photo of this trip to Asilomar:

Black turban snail with two attached slipper snails
Black turban snail (Tegula funebralis) wearing two slipper snails (Crepidula adunca)
2021-05-15
© Allison J. Gong

I don't know why I like this photo so much. It certainly isn't the best shot I've ever taken. There isn't any vibrant color at all. The subjects are the same color as the background. But it works for me.

When it comes to a snail's pace, you can't find anything slower than Thylacodes. That's because Thylacodes squamigerus is the snail that lives in a calcareous tube. Much like a barnacle, or the serpulid worms that have similar tubes, Thylacodes makes one decision about where to live and lives there for the rest of its life. I see Thylacodes at places like Pigeon Point up north, but they are much more abundant on the Monterey Peninsula.

Tube snail (Thylacodes squamigerus)
2021-05-15
© Allison J. Gong

And the snail winners in the Most Likely to be Overlooked have got to be the littorines. These little snails (most of which are smaller than 15 mm) live in the highest intertidal, where they get splashed by the ocean just often enough to keep their gill sufficiently moist. They are never entirely submerged, but they do tend to gather in cracks, even the tiniest of which will hold water longer than a flat rock surface.

Littorines (Littorina keenae) in the splash zone
2021-05-15
© Allison J. Gong

If you look closely at the photo above, you might see pairs of mating snails. Given where they live, high up in the intertidal where they are rarely covered by water, broadcast spawning isn't a viable option for the littorines. They have to copulate. There are, I think, eight copulating pairs in this group of ~30 snails.

Copulating pairs of Littorina keenae
2021-05-15
© Allison J. Gong

Because Littorina's habitat makes broadcast spawning an unfeasible option, the snails must lay eggs. But the splash zone isn't a very friendly place for the eggs of marine animals. The littorines lay eggs in gelatinous masses in crevices or depressions where water will remain. After a week or so of development, the egg mass dissolves as it gets splashed, and veliger larvae emerge. They recruit back to the intertidal after spending some period of time in the plankton.

When all is said and done it's difficult to make the claim that snails live exciting lives. Nonetheless, they are interesting animals. The diversity of morphology and lifestyle we see in the intertidal snails makes them eminently worthy of study and appreciation. I like to think that, as biologists once again "discover" the usefulness of natural history, students will be encouraged to fill in some of the gaps in our understanding of these and other abundant animals.

For animals that do essentially nothing when you see them where they live, chitons have a lot of charm. They are the kind of animal that, once you develop the search image for them, you start seeing everywhere. It helps that they are easily recognized as being chitons because of their eight dorsal shell plates—nothing else looks like them. Depending on species, those shell plates can be smooth or sculpted, and pigmented or not. Patterns of sculpting and pigmentation (or lack thereof) are diagnostic features used to distinguish different species. Some species are reliably consistent in appearance and look the same wherever you happen to see them. Other species show a lot of phenotypic variation, often even at a single site.

One of my favorite chitons is Mopalia muscosa, the mossy chiton. It's one of the easiest of our chitons to identify, because its girdle (the layer of tough tissue in which the shell plates are embedded) is densely covered by long, curved spines. They're called spines, but they're quite soft and flexible. Your basic Mopalia muscosa looks like this:

Mossy chiton with bare shell plates, in the rocky intertidal
Mossy chiton (Mopalia muscosa) at Pigeon Point
2016-04-24
© Allison J. Gong

Mopalia muscosa is one of the species whose appearance is quite variable. Many of them wear algae, usually reds but occasionally greens or browns, on their shell plates. Not all species of chiton do this. I've often wondered why some chiton species wear algae and others do not. This individual is probably fairly old, judging by the worn condition of the shell plates. The plates show signs of erosion, but are not decorated. There are some small pieces of coralline algae amongst the spines of the girdle, though, which I always associate with age. Smaller, and presumably younger, M. muscosa tend not to have algae on the girdle even if they are wearing some on the shell plates.

The degree of shell decoration in M. muscosa varies from none, as above, to heavy encrustation. This individual below has been colonized by only a small bit of coralline algae and perhaps some brown diatom-ish film on the edges of the shell plates:

Mossy chiton (Mopalia muscosa) at Pistachio Beach
2021-02-09
© Allison J. Gong

This next one has only a small bit of coralline alga, but sports a jaunty sprig of something quite a bit larger.

Mossy chiton (Mopalia muscosa) at Asilomar
2019-07-04
© Allison J. Gong

This season's fashionable chiton will go all out with the coralline algae, wearing both encrusting and upright branching forms. Look at this:

Mossy chiton (Mopalia muscosa) at Pigeon Point
2017-06-28
© Allison J. Gong

and this:

Mossy chiton (Mopalia muscosa) at Pigeon Point
2018-01-01
© Allison J. Gong

Sometimes the chitons wear the larger leafy red algae, in addition to or in place of the coralline algae. I always think that these individuals must be very old, by chiton standards.

Mossy chiton (Mopalia muscosa) at Pigeon Point
2020-11-14
© Allison J. Gong

And sometimes the chitons are so covered with algae that they blend in perfectly with the surrounding environment.

Mossy chiton (Mopalia muscosa) at Pistachio Beach
2021-04-06
© Allison J. Gong

These chitons can get very heavily fouled by algae. Is there any benefit to the chiton, to carry around a load of red algae? And if wearing algae is for some reason advantageous, is there a way for a chiton to attract algae to settle on their shell plates? Well, let's think about that. Chitons' main predators would be sea stars, crabs, and birds. Sea stars do not locate prey visually, so camouflage would not be very helpful in avoiding them. Birds such as oystercatchers and surfbirds certainly do pry up chitons and limpets, and blending in with the background just might help a chiton go unnoticed by an avian hunter.

Regarding the matter of how the algae end up living on chitons' bodies, I want to start with the question of how prevalent algal fouling is on Mopalia muscosa, and the extent of fouling on the chitons that are wearing algae. A little research study might be a fun way to spend my time in the intertidal. Pigeon Point is a lovely site on a foggy summer morning, and many of the most heavily decorated M. muscosa in my photo library are from there. Yes, I can foresee several visits up the coast over the next few months. Laissez les bons temps rouler!

This morning I went to Pigeon Point to poke around and do some collecting. It's a favorite site of mine, as it's exposed and dynamic, with the diversity you'd expect. Of the sea stars, the most common by far are the six-armed stars in the genus Leptasterias. They are small (less than 8 cm in diameter, often smaller than 1 cm), somewhat drably colored, and sometimes on the underside of rocks, all of which means that they are not always conspicuous. But once you get the right search image, you see them everywhere.

Six arms, see?

Sea stars are well known for their ability to regenerate lost arms. It is not uncommon to see a star that looks healthy in every way except that one of its arms is shorter than the others. This must happen in Leptasterias, too. Searching through my library of pictures of Leptasterias, I did find a couple of examples of regeneration.

When these stars finish regrowing those arms, they will have the typical number of arms for the genus, which is six.

Today I saw something that I'd never seen before. It was a Leptasterias that was regenerating arms. Only this was weird. It had three full-size arms and was growing four!

Sea star with 3 arms and regenerating 4 arms
Leptasterias star regenerating lost arms at Pigeon Point
2021-04-03
© Allison J. Gong

When (if) this star survives, it will eventually have seven arms. And that's strange. I asked my friend Chris Mah, who is the sea star systematist at the Smithsonian, if it was common for Leptasterias to do this. He said he'd never seen it, either. So it is indeed a rare phenomenon.

Now, there are stars in the genus Linckia that actually reproduce by deliberately leaving behind an arm, which then goes on to regenerate the rest of the body. While they do so they look like comets:

"comet" star. One arm is regenerating the remaining arms and central disc.
Linckia multiflora comet in the Maldives
Ahmed Abdul Rahman and Frédéric Ducarme for MDC Seamarc Maldives (CC BY-SA 4.0)

My regenerating Leptasterias isn't quite a comet, but it is doing something equally strange and wonderful. I really wished I could bring it to the lab and keep an eye on it over the next several months. However, Leptasterias are on the no-take list, I think because their populations are so patchy. It is extremely unlikely that I will ever see that same individual again, so we will never know what happens to it. Unless, of course, I happen to come across a 7-armed Leptasterias at Pigeon Point sometime in the future. If you see it, take a photo and let me know!

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.

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!

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.

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.

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