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Tag: marine invertebrates

A steady diet of worms

Posted on 2018-06-132023-01-06 by Allison J. Gong

Today is the first day of the week of low tides dedicated to Snapshot Cal Coast, a statewide citizen science project headed in my area by the California Academy of Sciences. This week groups and individuals will be making photographing the organisms they see in the ocean or along the coast, and uploading observations to iNaturalist. Participants will include both scientists and non-scientists, making the week-long event one of the biggest citizen science projects that I regularly take part in. Next Monday I’ll be taking a group of Seymour Center volunteers and staff up to Davenport to conduct a Bioblitz. The other days I’ll be out on my own, or with 1 or 2 people.

This morning the low tide was very early (-1.3 feet at 05:09), so I stayed close to home and went to Natural Bridges. The tide was low but the swell was big and I wasn’t able to get down to the low spots I could normally reach with this kind of tide. However, this meant that I could spend more time in the low-mid-intertidal, where there is a lot of biodiversity to document.

Today I want to write about polychaete worms. These are the segmented marine worms in the Phylum Annelida, which also includes earthworms and leeches.

Worm #1: Phragmatopoma californica

One of the most conspicuous inhabitants of this zone is the tube-dwelling polychaete worm, Phragmatopoma californica. This worm has a couple of common names: honeycomb worm, which refers to the mounds of tubes they build; and sandcastle worm, for the fact that the tubes are built of cemented sand grains. In effect, these worms are tiny masons!

Mound of Phragmatopoma californica tubes at Natural Bridges
26 May 2017
© Allison J. Gong

Each of the tubes is inhabited by a single worm. Mounds form because competent Phragmatopoma larvae, looking for a place to settle out and live permanently, are attracted to the tubes of existing adults. This phenomenon is called gregarious settlement. Once settled and metamorphosed, juvenile worms build their tubes by selecting sand grains and cementing them together around a lining of chitin-like material. How they do it, underwater, nobody knows. And these tubes are tough! The worm inside is skinny, and a humongous one would be all of 4 cm long, but it takes a lot of force to pry apart those sand grains. The openings to the tubes are 5-10 mm in diameter. Each worm can close off its tube with a circular-ish disc of stiff, fused chaetae called an operculum; this protects the worm from both predators and desiccation.

When the tide is out the worms withdraw into their tubes and clap the operculum down. They wait for the water to return. Phragmatopoma is a filter feeder; like most of the tube-dwelling polychaetes these worms use a crown of ciliated tentacles to create water currents that draw food particles to the mouth. When the tide is in the worms pull down the operculum and extend their feeding tentacles into the water. In the field, this is the most you can see of the worm’s body.

Feeding tentacles of Phragmatopoma californica at Natural Bridges
13 June 2018
© Allison J. Gong

Worm #2: Serpula columbiana

Many polychaetes live in tubes, and tubes can be made of a variety of materials. Phragmatopoma californica builds tubes out of sand grains. Another worm that I saw today, Serpula columbiana, builds tubes out of CaCO3 precipitated from seawater. Like other animals that build calcareous skeletons, S. columbiana may in the future have difficulty precipitating CaCO3 in an increasingly acidic ocean. Tubes of Serpula worms are white when new and soon become fouled with algal growth, and tend to wander over the substrate. The best photo I could take this morning is a little blurry but you can see the general morphology of the tubes.

Calcareous tubes of Serpula columbiana at Natural Bridges
13 June 2018
© Allison J. Gong

These worms are incredibly shy, and react to any perceived threat by pulling into their tubes. Their tentacles have tiny eyespots that can detect changes in light, so passing a hand over them can cause them to withdraw. Fortunately I was able to sneak up on one lazy worm in a pool, and grab a shot of its ‘head’ region. Worms that live in tubes are poorly cephalized, with none of the structures that we generally associate with a head. Serpula columbiana‘s ‘head’ looks like this:

Anterior end of Serpula columbiana at Natural Bridges
13 June 2018
© Allison J. Gong

The tentacles of S. columbiana are morphologically complex compared to those of Phragmatopoma. Serpula‘s tentacles are pinnate, or feather-shaped, and in cross-section look like a V. Cilia on the side branches of the tentacle create the feeding current, and food particles are transported by other cilia down the trough of the V to the mouth.

See that long, trumpet-shaped structure? That’s the worm’s operculum!

Worm #3: Unidentified cirratulid

Unlike Serpula and Phragmatopoma, worms of the Family Cirratulidae don’t live in tubes. Instead, they live with most of the body hidden in crevices, and extend tentacles to feed.

Feeding tentacles of an unidentified cirratulid polychaete worm at Natural Bridges
13 June 2018
© Allison J. Gong

As you can imagine, it is extremely difficult to identify a worm when all you can see of it are its tentacles; with the rest of the body hidden in a crevice, there are no visible characteristics to use to distinguish species. Cirratulids use their tentacles to feed, but in a way that is entirely unlike how Phragmatopoma and Serpula use theirs. Instead of feeding on particles suspended in the water, cirratulids are deposit feeders. They sweep their tentacles across the surface and collect organic deposits. Sticky mucus on the tentacles picks up organic matter, and cilia on the tentacles sweep the organic matter to the worm’s mouth.

Don’t believe me? Watch this!

It doesn’t matter if the surrounding substrate is sand or rock. The cirratulid’s sticky tentacles are very effective at gathering organic muck.

Worm #4: Flabesymbios commensalis

This worm remains an enigma. There doesn’t seem to be much known about its biology. I have seen them twice, both times on the body of purple urchins (Strongylocentrotus purpuratus), and although the genus name has changed twice since the first time, I’m pretty sure it’s the same worm. As the species epithet commensalis implies, this worm is a commensal on sea urchins. This means that it neither harms nor benefits its echinoderm host. Similar to the worm I’ve seen on bat stars, F. commensalis presumably cruises over the urchin’s body and feeds on detritus or scraps of kelp that the urchin grabs.

When I took the photo in a tide pool this morning I didn’t see the worm. It wasn’t until I downloaded the pictures from the camera onto my computer that I saw it. See how well it blends in with the urchin’s color?

Purple urchin (Strongylocentrotus purpuratus) at Natural Bridges
13 June 2018
© Allison J. Gong

Here’s a tighter crop of that photo:

Flabesymbios commensalis on aboral surface of a purple urchin (Strongylocentrotus purpuratus) at Natural Bridges
13 June 2018
© Allison J. Gong

For many polychaete worms, another animal’s body seems to be the ideal habitat. And for some reason, echinoderms are likely hosts for such commensal worms. I’ve written about the bat star worms, here is the urchin worm, and there’s also a scale worm that I’ve seen crawling around on the body of a sea cucumber. What is it about echinoderms that makes them habitat for worms? Or is this type of commensalism also common, but less observed, between polychaetes and other non-echinoderm invertebrates? I don’t know the answer to either of those questions, but am very intrigued.

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Things strange and beautiful

Posted on 2018-05-202023-01-06 by Allison J. Gong

This weekend I was supposed to take a photographer and his assistant into the field to hunt for staurozoans. I mean a real photographer, one who has worked for National Geographic. He also wrote the book One Cubic Foot. You may have heard of the guy. His name is David Liittschwager. Anyway, his assistant contacted me back in March, saying that he was working on something jellyfish-related for Nat Geo and hoped to include staurozoans in the story, and did I know anything about them? As in, maybe know where to find them? It just so happens that I do indeed know where to find staurozoans, at least sometimes, and we made a date to go hunting on a low tide. Then early in May the assistant contacted me to let me know that David’s schedule had changed and he couldn’t meet me today, and she hoped they’d be able to work with me in the future, and so on.

None of which means that I wouldn’t go look for them anyways. I’d made the plans, the tide would still be fantastic, and so I went. And besides, these are staurozoans we’re talking about! I will go out of my way to look for them as often as I can. Not only that, but I hadn’t been to Franklin Point at all in 2018 and that certainly needed to be remedied.

Pigeon Point, viewed from Franklin Point trail
19 May 2018
© Allison J. Gong

The sand has definitely returned. The beach is a lot less steep than it was in the winter, and some of the rocks are completely covered again. This meant that the channels where staurozoans would likely be found are shallower and easier to search. But you still have to know where to look.

Tidal area at Franklin Point
19 May 2018
© Allison J. Gong

See that large pool? That’s where the staurozoans live. They like areas where the water constantly moves back and forth, which makes them difficult to photograph in situ. And given that the big ones are about 2 cm in diameter and most of them are the same color as the algae they’re attached to, they’re a challenge to find in the first place. I looked for a long time and was about to give up on my search image when I found a single small staurozoan, about 10 mm in diameter, quite by accident. It was a golden-brown color, quite happily living in a surge channel. I took several very lousy pictures of it before coming up with the bright idea of moving it up the beach a bit to an area where the water wasn’t moving quite as much. I sloshed up a few steps and found a likely spot, then placed my staurozoan where the water was deep enough for me to submerge the camera and take pictures.

Staurozoan (Haliclystus sp.) at Franklin Point
19 May 2018
© Allison J. Gong
Staurozoan (Haliclystus sp.) at Franklin Point
19 May 2018
© Allison J. Gong

Cute little thing, isn’t it? I had my head down taking pictures of this animal, congratulating myself on having found it. When I looked around me I saw that I had inadvertently discovered a whole neighborhood of staurozoans. They were all around me! And some of them were quite large, a little over 2 cm in diameter. All of a sudden I couldn’t not see them.

Staurozoan (Haliclystus sp.) at Franklin Point
19 May 2018
© Allison J. Gong

I know I’ve seen staurozoans in the same bottle green color as the Ulva, but this time I saw only brown ones. As you can see even the animals attached to Ulva were brown. Staurozoans seem to be solitary creatures. They are not permanently attached but do not aggregate and are not clonal. Most of the ones I found were as singles, although I did find a few loose clusters of 3-4 animals that just happened to be gathered in the same general vicinity.

Trio of staurozoans (Haliclystus sp.) at Franklin Point
19 May 2018
© Allison J. Gong

Not much is known about the biology of Haliclystus, or any of the staurozoans. I collected some one time many years ago, and brought them back to the lab for closer observation. They seemed to eat Artemia nauplii very readily, and I did get to observe some interesting behaviors, but they all died within a week or so. Given that I can find them only in certain places at Franklin Point, they must be picky about their living conditions. Obviously I can’t provide what they need at the marine lab. The surging water movement, for example, is something that I can’t easily replicate. I need to think about that. The mid-June low tides look extremely promising, and my collecting permit does allow me to collect staurozoans at Franklin Point. Maybe I’ll be able to rig up something that better approximates their natural living conditions in the lab.

In the meantime, I just want to look at them.

Staurozoan (Haliclystus sp.) at Franklin Point
19 May 2018
© Allison J. Gong
Pair of staurozoans (Haliclystus sp) at Franklin Point
19 May 2018
© Allison J. Gong

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Squidlets

Posted on 2018-05-142023-01-06 by Allison J. Gong

Every once in a while some random person drops off a creature at the marine lab.  Sometimes the creature is a goldfish that had been a take-home prize at a wedding over the weekend (now weddings taking place at the Seymour Center are not allowed to include live animals in centerpieces). Once it was a spiny lobster that spent the long drive up from the Channel Islands in a cooler, and became the Exhibit Hall favorite, Fluffy. This time the objects had been collected off the beach and brought in by somebody who thought they might still be alive.

16 April 2018
© Allison J. Gong

These white objects are egg masses of the California market squid, Doryteuthis opalescens, that had been cast onto the beach at Davenport. Sometimes the masses are called fingers or candles, because they’re about finger-sized. Each contains dozens of large eggs. Squids, like all cephalopods, are copulators, and after mating the female deposits a few of these fingers onto the sea floor. Many females will lay their eggs in the same spot, so the eggs in this photo represent the reproductive output of several individuals. The cephalopods as a group are semelparous, meaning that they reproduce only once at the end of their natural life; salmons are also semelparous. After mating, the squids die. Not coincidentally, the squid fishing season is open right now, the idea being that as long as the squids have reproduced before being caught in seines, little harm is done to the population. Most of the time the squids are dispersed throughout the ocean, and the only time it is feasible to catch them in large numbers is when they gather to mate.

These egg masses look vulnerable, but they’re very well protected. The outer coating is tough and leathery, and the eggs must taste bad because nothing eats them. I’ve fed them to anemones, which will eat just about anything, and they were spat out immediately.

The eggs were brought to the Seymour Center because the person who brought them in thought they might make a good exhibit. I happened to be there that day and got permission to take a small subset of the bunch so I could keep an eye on them. And they did and still do make a good exhibit.

16 April 2018: I obtain squid eggs!

Egg mass, or ‘finger, of the California market squid Doryteuthis opalescens
16 April 2018
© Allison J. Gong

At this stage it is impossible to tell whether or not the eggs are alive. The only thing to do was wait and see.

30 April 2018: After waiting two weeks with apparently no change, I decided it was time to look at the egg fingers more closely again. Lo and behold, they are indeed alive! Look at the pink spots in the individual eggs–those are eyes. And if you can see the smaller pink spots, those are chromatophores, the ‘color bodies’ in the squids’ skin that allow them to perform their remarkable color changes.

Developing embryos of Doryteuthis opalescens
30 April 2018
© Allison J. Gong

9 May 2018: A week and a half later, the embryos definitely look more like squids! Their eyes and chromatophores have darkened to black now. The embryos are also more active, swimming around inside their egg capsules. You can see the alternating contraction and relaxation of the mantle, which irrigates the gills. Squids have two gills. More on that below.

At this point the squid fingers began to disintegrate and look ragged. They became flaccid and lightly fouled with sediment.

14 May 2018 (today): Almost a month after they arrived, my squid eggs look like they’re going to hatch soon! I didn’t see any chromatophore flashing, though.

In the meantime, some of the eggs on exhibit in the Seymour Center have already started hatching. The first hatchlings appeared on Friday 11 May 2018. The hatchlings of cephalopods are called paralarvae; they aren’t true larvae in the sense that instead of having to metamorphose into the adult form, they are miniature versions of their parents.

Peter, the aquarium curator at the Seymour Center, allowed me to take a few of the paralarvae in his exhibit and look at them under the scope. The squidlets are about 3mm long and swim around quite vigorously. Trying to suck them up in a turkey baster was more difficult than I anticipated. But I prevailed!

Paralarva of Doryteuthis opalescens
14 May 2018
© Allison J. Gong

You can actually see more of what’s going on in a video:

The cup-shaped layer of muscular tissue that surrounds the squid’s innards is the mantle. When you eat a calamari steak, you are eating the mantle of a large squid.The space enclosed by the mantle is called the mantle cavity. Because the paralarvae are transparent you can see the internal organs. Each of those featherlike structures is a ctenidium, which is the term for a mollusk’s gill. The ventilating motions of the mantle flush water in and out of the mantle cavity, ensuring that the gill is always surrounded by clean water.

And now we get to the hearts of the matter. At the base of each gill is a small pulsating structure called a branchial heart (‘branch’ = Gk: ‘gill’). It performs the same function as the right atrium of our own four-chambered heart; that is, boosting the flow of blood to the gas-exchange structure. So that’s two hearts. Between the pair of branchial hearts is the systemic heart, which pumps the oxygenated blood from the gills to the rest of the squid’s body. This arrangement of multiple hearts, combined with a closed circulatory system, allows cephalopods to be much more active swimmers and hunters than the rest of their molluscan kin.

I expect that my fingers will hatch very soon. If and when they do, it will be a challenge getting them to eat. I’ve never tried it myself, and cephalopods are known to be difficult to rear in captivity. But I’m willing to give it a shot!

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Trailblazers

Posted on 2018-02-152023-01-06 by Allison J. Gong

Who do you think makes these tracks in the sand?

15 February 2018
© Allison J. Gong

Any guesses?

Here’s another photo, taken from farther away to give you a bigger picture of the scale of things.

15 February 2018
© Allison J. Gong

Believe it or not, the maker of these trails is the little black turban snail, Tegula funebralis. They are one of my favorite animals in the intertidal, for a number of reasons:

  1. I always root for the underdog and the under-appreciated, and these snails are so numerous in the intertidal that they are practically invisible. People literally do not see them. I know, because I ask.
  2. They are very useful creatures to keep as lab pets. I throw a few of them into each of my seawater tables, except for the table that contains a resident free-ranging sea star, and they do a fantastic job keeping algal growth to a tolerable minimum. They’re my little marine lawnmowers!
  3. They come in very handy when I’m teaching invertebrate zoology. Students study them live to observe behavior, and the snails are not shy. They are very tolerant of being picked up and gently prodded, and soon emerge from their shells and carry on their little snail lives. Students also dissect them in lab to learn about gastropod anatomy.

So yes, these tracks in the sand are made by T. funebralis in the high intertidal. In areas where a layer of sand accumulates either at the bottom of a pool or on a flat exposed rock, it is not uncommon to see a turban snail pushing sand out of the way as it crawls along, like a miniature snow plow.

A black turban snail (Tegula funebralis) plowing through sand on a high intertidal rock at Natural Bridges
15 February 2018
© Allison J. Gong

Tegula funebralis and its congeners are called turban snails because their shells are shaped like turbans. Given their small size (a big T. funebralis would have a shell height of 2.5-3 cm), pushing sand around must be a tiresome chore. They do it because they have no choice. Most grazing gastropods, such as turban snails and limpets, can feed only when they are crawling. There may very well be a nice yummy layer of algal scum on the surface of this rock, but the snail has to push the sand out of the way before it can feed on it.

Here’s another photo, taken at the snail’s level.

Tegula funebralis plowing through sand at Natural Bridges
15 February 2018
© Allison J. Gong

This snail is pushing through a wall of sand as tall as itself! I don’t know about you, but I sure as heck couldn’t do that. Props to these little snails!

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Creepy crawlies

Posted on 2017-10-312023-01-06 by Allison J. Gong

There are certain creatures that, for whatever reason, give me the creeps. I imagine everyone has them. Some people have arachnophobia, I have caterpillarphobia. While fear of some animals makes a certain amount of evolutionary sense—spiders and snakes, for example, can have deadly bites—my own personal phobia can be traced back to a traumatic childhood event involving an older cousin and a slew of very large tomato hornworms. Even typing the words decades later makes me want to rub my hands on my jeans.

But enough about caterpillars. This Halloween I want to share something that isn’t nearly as disgusting, but can still creep me out sometimes. Commonly called skeleton shrimps, caprellid amphipods are a type of small crustacean very common in certain marine habitats. They are bizarre creatures, but a close look reveals their crustacean nature. For example, they possess the jointed appendages and compound eyes that only arthropods have.

Female caprellid amphipod (Caprella sp.)
22 October 2017
© Allison J. Gong

Around here the easiest place to find caprellids is at the harbor, where they can be extremely abundant. The last time I went to the harbor to collect hydroids for my class, the caprellids were swarming all over everything. When I brought things back to the lab I had to spend an hour or so picking the caprellids off the hydroids. I don’t think they eat the ‘droids, but they gallop around and keep messing up the field of view, making observation difficult. They’re essentially just a PITA to deal with, and everything is easier after they’ve been removed.

Caprellid amphipods (Caprella sp.) at the Santa Cruz Yacht Harbor
23 June 2017
© Allison J. Gong

Caprellids are amphipods, members of a group of crustaceans called the Peracarida (I’ll come back to the significance of the name in a bit). They have the requisite two pairs of antennae that crustaceans have, and seven pairs of thoracic appendages of varying morphology. Some of these thoracic legs are claws or hooked feet that like to grab onto things. A caprellid removed from whatever it’s attached to and placed by itself in a bowl of seawater thrashes around spastically. Only when it finds something to grab does it calm down. Even then, they attach with their posterior appendages and wave around the front half of the body in what I call the caprellid dance: they extend up and forward, and sort of jerk front to back or side to side. It isn’t pretty.

A bunch of caprellids removed from their substrate and dumped into a bowl together will use each other as something to grab. This forms the sort of writhing mass that makes my skin crawl. I was nice enough to give them a piece of bryozoan colony to hang onto, but even so they ended up glomming together.

Now, back to the thing about caprellids being peracarids. The name Peracarida means “pouch shrimp” and refers to a ventral structure called a marsupium, in which females brood their young. Males don’t have a marsupium, so adult caprellids are sexually dimorphic. When carrying young, a female caprellid looks like she’s pregnant. See that caprellid in the top photo? She’s a brooding female. That’s all fine, until her marsupium itself starts writhing. This ups the creepiness factor again. Here’s that same brooding female, in live action:

Crustaceans obviously don’t get pregnant the way that mammals do, but many of them spend considerable energy caring for their young. Well, females do, at least. A female caprellid doesn’t just carry her babies around inside a pouch on her belly. Although she isn’t nourishing them from her own body in the way of mammals (each of the youngsters in the marsupium is living off energy stores provisioned in its egg), the mother does aerate the developing young by opening and closing the flaps to the marsupium. This flushes away any metabolic wastes and keeps the juveniles surrounded by clean water. As the young caprellids get bigger, they begin to crawl around inside the pouch, and eventually leave it. They don’t depart from their mother right away, though; rather they cling to her back for a while, doing the caprellid dance in place as she galumphs along herself.

Until the juveniles strike out on their own they form a small writhing mass on top of a female who can herself be part of a larger writhing mass. And the sight through the microscope of all these long skinny bodies jerking around spasmodically can indeed be very creepy. Fortunately not as creepy as caterpillars, or I wouldn’t be able to teach my class or go docking with my friend Brenna. And it’s a good thing caprellids are small, ’cause if they were any bigger. . . just, no.

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The tiniest advantage

Posted on 2017-10-132023-01-06 by Allison J. Gong

Although the world’s oceans cover approximately 70% of the Earth’s surface, most humans interact with only the narrow strip that runs up onto the land. This bit of real estate experiences terrestrial conditions on a once- or twice-daily basis. None of these abiotic factors, including drying air, the heat of the sun, and UV radiation, greatly affects any but the uppermost few meters of the ocean’s surface so most marine organisms don’t need to worry about them. Despite the apparent paradox of where they live, intertidal organisms are also entirely marine–they cannot survive prolonged exposure to in air or freshwater. So how do they manage to live here?

Some organisms have a physiological tolerance for difficult conditions. These tidepool copepods and periwinkle snails, for example, are able to survive in the highest pools in the splash zone, where salinity can be either very high (due to evaporation) or very low (due to rain or freshwater runoff), dissolved oxygen is often depleted due to high temperature, and temperature itself can be quite warm. Sculpins and other tidepool fishes cope with low oxygen levels by gulping air and/or retreating to deep corners of their home pools.

Of course, animals that can locomote have the option of moving to a more favorable location. Other creatures, living permanently attached to their chosen site, aren’t quite so lucky. Let’s take barnacles as an example.

Nauplius larva of the barnacle Elminius modestus
© Wikimedia Commons

Barnacles have two planktonic larval stages: the nauplius and the cyprid. The nauplius is the first larval stage and hatches out of the egg with three pairs of appendages. It can be distinguished from the nauplius of other crustaceans by the presence of two lateral “horns” on the anterior edge of the carapace. The nauplius’s job is to feed and accumulate energy reserves. It swims around in the plankton for several days or perhaps a couple of weeks, getting blown about by the currents and feeding on phytoplankton.

Cyprid larva of a barnacle

After sufficient time feeding in the plankton, a barnacle nauplius metamorphoses into the second larval stage, the cyprid. A cyprid is a bivalved creature, with the body enclosed between a pair of transparent shells. It has more appendages than the nauplius, and these are more differentiated. If the nauplius has done its  job well, then the cyprid also contains a number of oil droplets under its shell. These droplets are of crucial importance, because the cyprid itself does not feed. For as long as it remains in the plankton it survives on the calories stored in those droplets. The cyprid’s job is to return to the shore and find a suitable place on which to settle. Somehow, a creature about 1 mm long, being tossed about by waves crashing onto rocks, has to find a place to live and then stick to it.

Returning to the topic of the challenges that marine organisms face when they live under terrestrial conditions, let’s see how these barnacles manage. Along the northern California coast we have a handful of barnacle species living in the intertidal. In the higher mid-tidal regions at some sites, small acorn barnacles of the genera Balanus and Chthamalus may be the most abundant animals.

Mixed population of the acorn barnacles Balanus glandula and Chthamalus dalli/fissus at Davenport Landing
27 June 2017
© Allison J. Gong

However, nowhere is a particular pattern of barnacle distribution more evident than at Natural Bridges. Here, the barnacles in the high-mid intertidal are small, and concentrated in little fissures and cracks in the rock.

I think most of these small (~5 mm) barnacles are Balanus glandula:

Small acorn barnacles (Balanus glandula) at Natural Bridges
11 October 2017
© Allison J. Gong

And here’s a closer look:

Small acorn barnacles (Balanus glandula) at Natural Bridges
11 October 2017
© Allison J. Gong

If all of the rock surfaces were equally suitable habitat, the barnacles would be distributed more randomly over the entire area. Instead, they are clearly segregated to the cracks in the rock. Each of these barnacles metamorphosed from a cyprid into a juvenile exactly where it is currently located. The cyprid may be able to move around to fine-tune its final location, but once the decision has been made that X marks the spot and the cyprid has glued its anterior to the rock, the commitment is real and lifelong. The barnacle will live its entire life in that spot and eventually die there. It is quite probable that cyprids landed in those empty areas on the rock, but they didn’t survive to adulthood.

How did this distribution of adult barnacles come to be?

There is one very important biological reason for barnacles to live in close groups, and that is reproduction. They are obligate copulators, which I touched on in this post, and as such need to live in close proximity to potential mates. But today I’m thinking more about abiotic factors. In a habitat like the mid-mid rocky intertidal, desiccation is a real and daily threat. Even a minute crack or shallow depression will hold water a bit longer than an exposed flat surface, giving the creatures living there a tiny advantage in the struggle for survival. No doubt cyprid larvae can and do settle on those empty areas of the rock. However, they likely die from desiccation when the tide recedes, leaving only the cyprids that landed in one of the low areas to survive and metamorphose successfully. There are other factors as well, such as the presence of adult individuals, that make a location preferable for a home-hunting cyprid. In addition to facilitating copulation, hanging out in a cluster slows down the rate of water evaporation, giving another teensy edge to animals living at the upper limit of their thermal tolerance.

Lower in the intertidal, where terrestrial conditions are mitigated by more time immersed, barnacles and other organisms do indeed live on flat rock spaces. But at the high-mid tide level and above, macroscopic life exists mostly in areas that hang onto water the longest. Pools are refuges, of course, but so are the tiniest cracks that most of us overlook. Next time you venture into the intertidal, take time on your way down to stop and salute the barnacles for their tenacity.

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My favorite larva — the actinotroch!

Posted on 2017-09-232023-01-06 by Allison J. Gong

Five days ago I collected the phoronid worms that I wrote about earlier this week, and today I’m really glad I did. I noticed when I first looked at them under the scope that several of them were brooding eggs among the tentacles of the lophophore. My attempts to photograph this phenomenon were not entirely successful, but see that clump of white stuff in the center of the lophophore? Those are eggs! Oh, and in case you’re wondering what that tannish brown tube is, it’s a fecal pellet. Everyone poops, even worms!

Lophophore of a phoronid worm (Phonoris ijimai)
18 Septenber 2017
© Allison J. Gong

Based on species records where I found these adult worms, I think they are Phoronis ijimai, which I originally learned as Phoronis vancouverensis. The location fits and the lophophore is the right shape. Besides, there are only two genera and fewer than 15 described species of phoronids worldwide.

Two days after I first collected the worms, I was watching them feed when I noticed some tiny approximately spherical white ciliated blobs swimming around. Closer examination under the compound scope showed them to be the phoronids’ larvae–actinotrochs! Actinotrochs have been my favorite marine invertebrate larvae–and that’s saying quite a lot, given my overall infatuation with such life forms–since I first encountered them in a course in comparative invertebrate embryology at the Friday Harbor Labs when I was in graduate school.

2-day-old actinotroch larva of Phoronis ijimai
22 September 2017
© Allison J. Gong

The above is a mostly top-down view on an actinotroch, which measured about 70 µm long. They swim incredibly fast, and trying to photograph them was an exercise in futility. They are small enough to swim freely in a drop of water on a depression slide, so I tried observing them in a big drop of water under a coverslip on a flat glass slide. At first they were a bit squashed, but as soon as I gave them enough water to wiggle themselves back into shape they took off swimming out of view.

Here’s the same photo, with parts of the body labelled:

2-day-old actinotroch larva of Phoronis ijimai
22 September 2017
© Allison J. Gong

The hood indicates the anterior end of the larva and the telotroch is the band of cilia around the posterior end. The hood hangs down in front of the mouth and is very flexible. At this stage the larva possesses four tentacles, which are ciliated and will get longer as the larva grows. These are not the same as the tentacles of the adult worm’s lophophore, which will be formed from a different structure when the larva undergoes metamorphosis.

As usual, a photograph doesn’t give a very satisfactory impression of the larva’s three-dimensional structure. There’s a lot going on in this little body! The entire surface is ciliated, and this actinotroch’s gut is full of phytoplankton cells. You can see a lot more in the video, although this larva is also a little squished.

I’ve been offering a cocktail of Dunaliella tertiolecta and Isochrysis galbana to the adult phoronids, and these are the green and golden cells churning around in the larva’s gut. However, good eaten is not necessarily food digested, and the poops that I saw the larvae excrete looked a lot like the food cells themselves. Today I collected more larvae from the parents’ bowl and offered them a few drops of Rhodomonas sp., a cryptonomad with red cells. This is the food that we fed actinotrochs in my class at Friday Harbor. We didn’t have enough time then to observe their long-term success or failure, but I did note that they appeared to eat the red cells.

I don’t know if phoronids reproduce year-round. It would be a simple task to run down and collect a few every month or so and see if any worms are brooding. Now that I know where they are, it would also be a good idea to keep an eye on the size of the patch. Some species of phoronid can clone themselves, although I don’t know if P. ijimai is one of them. In any case, even allowing for the possibility of clonal division, an increase in the size of the adult population would be at least partially due to recruitment of new individuals. If recruitment happens throughout the year, it follows logically that sexual reproduction is likewise a year-round activity. Doesn’t that sound like a nifty little project?

Besides, it’s never a bad idea to spend time at the harbor!

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A different take on ‘vermiform’

Posted on 2017-09-202023-01-06 by Allison J. Gong

If I asked you to draw a worm and designate the front and back ends, you’d most likely come up with something that looks like this:

And you would be entirely correct. A worm, or any creature described as ‘vermiform’ for that matter, has an elongated, wormlike body. Some worms have actual heads with eyes and sensory tentacles, but many don’t. The great many polychaete worms that live in tubes don’t have much of a head at all: usually all you can see sticking out of the tube is a crown of tentacles used for feeding. Although even the use of the word ‘crown’ more than suggests the presence of a head, doesn’t it? After all, where else does one wear a crown?

Polychaete worms, Phragmatopoma californica, sticking their ‘heads’ out of their tubes at Natural Bridges
26 May 2016
© Allison J. Gong

Most worms, including the worm that we imagined above, are bilaterally symmetrical, with bodies elongated along the Anterior-Posterior axis. This means the head is at the anterior end and the rear is the posterior end. For animals that don’t have a prominent head, the Anterior can also be defined by the direction of locomotion. Worms crawl with their bellies against the ground, which sets up a second axis of symmetry, the Dorsal-Ventral axis. The third axis of symmetry is the Left-Right axis. These axes should sound familiar, because they apply to our own bodies, as well of those of all other vertebrates and many invertebrates. Because of our upright stance we actually walk with our ventral surface forward, which is a little confusing, but if you don’t trust me you can see for yourself by crawling around on hands and knees for a while.

Now back to our worms, hypothetical and otherwise. Consider a worm that is elongated not along its Anterior-Posterior axis, but along its Dorsal-Ventral axis. It sounds strange, but such worms do exist. They are called phoronid worms, and are classified within their own phylum, the Phoronida. They all live in tubes, and the few times I’ve seen them they have been in pretty dense aggregations. As with most tube-dwelling worms the only part of the body that you can usually see is the crown of feeding tentacles, which in these animals (as well as in the Bryozoa and Brachiopoda) is called a lophophore.

The other day I was at the harbor looking for slugs with my friend Brenna, and spotted these pale tentacles swaying in the current.

Phoronids at the Santa Cruz Yacht Harbor
18 September 2017
© Allison J. Gong

These are the lophophores of an aggregation of phoronids! I’d never seen them at the harbor before, so I was pretty excited about it. They were on the side of a floating walkway, down almost beyond the reach of my outstretched arm. The current caused the lophophores to sway continuously and I was barely able to snap some blurry photos without falling in (I couldn’t really see what I was doing and just hoped for the best) when I accidentally caught this one shot. I wanted to have at least one clear-ish shot to submit to iNaturalist. I did manage to scrape off some bits of stuff that I hoped contained intact phoronids, so I could observe them under the dissecting scope at the lab.

And these are some lovely little worms!

The tubes that these phoronids inhabit are more like burrows of slime to which the surrounding sediments adhere. The tube itself isn’t anything particularly interesting, but the bodies of the worms are beautifully transparent. One of the coolest things you can see in a living phoronid is its circulatory system. They have red blood that, like ours, contains hemoglobin, so it’s easy to see the vessels that run along the length of the worm (which is the Dorsal-Ventral axis, remember) and the two blood rings around the base of the lophophore. If you get the lighting right you can even see the vessels that extend into each tentacle of the lophophore.

Single phoronid worm extending its lophophore
18 September 2017

I was disappointed to see that none of the video clips I took really do justice to these worms. They are so pretty when I look at them through the microscope, and I wish I could capture their beauty. You may at least be able to see blood moving through the larger vessels of the body in this short video.

Seems I need to upgrade my photomicroscopy set-up. Anybody have a few thousand bucks they want to donate to the cause?

I’m keeping the phoronids for as long as I can, although I don’t know what to feed them. I had time to take just a quick look at them this morning, and they look fine. Just for kicks I offered them a little phytoplankton to see what they’d do with it and couldn’t see if they were reacting at all. Still, they are filter feeders, and if I can adjust the lighting and get a good view of those ciliated tentacles I should be able to see if they are creating a water current that is bringing food to the mouth. Friday is the next day I have time to spend with these animals that I don’t get to see very often. Maybe then I’ll have something else to report.

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Lunacies

Posted on 2017-09-072023-01-06 by Allison J. Gong

For several centuries now, Earth’s only natural satellite has been associated with odd or unusual behavior. Lunatics were people we would describe today as mentally ill, who behaved in ways that couldn’t be predicted and might be dangerous. The erratic behaviors were attributed to the vague condition of lunacy. These words are derived from the Latin luna, which means ‘moon’. The cycles of the moon have long been thought to influence human behavior as well; hence such legends as the werewolf.

We do know that the moon indeed has a very strong influence on aspects of many organisms, primarily through the tides. For example, reproduction in many marine animals is timed to coincide with a particular point in the tidal cycle. Grunion (Leuresthes tenuis, small, silver, finger-shaped fishes) run themselves up onto California beaches at night to spawn following the full and new moon high tides in the early summer months. Corals in the Great Barrier Reef spawn together in the handful of nights after the full moon in November. Animals such as these, which reproduce via broadcast spawning, are the ones most likely to benefit from synchronized spawning; after all, there is no point in spawning if you’re the only one doing it. Invertebrates don’t have watches or calendars; they keep time by sensing the natural cycles of sun and moon. The moon’s strong effect on the tides is a signal that all marine creatures can sense and use to coordinate spawning, increasing the probability of successful fertilization for all.

Last night, Wednesday 6 September 2017, the moon was full. Yesterday at the lab, I noticed that  the large Anthopleura sola anemones living in the corner of my table had spawned.

A male Anthopleura sola anemone that had spawned
6 September 2017
© Allison J. Gong

That diffuse grayish stuff in the right-hand side of the photo is a pile of sperm. I looked at a sample under the microscope, just to be sure. By this time they had been sitting at the bottom of the table for several hours and most of them were dead. But they were definitely sperm:

Whenever I see something unusual like this my first impulse is to see if it’s happening anywhere else at the lab. So I started poking around. The aquarists at the Seymour Center told me that some of their big anemones had spawned in the past couple of days; however, since they clean and vacuum the tanks every day all evidence was long gone.

Fortunately there are several A. sola anemones in other labs that aren’t cleaned as regularly as the public viewing areas. One of the animals in the lab next door to where I have my table had also spawned. . .

Female Anthopleura sola
6 September 2017
© Allison J. Gong

. . . and this one is a female! What looks like a pile of fine dust is actually a pile of eggs.

Eggs of Anthopleura sola
6 September 2017
© Allison J. Gong

And the eggs are really cool. See those spines? They are called cytospines and apparently deter predation. Other species in the genus Anthopleura (A. elegantissima and A. xanthogrammica) are known to have spiny eggs, so it appears that this is a shared feature. Now, if only I could get my hands on eggs of the fourth congeneric species–A. artemisia, the moonglow anemone–that occurs in our area, I’d know for certain, at least for California species. I examined the eggs under higher magnification, but due to their opacity I couldn’t tell if the had been fertilized. Most appeared to be solid single undivided cells; they could, however, be multicellular embryos.

All told, of the anemones that had obviously spawned, 1 was female and 4 were male. I sucked up some of the eggs and put them in a beaker of filtered seawater. I doubt that anything will happen, but I may be in for a pleasant surprise when I check on them tomorrow.

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The fluidity of sex

Posted on 2017-09-022023-01-06 by Allison J. Gong

We humans are accustomed to thinking of sexual function as being both fixed and segregated into bodies that we designate as either Female or Male. And while we, as a species, generally do things this way, in the larger animal kingdom sexual function doesn’t always follow these rules. Many animals are monoecious, or hermaphroditic, having both male and female sex organs in the same body. Not only that, but lots of animals change from one sex to the other. As in so many aspects of biology, the way humans do things may be thought of by us as “normal,” but it isn’t the most interesting way.

Take, for example, the slipper shell Crepidula adunca. This is a small limpet-like creature that lives on the shell of a larger snail. Around here the usual host is a turban snail, either Tegula funebralis or T. brunnea.

Slipper shell (Crepidula adunca) on its host, the turban snail Tegula brunnea, at Pigeon Point
1 May 2017
© Allison J. Gong

There are several species in the genus Crepidula, including C. fornicata, which lives on the Atlantic coast of North America. The species epithet gives an inkling of how reproduction occurs in at least these two species of the genus.

Sometimes C. adunca is found in stacks. I’ve never seen a stack taller than three individuals, but C. fornicata occurs in stacks of about six. The animal at the bottom of the stack is always the largest, and a given turban snail can play host to more than one stack at a time.

Two stacks of Crepidula adunca on the turban snail Tegula funebralis, at Pigeon Point
28 June 2017
© Allison J. Gong

As you might guess, it isn’t mere happenstance that these stacks of C. adunca occur. It turns out that this unusual living arrangement is key to both sexual function and eventual reproduction in this species. The individual on the bottom of the stack (i.e., the oldest) is always a female; those at the top of the stack (i.e., the youngest) are males. However, every stack begins with a single individual, and the default sex in newly settled C. adunca is male. An experiment conducted at Friday Harbor in Washington State1 showed the change from male to female began when the snails reached a size of 7 mm, and all animals larger than 10 mm were female. Animals that begin life as male and transform into females are described as protandrous hermaphrodites. How common is this phenomenon? Not uncommon among fishes, actually. Clownfishes in the genus Amphiprion are protandrous. Remember how in the beginning of the moving Finding Nemo, Nemo’s mom dies? Well, in real life Nemo’s dad would have become his new mom!

In any case, all C. adunca begin adult life as males. If they live long enough to reach about 7 mm in length, they might get to become females. Crepidula adunca‘s unusual living arrangement also facilitates reproduction. Unlike most limpet-like gastropods, C. adunca isn’t a broadcast spawner. Rather, it copulates, as hinted at by the species epithet of its congener C. fornicata. A female slipper shell with a male on her back has a convenient source of sperm with which to fertilize her eggs:  the male reaches into her mantle cavity and transfers sperm to her. Given the constraint of copulation, a female cannot mate until she carries at least one male on her back, and a male cannot reproduce unless he settles atop a female. Once the eggs have been fertilized, they develop within the mother’s mantle cavity until she pushes them out as little miniatures of herself.

Crepidula adunca on the turban snail Tegula brunnea, at Davenport Landing
27 May 2017
© Allison J. Gong

Cool little animals, aren’t they? They remind us not to think of ourselves as The Way Things Are Done. We have a lot to learn from creatures that are not like us, and it’s stories like these that ensure I will never lose my appreciation and love for the marine invertebrates.


1 Collin, R. 2000. Sex Change, Reproduction, and Development of Crepidula adunca and Crepidula lingulata (Gastropoda: Calyptraeidae). The Veliger 43(l):24-33.

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