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Category: Marine invertebrates

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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Now you see it, now you don’t

Posted on 2017-07-112023-01-06 by Allison J. Gong

This morning in the intertidal I was reminded of how often I encounter animals I wasn’t looking for and almost missed seeing at all. That got me thinking about color and pattern in the intertidal, and how they can be used either to be seen or to avoid being seen. Some critters–the nudibranchs immediately come to mind–are so brightly colored that they are impossible to miss, while others are camouflaged to the point that it takes a trained eye to see them.

Truth be told, however, most of the animals in the intertidal don’t have eyes, or at least eyes that can form images the way ours do. While just about any animal might be preyed upon by birds at low tide, most of the predators a creature of the tidepools might face would not be visual predators. This in turn begs the question of just how adaptive or not a species’ crypticness is. The way I see it, there are three options, or hypotheses about the potential benefit of an animal’s coloration and patterning:

  1. Colors and patterns that make an animal conspicuous are advantageous.
  2. Colors and patterns that make an animal cryptic or camouflaged are advantageous.
  3. Colors and patterns are neither advantageous or disadvantageous.

Today I’m going to consider hypothesis #2, as it is the most interesting one. Let’s put aside for now the question of how an animal’s color comes to be and consider only its effect on visibility to Homo sapiens (specifically, me).

Example #1 (obvious): Tonicella chitons

These are the pink chitons that I find on exposed coasts. They eat encrusting coralline algae, and I suspect their color derives at least in part from their diet. Here’s one that perfectly matches its food:

The chiton Tonicella lokii at Pistachio Beach
29 May 2017
© Allison J. Gong

On the other hand, Tonicella isn’t always this entirely pink, nor is it always seen on coralline algae:

The chiton Tonicella lokii at Pistachio Beach
29 May 2017
© Allison J. Gong
The chiton Tonicella lokii at Monastery Beach
27 October 2015
© Allison J. Gong

The chiton I saw at Monastery Beach wasn’t anywhere near coralline algae. It has obviously been eating something, probably algal films of whatever sort it comes across. Correlation is not causation, but it may not be mere coincidence that this pale version of Tonicella lokii lives on rock devoid of coralline algae.

Example #2 (obvious): Decorator crabs

Tonicella doesn’t intentionally alter its appearance by eating pink food. Given the extremely rudimentary nature of a chiton’s nervous system, it likely can’t intentionally do much of anything. It doesn’t have eyes so it cannot see, although there are light-sensing organs called aesthetes in the dorsal shell plates and light-sensitive cells in the lateral girdle. Chitons make their way through the world largely by following chemical gradients, either in the water current or on the substrate.

Crabs, on the other hand, have very complex compound eyes and can, to some extent, see what’s going on around them. The compound eyes of arthropods are highly effective motion sensors, certainly much more sensitive than our eyes are, which is why it’s so hard to sneak up on a fly even if you’re extending your reach by using a fly swatter. Crabs certainly are aware of the visual aspects of their surroundings. They can see potential threats and typically respond in one of three ways: (1) scuttling away; (2) coming out fighting; and (3) remaining still and trying not to be noticed.

It takes energy to scuttle back and forth, and the little shore crabs (Pachygrapsus crassipes) are always on the move. They are quick to run for cover when approached, but will come out and resume their explorations if you sit still for about a minute. They are really fast and difficult to catch, perhaps not quite as challenging as the Sally Lightfoot crabs that so enraged the crew of the Western Flyer during Ed Ricketts’ and John Steinbeck’s excursion to the Sea of Cortez, but hard enough to be not worth my effort. Fighting is an option only for those equipped to fight. Rock crabs (for example, Romaleon antennarium) remain hidden under algae or partially buried in sand, but when exposed they come out with big claws open and ready to pinch the hell out of anything that comes close. These are the only animals that I really worry could hurt me in the intertidal.

Which leaves the hold-still-and-hope-not-to-be-seen option. This is what decorator crabs do. In terms of temperament, decorator crabs (of which there are several species) are placid and unaggressive: they will pinch when provoked and it can hurt, but they won’t do the kind of damage that a rock crab would happily inflict. Decorator crabs hide in plain sight by covering their carapace and legs with little bits of the environment, usually algae. A well-decorated crab can be sporting several species of algae on its back.

This morning I saw and collected this small crab:

A small decorator crab, Pugettia richii, on a bed of Egregia menziesii at Davenport Landing
11 July 2017
© Allison J. Gong

I actually didn’t see it at first. I was pawing through the thick algal growth and felt its little feet scratching my hand. I peeked under the algae and there was the crab. Its carapace is about 2.5 cm across, and its claws probably wouldn’t be able to pinch human skin even if the crab tried to. Which it certainly didn’t. I wanted to observe the crab more closely in and keep it for use when I teach the crustacean diversity lab this fall, so I brought it back to be examined under the dissecting scope.

A decorated Pugettia richii, observed in the lab
11 July 201
© Allison J. Gong

The crab’s own color is a dark brownish red, which helps it hide amongst the red algae. It adds to the environment-as-appearance effect by attaching at least three species of red algae to its carapace. The crab does this by grabbing a piece of algae with one of its claws, then reaching up and behind its head to put it on the carapace, which has has tiny hooks that hang onto the decoration. It’s a very nifty scheme, but there’s one big problem. Each time the crab molts it loses its decoration and has to acquire its accessories all over again.

Example #3 (not obvious at all): Lottia digitalis

We have about a gazillion species of limpets on the California coast. Well, not really but it certainly does feel like it. To make things even more difficult I can’t seem to keep the current scientific names straight. I know that many of the commonly encountered intertidal limpets have been consolidated into the genus Lottia (this includes species that I learned by another name way back when) and I’m slowly getting used to recognizing the Lottia “look”. However, aside from the owl limpet (L. gigantea), which is much bigger and more conspicuous than any others, the other species are difficult to distinguish and I can never remember if species x has the deep ridges or if that’s species y. Ugh.

Earlier this spring I was in the field with my friend Brenna, and she was showing me the differences between Lottia scabra and L. digitalis. Brenna studies molluscs so I know she knows what she’s talking about. Lottia scabra is now easy for me to recognize, but L. digitalis is both trickier and more interesting.

Limpets Lottia scabra (upper right) and L. digitalis (left and lower right) among barnacles at Natural Bridges
25 June 2017
© Allison J. Gong

See how those all look like limpets? Now look at this:

Davenport Landing
11 July 2017
© Allison J. Gong

Do you even see the limpets?

The large animals in the photo are gooseneck barnacles, Pollicipes polymerus. They live on and amongst mussels in the mid-intertidal. This spring Brenna told me that Lottia digitalis comes in a morph that lives on and looks like Pollicipes. I’d never seen it until today. Look at the photo again. Can you see the limpets now?

Here are some more photos.

Lottia digitalis (“Pollicipes morph”) at Natural Bridges
11 July 2017
© Allison J. Gong
Lottia digitalis (“Pollicipes morph”) at Natural Bridges
11 July 2017
© Allison J. Gong

Isn’t it remarkable how these limpets have exactly the colors and pattern as the plates of Pollicipes? And I didn’t even know about them six months ago. I love having new things to learn and more reasons to pay closer attention to creatures I tend to take for granted. I think it’s time for me to tackle the challenge of identifying limpets in the field. Next season, that is. Today was probably my last day in the intertidal for a few months. We won’t have decent low tides during daylight hours until November.

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Peanut worms!

Posted on 2017-05-232023-01-06 by Allison J. Gong

If I ask my invertebrate zoology students to name three characteristics of the Phylum Annelida, they would dutifully include segmentation and chaetae (bristles) in the list. And they would be correct. Annelids, for the most part, are segmented and many of them have chaetae. But in biology there are many exceptions for every rule we teach, and it’s these exceptions that make a deeper study of biology so rewarding.

Peanut worms (Phascolosoma agassizii) at Pigeon Point
30 April 2017
© Allison J. Gong

A couple of weeks ago I did some collecting in the intertidal at Pigeon Point. It was a very accommodating low tide, and I had a lot of time to poke around and explore. I found an area that had several decently sized rocks that I could turn over, and had fun seeing what lives on the side away from the light. Some of the animals on the underside of rocks are the common ones you see everywhere–turban snails, limpets, Leptasterias stars, and the like. Some, however, prefer a life of darkness and actively move away from the sun when their rock is turned over. And others happen to live in the sand under the rock and might not care one way or the other about the light.

Peanut worms, scientifically known as sipunculans, are delightful small worms that in my opinion are vastly underappreciated. This is understandable, as they are usually hidden in sand or rubble and aren’t exactly conspicuous even when uncovered. Phascolosoma agassizii is our local sipunculan. Like all sipunculans it is unsegmented, and it has no chaetae. Peanut worms used to be elevated to their own phylum, the Phylum Sipuncula; however, molecular evidence shows that they are indeed annelids despite their apparent loss of key features such as body segmentation and chaetae.

Peanut worms (Phascolosoma agassizii) at Pigeon Point
30 April 2017
© Allison J. Gong

They do look vaguely peanut-ish, don’t they? They’re small, maybe 6 cm all stretched out, which you hardly ever see. Phascolosoma agassizii is a grayish pink color, with irregular black stripes that usually don’t form complete hoops around the body. Peanut worms are sedentary, living with most of the body buried in sand, rubble, shell debris, kelp holdfasts, etc. One of the weird things about them is that the mouth in located on the distal end of a long tube called the introvert. Most of the time the introvert is stuffed inside the main body region, or trunk. It is eversible and unrolls from the inside out, sort of like when you remove a long sock by pulling the top edge down over your leg and off your foot. The mouth on the end of the introvert is surrounded by short sticky tentacles, and the introvert dabs around to pick up organic deposits from the surfaces. Mucus and cilia on the tentacles convey the yummy organic gunk to the mouth, and a pharynx pushes food through to a long esophagus that runs the length of the introvert and leads to the long coiled intestine in the trunk.

Watch these peanut worms extending and retracting their introverts. Cute, aren’t they?

I brought three peanut worms back to the lab with me, where they are happily living in my sand tank. Their housemates are ~15 sand crabs (Emerita analoga) and a clump of tube-dwelling polychaetes (Phragmatopoma californica). I never see them unless I dig them up from the sand, which leads me to believe that they do most of their feeding at night. Either that or they actually do actively shy away from the light.

Despite not sharing much in the way of apparent morphological similarity with more typical annelids, sipunculans are indeed annelid-like in other ways. Many of their internal structures are like those of annelids, and at least their early development (cleavage pattern and differentiation of tissue layers) follows the annelidan pathway. The species that have indirect development have a trochophore larva, typical of the marine annelids, that in some cases morphs into a second larval stage called a pelagosphera.

Sipunculans are the poster child for Animals That Are Not What They Seem. But they are interesting in their own way, and I always have a “yay!” moment when I find them in the field. It’s really hard not to make sound effects as they’re rolling their introverts in and out. You should try it yourself some time.

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Eight is enough

Posted on 2017-05-162023-01-06 by Allison J. Gong

One of the defining characteristics of the Phylum Mollusca is the possession of a shell, which serves both as a protective covering and an exoskeleton. We’ve all seen snails, and some people may have noticed that snails often withdraw entirely into their shells and even have a little door that they can use to seal up the opening of the shell. That little door is called the operculum. Opercula occur in non-molluscan animals, too, such as some of the tube-dwelling polychaete worms and some of the thecate hydroids. Snail opercula come in lots of different shapes, depending on the aperture of their owner’s shell.

Calliostoma ligatum at Mitchell’s Cove
2 April 2017
© Allison J. Gong

Given the enormous morphological diversity within the Mollusca it shouldn’t be surprising that their shells vary immensely in prominence and shape. In fact, molluscan shells demonstrate quite beautifully the relationship between form and function. The benthic and most familiar molluscs, the gastropod snails, generally have coiled shells. Notable exceptions to this generality are the marine opisthobranchs (nudibranchs and sea hares) and the terrestrial slugs. And for the most part snail shells look recognizably like snail shells, even though some are plain coils, others may be flattened (e.g., abalones), and still others may be crazily ornamented. Aquatic animals crawl around in water, which helps to support the weight of heavily calcified shells. Terrestrial snails, on the other hand, live in a much less dense medium (air) and have lighter, less calcified shells. The trade-off for a more easily transportable shell is that air is also very drying, and a thinner shell provides less protection from desiccation.

I should state for the record right now that I’m not talking about the many molluscs that don’t have shells at all, or that have much reduced shells.

View into the exhalant opening of a mussel (Mytilus sp.) at the Santa Cruz Yacht Harbor
29 August 2015
© Allison J. Gong

The bivalve molluscs (mussels, clams, oysters, etc.) live inside a pair of shells. They are sedentary animals, living either attached to a hard surface or buried in sand or mud. Not being able to run from predators (although some scallops can swim!), their only defense is the toughness of their shells and the strength of the adductor muscles that hold the shells closed. Most bivalves feed by sucking water into the shells through an incurrent siphon, using their gills to filter food particles from the water, and expelling the water through an excurrent siphon. To do so they must open their shells enough to extend their siphons, or at least expose inhalant and exhalant openings, to the water current surrounding them.

So, snails have one shell and bivalves have two. Some of the most interesting molluscs, in terms of shell morphology, are the chitons. The Polyplacophora (Gk: ‘many plate bearer’) have a shell that is divided into eight dorsal plates. This makes them immediately distinguishable from just about any other animal.

Chiton (Lepidozona mertensii) at Point Piños
6 February 2016
© Allison J. Gong

Chitons live from shallow water to the deep sea, but the majority of species live in the intertidal. This is a high-energy habit characterized by the bashing of waves as the tide rises and falls twice daily. Any organism living here must be able to hang on for dear life or risk being swept away to certain death. Chitons are certainly well equipped to survive in this habitat. They have a low profile, offering minimal resistance to the waves. Rather than stand tall and face the brunt of the wave energy, chitons cling tightly to the rocks and let the waves wash over them.

Tonicella lokii at Monastery Beach
27 October 2015
© Allison J. Gong

The chiton’s shell, divided into eight articulating plates, gives the animal a much more flexible shell than is found in any other mollusc. This allows them to conform to the topography of the rocks, giving them an even lower profile than, say, a limpet of the same overall shape and size.

While most chitons are pretty sedentary, at least during the low tides when we can see them, some of them can move pretty quickly when they want. So what, exactly, motivates a chiton to run? One species, Stenoplax heathiana, lives on the underside of rocks in the intertidal; it comes out at night to forage on algal films and retreats back under its rock with the dawn. I’ve seen them at Pistachio Beach, where I turned over rocks and watched them run away from the light. This video is shot in real-time; the chitons are really running fast!

When the eight shell plates are visible it’s easy to identify a chiton as a chiton. But not all chitons are quite so obliging with their most chiton-ish characteristic, and one is downright misleading.

Below is Katharina tunicata, one of the largest chitons on our coast. Its shell plates are barely visible, as they are almost entirely covered by the animal’s mantle, the layer of tissue that covers the visceral mass and encloses an open space called the mantle cavity in which the gills are located. In chitons, the mantle extends onto the dorsal side of the animal and is called the girdle. Katharina‘s girdle is smooth and feels like wet leather.

The chiton Katharina tunicata at the Great Tidepool in Pacific Grove
26 October 2015
© Allison J. Gong

The largest chiton in the world is the gumboot chiton, Cryptochiton stelleri, and it lives on our coast. This beast is about the size of a football, reaching a length of 30 cm or so. It lives mostly in subtidal kelp forests, but can be found in the very low intertidal, which is where I usually see it. At first encounter it’s hard to figure out what this animal is. It certainly doesn’t look like a chiton.

A large Cryptochiton stelleri at Mitchell’s Cove
6 June 2016
© Allison J. Gong

If anything, it looks like a mostly deflated football, doesn’t it? Turning it over to look at the underside doesn’t help much, either, although this photo does give an idea of how big the animal can get:

Ventral view of Cryptochiton stelleri at Pigeon Point
24 April 2016
© Allison J. Gong

Cryptochiton goes one beyond Katharina and covers its plates entirely. Just looking at the animal you’d have no idea that there are eight plates underneath the tough reddish-brown mantle, but you can feel them if you run your finger along the midline of the dorsum. Living subtidally as it does, Cryptochiton doesn’t have the ability to cling tightly to rocks that its intertidal relatives do, and it tends to get washed off its substrate and cast onto the beach during storms. I’ve never seen one on the beach that wasn’t very dead. Once a friend and I were trudging back up the beach after working a low tide, and encountered a dead softball-sized Cryptochiton. I mentioned that it would be nice to have a complete set of shell plates from one of these animals. My friend always carries a knife in her pocket, so we started an impromptu dissection right there on the beach. It didn’t take long to learn that the mantle of a gumboot chiton is really tough and difficult to cut through with a pocket knife. And even once we got through the mantle, dissecting the plates from the underlying tissue wasn’t going to happen with the tools we had with us. Besides, the stench was godawful even with our unusual tolerance for the smell of dead sea things. We abandoned that corpse.

Single plate of the gumboot chiton, Cryptochiton stelleri
16 May 2017
© Allison J. Gong

Many beachcombers have found white butterfly-shaped objects in the sand, but not known what they are. They are definitely calcareous and feel like bone, but what kind of animal makes a bone shaped like this? Turns out this object is one of the shell plates from C. stelleri. They wash up frequently, never attached to their neighbors so they provide no clue as to what organism they came from.

In order to obtain a complete set of Cryptochiton plates, I’d have to start with an intact chiton corpse. I did happen upon another dead Cryptochiton on a beach somewhere I was allowed to collect organisms, and I brought it back to the marine lab. I remember spending a smelly afternoon cutting the plates out of the corpse and removing as much of the tissue as I could, then feeding the plates to various hungry anemones to take care of the rest. Some of the plates got a little broken during the extraction process, but I do have my very own full set!

Shell plates of Cryptochiton stelleri
16 May 2017
© Allison J. Gong

Some day I will figure out a way to mount those plates permanently.

One final question to ponder. Does a chiton have one shell, or eight shells?

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Friends in strange places

Posted on 2017-05-092023-01-06 by Allison J. Gong

Animal associations can be strange and fascinating things. We’re used to thinking about inter-specific relationships that are either demonstrably good or bad. Bees and flowering plants–good. Mosquitos on their vertebrate hosts–bad. In many cases the ‘goodness’ or ‘badness’ of these associations is pretty clear. However, there are cases of intimate relationships between animals of different species that cannot be easily categorized as good or bad.

Take, for example, the barnacles on the skin of gray and humpback whales. From the barnacles’ perspective the skin of a whale isn’t a bad place to live: as the whale swims through the water the barnacle is continually flushed by clean water, which should make feeding easier. But is the whale affected in any way by its barnacle passengers? I suppose they might increase the drag coefficient a little bit and make swimming marginally less efficient, and maybe they itch, although it’s hard to imagine that the whale would really care much one way or the other.

A week ago I went to the intertidal up at Pigeon Point. It’s a great spot for certain animals, especially the small six-rayed stars of the genus Leptasterias. These stars rarely get larger than 8 cm in diameter and always have six arms. I’ve been told by a friend who just happens to be a sea star taxonomist at the Smithsonian, that making species identifications in the field is very difficult for this genus, so I’ve stopped trying. I do know that some of the Leptasterias stars have slender rays and others have thicker rays.

Two stars of the genus Leptasterias, at Pigeon Point
9 May 2016
© Allison J. Gong

The most common large star at Pigeon Point is the bat star, Patiria miniata. These stars get about as big as my outstretched hand, and come in a variety of colors. Last week I didn’t see very many Patiria, but all of them were reddish orange, like this one:

Bat star (Patiria miniata) at Pigeon Point
30 April 2017
© Allison J. Gong

Unless they’re so abundant as to be annoying, I like picking up bat stars and looking at their underside. That’s because sometimes they have these little dark squiggles in their ambulacral groove:

Patiria miniata with commensal worm, at Pigeon Point
30 April 2017
© Allison J. Gong

That little squiggle is a polychaete worm, Oxydromus pugettensis. It is one of many polychaete worms that forms a symbiotic relationship with another animal species. Some symbiotic polychaetes live in the tubes of other worms, or within the shells of bivalves, for example. Oxydromus crawls around inside the ambulacral groove of Patiria, where it feeds on scraps of leftover food from the star’s meals. The worms don’t like light, and as soon as I picked up this star and flipped it over the worm started burrowing down between the star’s tube feet to get back to the dark. The next day I found another star with a worm and was able to take a picture of it before it disappeared.

Commensal worm (Oxydromus pugettensis) in the ambulacral groove of Patiria miniata, at Pigeon Point
1 May 2017
© Allison J. Gong

Oxydromus pugettensis is clearly segmented, evidence of its annelidan roots. It doesn’t look very different from many other free-crawling polychaetes. A member of the family Hesionidae, it lives in fine silty sediments in the intertidal as well as in the ambulacral grooves of sea stars. According to one source, it is the most common intertidal member of its family along the California and Oregon coast. For reasons as yet undetermined, P. miniata seems to be the favored host, although I have also seen the worms in the ambulacral grooves of the leather star Dermasterias imbricata.

Over two days at Pigeon Point last week I examined a total of five bat stars, and all of them had worms. One of the stars had three worms! It’s possible that more worms were hiding deep within the ambulacral grooves, too. I always wonder how, in this type of association, the partners manage to find each other. How does one “lucky” star end up with three worms? Do the worms every migrate from one star to another? Does the star do anything to attract the worms? In what way(s) would the star benefit from having a few worms in its ambulacral regions? It does seem that the worms don’t stick around very long once a star is brought into the lab–I don’t know if they die or just leave on their own–but since they also live in the sand maybe they do actively migrate between stars. There hasn’t been much work done on these worms in recent decades, probably because of the overall decline in natural history studies. However, I’ll keep this worm in mind for my Marine Invertebrate Zoology students this fall, when one of them asks me for help coming up with an idea for his or her independent research project.

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