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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!

 

This week's field trip for my Ecology class was the first of two visits to the Santa Cruz harbor. The students' task was to select a site to monitor for a semester-long study of ecological succession. The floating docks at the harbor are the ideal site for this kind of study because I know from experience that the biota changes from season to season throughout the year, on a time scale that can be observed within the confines of a 16-week semester. We will return to the harbor in nine weeks and students will document how their sites have changed in that time.

California is swinging back into the severe drought situation we had before the epic 2016-2017 rainy season. Since the current rainy season began on 1 October 2017, we've had hardly any rain at all and very little snow in the Sierra. Fools who thought that one rainy season would get us out of drought are just that--fools. However, one nice thing about drought conditions is that visibility at the harbor is pretty good. Without any significant runoff the water is nice and clear, making it easy for the students to see what's growing on their section of the docks.

Students examining their study plot
9 February 2018
© Allison J. Gong
Sometimes a little ballast is required!
9 February 2018
© Allison J. Gong

The assignment for this first visit to the harbor was to choose a site, identify what lives on the site, and draw a map of it. I had warned them that all the interesting biology on the docks occurs below the level of their feet, and that they would have to lie or kneel on the dock to get a good look at what's going on down there. Some of them tried to take a photo of the entire site, but it's impossible to get far enough away. Unless you're actually in the water, from where it would be easy. Yeah, you could don a wetsuit and get in the water, but the harbor isn't the most ideal place to go for a morning swim.

A little back story on the docks at the Santa Cruz harbor

Remember the magnitude 9.0 earthquake and subsequent tsunami that occurred in northern Japan several years ago? That was on 11 March 2011 at 14:46 local time. That morning in Santa Cruz we received a tsunami warning. I didn't venture down to the harbor (I think I was working at the marine lab that day) but here's a video shot by a woman who watched the ~0.5 meter tsunami tear through the upper harbor:

Amazing, the destructive power of such a small wave, isn't it? Boats were wrenched from their moorings and slammed into other boats and harbor infrastructure. I forget the total dollar amount of damage that our harbor sustained, but as a result all of the docks were replaced in the next few years. I did happen to be at the harbor with a group of students on one of the days that the old docks were being removed. It was heartbreaking to see the docks, carrying decades of biological growth on them, dumped in the parking lot to dry out in the afternoon sun. I imagine they were eventually hauled out to the landfill. 

Since then, the biota on the new floating docks seems finally to be stabilizing. If I had been teaching Ecology back in 2013, we would have had pristine habitat in which to observe honest-to-goodness primary succession. As things are, however, I'm giving students the option of scraping all or part of their plot clear, to simulate primary succession. Their other option is to leave the plot as-is, and pick up the succession process somewhere in the middle and see what happens from this point forward.

So, what did they see down there? 

Well, even though the water was relatively clear, a lot of the photos looked like this:

9 February 2018
© Allison J. Gong

I can identify much of the stuff in this photo, but this isn't the best shot to showcase the biodiversity on the docks. I decided that the camera would do a better job if I used it to photograph individual organisms instead. Here are some of my favorites.

This shot is looking straight down along the edge of one of the docks. The macroscopic life begins 2-3 cm below the waterline, and even above that the dock surface is covered with microscopic scuzzes.

White plumose anemones (Metridium senile) at the Santa Cruz harbor
9 February 2018
© Allison J. Gong
Oral view of white plumose anemones (Metridium senile) at the Santa Cruz harbor
9 February 2018
© Allison J. Gong

I had shown the students pictures of organisms they would be likely to see at the harbor. One of the critters that shows up sporadically is the introduced hydroid Ectopleura crocea. Later in the semester we will discuss species introductions and invasions in more detail. Harbors generally tend to be heavily populated by non-native species, and our local harbor is no exception. The species of Ectopleura found in harbors has hydranths that can be 8-10 cm long, and when it occurs it tends to be quite conspicuous. The congeneric species, E. marina, lives in intertidal in some areas on the open coast; I've seen it in a few tidepools at Davenport Landing, for example. The intertidal species is much smaller, about 2-3 cm tall and doesn't form the dense clumps that typifies E. crocea.

The non-native hydroid, Ectopleura crocea, at the Santa Cruz harbor
9 February 2018
© Allison J. Gong
Caprellid amphipods at the Santa Cruz harbor
9 February 2018
© Allison J. Gong

The ubiquitous caprellid amphipods were crawling all over everything, as usual. Some of the students really didn't like these guys and one of them had the same reaction to them that I do, which is a general shudder. They're sort of cute in still photos, but when they start inchworming around they look sort of creepy. And when there's a bunch of them writhing around in an oozy mass, they're REALLY creepy.

 

 

 

One of the most conspicuous worms at the harbor is Eudistylia polymorphora, the so-called feather duster worm. They come in oranges, purples, and yellows. This one was pure white. Lovely animal!

Feather duster worm (Eudistylia polymorpha) at the Santa Cruz harbor
9 February 2018
© Allison J. Gong

Tube-dwelling polychaete worms, such as Eudistylia, don't have much in the way of a head but they do have many light-sensitive eyespots on the tentacles. They react very quickly to many stimuli, and even a shadow passing over a worm causes it to yank its tentacles into its tube in the blink of an eye. Usually they're not too shy, though, and will extend their tentacles soon to resume feeding.

All told we were on the docks for about 2.5 hours. Not a bad way to spend a glorious morning, is it?

9 February 2018
© 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.

 

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.

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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