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Today was the first time I've gone out on a low tide since before the whole COVID19 shelter-in-place mandates began. Looking back at my records, which I hadn't done until today because it was much too depressing, I saw that my last time out was 22 February, when the low tides were in the afternoon. At the time I made what seemed to be the not-too-bad decision to stay away from the remaining afternoon lows and wait until the spring shift to morning lows, which I like much more. And then then COVID hit and we all had to stay home and beaches were closed. So yeah, it has been much too long and I really needed this morning's short visit to the intertidal.

Pair of black oystercatchers (Haematopous bachmani) at Mitchell's Cove
My companions for a short while this morning, a pair of black oystercatchers (Haematopus bachmani) at Mitchell's Cove
2020-05-08
© Allison J. Gong

Beaches in Santa Cruz County are closed between the hours of 11:00 and 17:00, except that we are allowed to cross the beach to get to the water. This means that surfers, kayakers, SUP-ers, and marine biologists can get out and do their thing. Of course, my particular thing took place hours before the beach restrictions began, so I was in the clear anyway. I didn't venture too far from home, as I wasn't quite certain how easy it would be to get down to the beach.

Spring is the prime recruitment season for life in the intertidal. The algae are coming back from their winter dormancy, and areas that had been scraped clean by sand scour or winter storms are being recolonized. Many of the invertebrates have or will soon be spawning. And larvae that have spent weeks or even months in the plankton are returning to the shore to metamorphose and begin life as an adult. Just as it is on land, spring is the time for life in the sea to go forth and multiply.

For several decades now, marine ecologists have been studying barnacles and barnacle recruitment. Barnacles are a nice system for studying, for example, recruitment patterns and mortality. The cyprid larva, the larval stage whose job it is to find a permanent home in the intertidal, readily settles and metamorphoses on a variety of man-made surfaces; this makes it easy to put out plates or tiles and monitor who lands there. The fact that barnacles, once metamorphosed, remain attached to the same place for their entire lives means an ecologist can measure mortality (or survivorship, which is the inverse) by counting the barnacles every so often.

These are young barnacles (Chthamalus sp.), about 4-5 mm in diameter. I don't know how old they are, but would guess that they recruited in the past couple of months. These individuals all found a nice place to set up, because as I've written before, barnacles need to be in close proximity to conspecifics in order to mate.

Young acorn barnacles (Chthamalus dalli/fissus) on a rock at Mitchell's Cove
Small barnacles (Chthamalus sp.) on a rock at Mitchell's Cove
2020-05-08
© Allison J. Gong

This is a mixed group of Chthamalus sp. and Balanus glandula. Balanus is taller and has straighter sides and a more volcano-like appearance. Larvae of both genera recruit to the same places on rocks in the intertidal, and it is not uncommon to see assemblages like this.

Mixed assemblage of Balanus and Chthamalus barnacles at Mitchell's Cove
Barnacles Balanus glandula and Chthamalus sp. at Mitchell's Cove
2020-05-08
© Allison J. Gong

Both species of barnacles are preyed upon by birds, sea stars, and snails. Predatory snails use their radula to drill a hole through the barnacle's plates and then suck out the body. Some of the barnacles in the photo below are dead--see the empty holes? Those are barnacles that were eaten by snails such as these.

Small barnacles and predatory snails at Mitchell's Cove
2020-05-08
© Allison J. Gong

What was unusual about this morning was the number of snails of the genus Acanthinucella. I don't know that I've ever seen this many of them before.

Large group of Acanthinucella snails at Mitchell's Cove
2020-05-08
© Allison J. Gong

Lots of Acanthinucella means that lots of barnacles are being eaten. And empty (i.e., dead) barnacle tests are more easily dislodged from the rock than live ones are. A lot of dead barnacles could result in bare patches. And guess what? That's what I saw this morning!

Bare patches in barnacle population
Bare patches in barnacle population at Mitchell's Cove
2020-05-08
© Allison J. Gong

And those aren't just empty spaces where nobody settled. Notice the clean edges. These empty spaces formed because barnacles were there, but died recently and fell off. The abundance of Acanthinucella may have indirectly caused these patches to form--by eating barnacles and weakening the physical structure of the population. Bare space is real estate that can be colonized by new residents. See?

Newly settled barnacles
2020-05-08
© Allison J. Gong

These brand new recruits are about 1 mm in diameter. No doubt more will arrive in the coming months, and this patch will fill up with barnacles again. Vacant space is a limited resource in the rocky intertidal, and the demise of one generation provides opportunity for new recruits. And if the barnacles themselves don't occupy all of the space, then other animals and algae will. That's one of the things I love about the intertidal--it is a very dynamic habitat, and every visit brings something new to light. No wonder I missed it so much!

4

I'm willing to bet that when you think about coral, what comes to mind is something like this:

Great Barrier Reef
A Blue Starfish (Linckia laevigata) resting on hard Acropora coral. Lighthouse, Ribbon Reefs, Great Barrier Reef
© 2004 Richard Ling

The reef-building corals of the tropics are indeed spectacular structures, incredibly rich in biodiversity and worthy of a visit if you ever get the chance. These coral colonies come in many shapes, as you can see in the photo above. Each colony consists of hundreds or thousands of tiny polyps, all connected by a shared gastrovascular cavity, or gut. The living polyps secrete a skeleton of CaCO3, which grows slowly over decades or even millennia as successive generations of polyps live their lives and then die. It's this slow accumulation of CaCO3 that makes up the physical structure of the reef.

Reef-building corals are members of the Scleractinia, the so-called stony corals. The stoniness refers to the calcareous skeleton that they all have. But not all corals live in the tropics. We actually have two species of stony corals in Northern California. The brown cup coral, Paracyathus stearnsi, lives subtidally, and I think I've seen maybe a handful in all my intertidal explorations. The orange cup coral, Balanophyllia elegans, extends up into the low intertidal, and can be very common at certain sites I visit regularly. When I see them at low tide they are emersed and look like orange blobs. But if you touch one with your finger, you can feel the hardness of the calcareous base.

Orange cup corals growing among coralline algae
Orange cup coral (Balanophyllia elegans) at Asilomar
2015-10-26
© Allison J. Gong

Stony corals they may be, but Paracyathus and Balanophyllia are both solitary; that is, they aren't colonial. Each polyp developed from its own larva and lives its own life independent from all other corals. Its bright orange color makes Balanophyllia pretty conspicuous, even though most of them are less than 10 mm in diameter. They do occur in patches, which makes one wonder. If they're solitary rather than clonal or colonial, how do these patches arise?

To answer this question we need to venture into the lab and examine the biology of Balanophyllia more closely. Fortunately, they grow in the lab quite happily. Years ago my friend Cris collected a bunch of Balanophyllia and glued them to small tiles so they could be moved around and managed in the lab. Cris has since moved on to other things, but the corals remain in the lab to be studied. They are beautiful animals, and can't really be appreciated in the intertidal because at low tide they're all closed up. But look at how pretty they are when they're relaxed and open:

Group of adult cup corals, Balanophyllia elegans
Cluster of orange cup corals (Balanophyllia elegans)
2020-05-01
© Allison J. Gong

Like all cnidarian polyps, these corals have long tentacles loaded with stinging cells, or cnidocytes. See the little bumps on the otherwise translucent tentacles? Those are nematocyst batteries, clusters of stinging cells.

Orange cup coral, Balanophyllia elegans
Orange cup coral (Balanophyllia elegans)
2020-05-01
© Allison J. Gong

Let's get back to the biology of this beast and how it is that they seem to live in groups. Balanophyllia is a solitary coral with separate sexes--each polyp is either male or female. They are also brooders. Males release sperm, which are ingested by a nearby female. The female broods fertilized eggs in her gastrovascular cavity. After a long period, perhaps several months, a large reddish planula larva oozes out of the mother's mouth and crawls around for a while, generally settling and metamorphosing near its parent.

Planula larva of Balanophyllia elegans
Planula larva of Balanophyllia elegans
29 April 2020
© Allison J. Gong

This planula is a very squishy elongate blob, and can measure anywhere from 1-4 mm in length. It is an opaque red color, has a ciliated epidermis, and lacks a mouth or digestive system. Instead of feeding, it survives on energy reserves that its mother partitioned in the egg. You might surmise that not being able to eat would necessitate a quick metamorphosis into a form that has a mouth, but you'd be wrong. While some of them do indeed settle and metamorphose very close to their parent, others crawl around for several weeks, showing no inclination to put down roots and take on life as a sessile polyp. Perhaps they can take up enough dissolved organic matter from the seawater to sustain them through a long period of fasting.

At some point, though, the larva settles and metamorphoses into a little polyp. In the lab at least, Balanophyllia larvae settle on a variety of surfaces--glass, various plastics, even the fiberglass of the seawater tables.

Juvenile cup coral and three planula larvae of Balanophyllia elegans
Recently metamorphosed coral and three planula larvae of Balanophyllia elegans
2020-04-29
© Allison J. Gong

The little coral measures about 2 mm in diameter and has 12 tentacles. It feeds very happily on brine shrimp nauplii and should grow quickly. Those three larvae, though, may hang around forever. I got tired of waiting for them to do something and released them into the seawater system. It might or might not have been an accident.

So there we have it. Our local cold-water coral, which doesn't form reefs or live in colonies. Balanophyllia may seem atypical for corals, but what it really demonstrates is the diversity within the Scleractinia. It reminds us that generalities do indeed have some value, and that for the discerning mind it is the exceptions to the generalities that are most interesting.

3

Of course, sea anemones don't have faces. They do have mouths, though, and since a mouth is usually part of a face, you can sort of imagine what I'm getting at. The sunburst anemone, Anthopleura sola, is one of my favorite intertidal animals to photograph. Of the four species of Anthopleura that we have on our coast, A. sola is the most variable, which is why it keeps catching my eye.

This afternoon I met the members of the Cabrillo College Natural History Club for the low tide at Natural Bridges. Here are some of the A. sola anemones we saw.

Sunburst anemone (Anthopleura sola) at Natural Bridges
2020-02-22
© Allison J. Gong
Sunburst anemone (Anthopleura sola) at Natural Bridges
2020-02-22
© Allison J. Gong
Sunburst anemone (Anthopleura sola) at Natural Bridges
2020-02-22
© Allison J. Gong
Sunburst anemone (Anthopleura sola) at Natural Bridges
2020-02-22
© Allison J. Gong
Sunburst anemone (Anthopleura sola) at Natural Bridges
2020-02-22
© Allison J. Gong

Such an amazingly photogenic animal, isn't it?

This past Fall semester the NHC went tidepooling at Pigeon Point. Today we were at Natural Bridges, and later in the spring we are going to Asilomar. I didn't intend it, but this school year the club is getting a look at three very different intertidal sites.

I love it when things work out that way!

A while back now I went out on a low tide even though the actual low was after sunset. I figured that it was low enough that I'd have plenty of time to poke around as the tide was receding. And given that there were promising clouds in the sky, I took my good camera along just in case the sunset proved to be photo-worthy. Having had enough of crowds in the intertidal at Natural Bridges the previous day, I decided to venture up to Pistachio Beach, which isn't as heavily visited.

I ended up spending only 45 minutes in the intertidal, all the while watching the sun sink lower in the sky. It was already too dark to take many photos in the tidepools, but there were some interesting things on the beach.

The majority of shells that wash up on any beach are going to be molluscs, usually either gastropods or bivalves. I've often seen living red abalone (Haliotis rufescens) hidden in nooks and crannies at this site, so it's not surprising to find their shells on the sand. Usually, though, the shells are a little beat up. This one was intact, with a lovely layer of nacre inside.

This butterfly-shaped object is one of the shell plates of Cryptochiton stelleri, also known as the gumboot chiton. Cryptochiton is the largest of all chiton species; the largest one I've ever seen is the length of my forearm from elbow to fingertip. Like all chitons, C. stelleri has a row of eight shell plates running down the dorsal side of the body. Unlike other chitons, however, in Cryptochiton the plates are covered by a layer of tissue called the girdle and not visible from the outside. If you run your finger down the back you can feel the plates under the girdle. I never thought about it before now, but it seems that the name Cryptochiton refers to the hidden chiton-ness of the animal.

Anyway, Cryptochiton lives mostly in the subtidal, although you can occasionally see them in the very low intertidal. As subtidal creatures they have neither the ability nor the need to cling tightly to rocks, as their intertidal cousins do. This means that when big swells come through at low tide, they can get dislodged and wash ashore. I know from personal experience that the tissue of Cryptochiton is really tough. Once a pal and I were trudging back after working on a low tide and came across several dead Crytochiton scattered over the beach. We decided to do an impromptu dissection and try to salvage the plates, hacking away with her pocket knife. The smell was horrendous, and after several minutes we made practically zero progress, so we gave up. I've seen gulls pecking at dead Cryptochiton, too, and they didn't seem to have any success either. However, their bodies do eventually disintegrate, or something manages to eat them, and their naked plates can often be found on beaches.

Shell plate of Cryptochiton stelleri
2020-01-12
© Allison J. Gong

One of the coolest pattern I've ever seen in the intertidal was this:

Leaf barnacle (Pollicipes polymerus), mussels (Mytilus californianus), and limpets (Lottia sp.)
2020-01-12
© Allison J. Gong

I've never seen anything like this before. It's hard to tell from the photo, but these two rock faces converge into the crevice, sort of like the adjacent pages in an open book. This side of the rock surface faces away from the ocean and will never be subject to the main force of pounding waves. The barnacle in the middle is attached pretty much in the deepest part of the crevice, and is surrounded by mussels, which are then surrounded by limpets.

Now, all of these animals recruited to this location after spending some period of time, from a few days to a few weeks, in the plankton. The barnacle certainly can't move once it has settled and metamorphosed. Newly settled mussels have a limited ability to scoot around a bit but are generally stationary once they've extruded their byssal threads and fastened them to something hard. The limpets, on the other hand, are quite mobile. The barnacle and mussels gave up their ability to move around after they became benthic, but limpets can and do locomote quite a bit--in fact, they have to, in order to feed. So in a sense, these limpets "chose" to aggregate together long after settlement.

What are the ecological implications of this pattern?

Well, for one thing, that barnacle is a genetic dead end. I've written before about the bizarre sex lives of barnacles. This one lone barnacle, far from any others of its species, is not able to reproduce. It has nobody to copulate with. It is possible that other barnacles will recruit to the mussels (Pollicipes is often associated with Mytilus), but until then there will be no sexy times for this individual.

Another ecological consequence concerns the limpets. If these are owl limpets (Lottia gigantea), then some of them will grow up to be the big females that maintain farms on the rocks where they manage and harvest the crop of algal film that grows. These big females are territorial, and will bump or scrape off any creature found to be trespassing on their farms. Clearly, none of the limpets in the photo above are demonstrating any type of territorial behavior! So they are either some other species of Lottia, or are younger individuals of L. gigantea that haven't yet made the change from male to female.

In any case, I do think the pattern is very interesting, even though I don't understand it. Or maybe because I don't understand it. I'm always intrigued by something that I can't explain, which is a good thing because it means I don't get bored very often. If anyone reading this has an explanation for this pattern, let me know about it!

4

It has been a while since I've spent any time in the intertidal. There isn't really any reason for this, other than a reluctance to venture out in the afternoon wind and have to fight encroaching darkness. There's also the fact that I much prefer the morning low tides, which we'll have in the spring. However, this past weekend we had some spectacular afternoon lows, and although I was working on Friday and couldn't spare the time to venture out, I went out on Saturday and Sunday.

Saturday was a special day, because I had guests with me. A woman named Marla, who reads this blog, contacted me back in the fall. She said she wanted to do something special for her husband's birthday, and asked if I'd be willing to take them to the intertidal. It turns out that Andrew's birthday was around this past weekend, and he had family coming out from Chicago to celebrate. They picked the perfect weekend, because the low tides we had were some of the lowest of the year. So on Saturday I met up with Marla, Andrew (her husband), and Betsy (Andrew's sister) and we all traipsed out to Natural Bridges.

This was our destination for the afternoon:

Intertidal "island" at Natural Bridges
2020-01-11
© Allison J. Gong

Taking civilians into the intertidal can be tricky, because they often come with expectations that don't get met. Like expecting to see an octopus, for example. I explain that the octopuses are there, but are better at hiding from us than we are at finding them, but that never feels very satisfactory. This trio, however, were fun to show around. The tide was beautifully low and we had fantastic luck with the weather. It had rained in the morning, but the afternoon was clear and sunny. I congratulated Marla on remembering to pay the weather bill. And the passing stormlet didn't come with a big swell, so the ocean was pretty flat. We were able to spend some quality time in the mid-tidal zone, with occasional forays into the low intertidal.

Andrew, Marla, and Betsy standing on intertidal mussel bed at Natural Bridges State Beach
Andrew, Marla, and Betsy at Natural Bridges
2020-01-11
© Allison J. Gong

The typical Natural Bridges fauna--owl limpets, mussels, chitons, anemones, etc.--were all present and accounted for. Of course, there isn't much algal stuff going on in mid-January.

Given the time of year (mid-January) and the time of day (late afternoon), the sun was coming in at a low angle. This was tricky for photographing, both in and out of water. However, sometimes good things happen, as in this photo below:

Tidepool at Natural Bridges
2020-01-11
© Allison J. Gong

That's a big kelp crab (Pugettia producta) nestled among four sunburst anemones (Anthopleura sola). Kelp crabs are pretty placid creatures, for crabs, and usually take cover when approached. But this one remained in plain sight, holding so still that I thought it was dead. Even when I hovered directly over it and blocked the sun, it didn't move at all. Then it occurred to me that maybe he was having the sexy times with a lady friend. So I very carefully reached down and gave him a tap on the carapace. He flinched a little, so I knew he wasn't dead, but made no move to get away. And I caught a glimpse of a more golden leg underneath him.

Pair of courting kelp crabs (Pugettia producta)
2020-01-11
© Allison J. Gong

Crabs live their entire lives encased in a rigid exoskeleton, and can mate only during a short window of opportunity after a female molts. Early in the breeding season, a female crab uses pheromones to attract nearby males. When a suitable male approaches, she may let him grab her in a sort of crabby hug. That's what this male kelp crab is doing to his mate. They may remain in this embrace for several days, waiting until the female molts and her new exoskeleton is soft. At that point the male will use specialized appendages to insert packets of sperm into the female's gonopores. The two will then go their separate ways.

We didn't disturb these crabs, and let them go on doing their thing. By now the sun was going down, so we headed back up and were rewarded with a glorious sunset.

Always a great way to end the day!

The intertidal sculpins are delightful little fish with lots of personality. They're really fun to watch, if you have the patience to sit still for a while and let them do their thing. A sculpin's best defense is to not be seen, so their first instinct is to freeze where they are. Then, if a perceived threat proves to be truly frightening, they'll scoot off into hiding. They can also change the color of their skin, either to enhance camouflage or communicate with each other.

Around here we have a handful of sculpin species flitting around in our tidepools. Sculpins can be tricky to identify even if you have the fish in hand--many of the meristics (things you count, such as hard spines and soft rays in the dorsal fin, or the number of scales in the lateral line) used to distinguish species actually overlap quite a lot between species. The fishes' ability to change color means that skin coloration isn't a very reliable trait. When I was in grad school there was another student in my department who was studying the intertidal sculpins, and she told me that most of the ones we see commonly are either woolly sculpins (Clinocottus analis) or fluffy sculpins (Oligocottus snyderi). I've developed a sort of gut feeling for the gestalt of these species, but I'm not always 100% certain of my identifications.

Sculpin in a tidepool at Asilomar State Beach. The fish is colored pink and brown, to match its surroundings in the tidepool.
Sculpin at Asilomar State Beach
2019-07-04
© Allison J. Gong

Anyway, back to the camouflaged sculpins. The ability to change the color of the skin means that sculpins can match their backgrounds, which comes in very handy when there isn't anything to hide behind. Since the environment is rarely uniformly colored, sculpins tend to have mottled skin. Some can be banded, looking like Oreo cookies. The fish in this photo lives in a pool with a granite bottom. The rock contains large quartz crystals and is colonized by tufty bits of mostly red algae. There is enough wave surge for these fist-sized rocks to get tumbled about, which prevents larger macroalgae from colonizing them.

Other shallow pools higher up in the intertidal at Asilomar have a different type of rocky bottom. The rocks lining the bottom of these pools are whitish pebbles that are small enough to be tossed up higher onto the beach. I don't know whether or not these pebbles have the same mineral content as the larger rocks lower in the intertidal, but they do have quartz crystals. The pebbles are white. So, as you may have guessed, are the sculpins!

Sculpins on a gravel bottom in a tidepool at Asilomar State Beach. The fish are white and gray in color, to match the color of the gravel in their pool.
Sculpins at Asilomar State Beach
2019-07-04
© Allison J. Gong

Other intertidal locations have different color schemes. On the reef to the south of Davenport Landing Beach, you will see a lot of coralline algae. Some pools are overwhelmingly pink because of these algae. Bossiella sp. is a common coralline alga at this location.

What color do you think the sculpins are in these pools?

Give yourself a congratulatory pat on the back if you said "pink"!

Sculpin in a tidepool at Davenport Landing. The fish is mottled pink and brown, for camouflage among the pink coralline algae in the pool.
Sculpin and coralline algae (Bossiella sp.) at Davenport Landing
2017-06-27
© Allison J. Gong

Sculpins aren't the only animals to blend in with coralline algae. Some crustaceans are remarkably adept at hiding in plain sight by merging into the background. Unlike the various decorator crabs, which tuck bits and pieces of the environment onto their exoskeletons, isopods hide by matching color.

Turning over algae and finding hidden creatures like these is always fun. For example, I saw these isopods at Pescadero this past summer. See how beautifully camouflaged they are?

Sometimes, when you're not looking for anything in particular, you end up finding something really cool. Last weekend I met up with students in the Cabrillo College Natural History Club for a tidepool excursion up at Pigeon Point. We were south of the point at Whaler's Cove, where a staircase makes for comparatively easy access to the intertidal.

Photo of Whaler's Cove just south of Pigeon Point, during an autumn afternoon low tide
Whaler's Cove at Pigeon Point
2019-11-24
©Allison J. Gong

It's fun taking students to the intertidal because I enjoy helping them develop search images for things they've never seen before. There really is so much to see, and most of it goes unnoticed by the casual visitor. Often we are reminded to "reach for the stars," when it is equally important to examine what's going on at the level of your feet. That's the only way you can see things like this chiton:

A chiton (Mopalia muscosa), heavily encrusted with a variety of red algae, at Whaler's Cove.
Mopalia muscosa at Whaler's Cove
2019-11-24
© Allison J. Gong

Mopalia muscosa is one of my favorite chitons. It is pretty common up and down the California coast. However, like most chitons it is not very conspicuous--it tends to be encrusted with algae! This individual is exuberantly covered with coralline and other red algae and has itself become a (slowly) walking bit of intertidal habitat. It is not unusual to see small snails, crustaceans, and worms living among the foliage carried around by a chiton. Other species can carry around some algae, but M. muscosa seems to be the most highly decorated chiton around here. I showed this one to some of the students, who then proceeded to find several others. A search image is a great thing to carry around!

Compared to the rocky intertidal, a sandy habitat can be a difficult place to live. Sand is inherently unstable, getting sloshed to and fro with the tides. Because of this instability there is nothing for holdfasts to grab, so there are many fewer algae for animals to eat and hide in. Most of the life at a sandy beach occurs below the surface of the sand, and is thus invisible to anyone who doesn't want to dig. There's a beach at Whaler's Cove where I've found burrowing olive snails (Olivella biplicata) plowing along just below the surface. I wanted to show them to the students, so I waded in and rooted around. I did find Olivella, but I also found a burrowing shrimp. I think it's a species of Crangon.

Shrimp on sandy bottom of a shallow tidepool at Whaler's Cove. The shrimp is colored to match the sand.
Shrimp (Crangon sp.) at Whaler's Cove
2019-11-24
©Allison J. Gong

Now that is some damn fine camouflage! If the shrimp didn't cast its own shadow, it would be invisible. Even so, it was clearly uneasy sitting on the surface like that. I had only a few seconds to shove the camera in the water and snap a quick photo before the shrimp wriggled its way beneath the sand again.

As I've said before, observation takes practice and patience. To look at something doesn't mean you truly see it. That's why it is so important to slow down and let your attention progress at the pace of the phenomenon you're observing. If the only things that catch your eye are the ones that flit about, then I can guarantee you will never find a chiton in the intertidal. And wouldn't that be a sad thing?

2

When we stop to marvel at the wonders of the natural world, we usually forget about all the life that is going on that we don't get to see. But there is a lot happening in places we forget to look. For example, any soil is an entire ecosystem, containing a variety of small and tiny animals, bacteria, and fungi. In fact, if a fungus didn't send up a fruiting body (a.k.a. mushroom) every once in a while, most observers wouldn't realize it was there at all. We humans tend to behave as though something unseen is something that doesn't exist, and I admit to the very same thinking with regards to my own kitchen: anything stored way up in cupboards I can't reach, may as well not be there at all.

But there are places where we can witness the life occurring below our feet, and floating docks in marinas and harbors are some of the best. Of course, the trick is to "get your face down where your feet are", a piece of advice about how to observe life in tidepools that applies just as well to investigating the dock biota. Once you get used to the idea of lying on the docks, which can be more or less disgusting depending on time of year and number of birds hanging around, a whole new world literally blossoms before your eyes.

Some of the flower-looking things are indeed anthozoans ('flower animal') such as this plumose anemone:

Plumose anemone (Metridium senile) at the Santa Cruz harbor
09 October 2019
© Allison J. Gong

and this sunburst anemone:

Sunburst anemone (Anthopleura sola) at the Santa Cruz harbor
09 October 2019
© Allison J. Gong

Other animals look like dahlias would look if they were made of feathers. Maybe that doesn't make sense. But see what I mean?

This is Eudistylia polymorpha, the so-called feather duster worm. These worms live in tough, membranous tubes attached to something hard. They extend their pinnate tentacles for feeding and are exquisitely sensitive to both light and mechanical stimuli. There are tiny ocelli (simple, light-sensing eyes) on the tentacles, and even casting a shadow over the worm causes it to pull in its tentacles very quickly. This behavior resembles an old-fashioned feather duster, hence the common name. These were pretty big individuals, with tentacular crowns measuring about 5 cm in diamter. Orange seems to be the most common color at the Santa Cruz harbor.

One of the students pointed down at something that he said looked like calamari rings just below the surface. Ooh, that sounds intriguing!

And he was right! Don't they look like calamari rings? But they aren't. These are the egg ribbons of a nudibranch. They appeared to have been deposited fairly recently, so I went off on a hunt for the likely parents. And a short distance away I caught the nudibranchs engaging in the behavior that results in these egg masses. Ahem. I don't know if the term 'orgy' applies when there are three individuals involved, but that's what we saw.

Mating aggregation of Polycera atra
09 October 2019
© Allison J. Gong

To give you some idea of how these animals are oriented, that flower-like apparatus is the branchial (gill) plume, which is located about 2/3 of the way down the animal's dorsum. The anterior end bears a pair of sensory organs called rhinophores; they look kind of like rabbit ears. You can see them best in the animal on the left.

When you see more than one nudibranch in such immediate proximity it's pretty safe to assume that they were mating or will soon be mating. Nudibranchs, like all opisthobranch molluscs, are simultaneous hermaphrodites, meaning that each can mate as both a male and a female. The benefit of such an arrangement is that any conspecific individual encountered is a potential mate. The animals pair up and copulate. I'm not sure if the copulations are reciprocal (i.e., the individuals exchange sperm) or not (i.e., one slug acts as male and transfers sperm to the other, which acts as female). In either case, the slugs separate after mating and lay egg masses on pretty much whatever surface is convenient. Each nudibranch species lays eggs of a particular morphology in a particular pattern. Some, such as P. atra, lay eggs in ribbons; others produce egg masses that look like strings of miniature sausages.

Nudibranch (Polycera atra) at the Santa Cruz harbor
09 October 2019
© Allison J. Gong

This is the first time I've seen big Polycera like these. The slugs were about 4 cm long. They eat a bryozoan called Bugula, and there is a lot of Bugula growing at the harbor these days. Maybe that's why there were so many Polycera yesterday. Nudibranchs are the rock stars of the invertebrate world--they are flamboyantly and exuberantly colored, have lots of sex, and die young. They can be very abundant, but tend to be patchy. Quite often an egg mass is the only sign that nudibranchs have been present.

The next time you happen to be at a marina poke your head over the edge and take a look at the stuff living on the dock. Even if you don't know what things are, you should see different textures and colors. With any luck, you'll be pleasantly surprised at the variety of life you find under your feet.

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We usually think of sea stars as the colorful animals that stick to rocks in the intertidal. You know, animals like Pisaster ochraceus (ochre star) and Patiria miniata (bat star). I see these animals all the time in the intertidal, and if you're a regular reader of this blog you've probably seen the photos that I post here. Given how prominent P. ochraceus and P. miniata can be in the rocky intertidal, it may be a bit of a surprise to learn that not all sea stars live on rocks. In fact, some can't even really stick to a rock.

This morning I was meandering through the Seymour Center when I stopped at a recently refurbished tank. The new inhabitants are a couple of curlfin sole (Pleuronichthyes decurrens) and their secretive and strange roommate. Here's one of the flat fish:

Photo of curlfin sole (Pleuronichthyes decurrens)
Curlfin sole (Pleuronichthyes decurrens) at the Seymour Marine Discovery Center
5 September 2019
© Allison J. Gong

The other fish was hiding up against the wall in one of the back corners and didn't come down until it was feeding time.

The secretive roommate was all but invisible. Here's a photo. Ignore the fish's tail. Do you see anybody else?

Who is this?

Fortunately for all of the tank's inhabitants, feeding time was just around the corner. I knew what would happen, so I stuck my phone on the glass and recorded some video. Keep an eye on the upper left-hand corner. Watching the fish eat is entertaining, too. Just how do they manage with those tiny sideways mouths?

It's not the greatest bit of video, but did you see what happened? That creature emerging from the sand is Astropecten armatus, a sea star that lives in sand. And did you notice how fast it moves? Most of the time it is buried under the sand and usually comes out only to grab food. Every once in a while I'll find it on one of the walls but most of the time it is essentially invisible to human viewers on the other side of the glass.

Astropecten armatus
5 September 2019
© Allison J. Gong

All spread out, this Astropecten is probably a little smaller than my hand. It has a smooth-ish aboral (i.e., top) surface, lacking the spiny protuberances that Pisaster has. The texture of the aboral surface is similar to that of the bat star, Patiria miniata. The species epithet, armatus, means 'armored' and refers to the row of marginal plates along the perimeter of the body. These plates bear a row of spines that point up and another row that point down. Astropecten is unusual among sea stars for having suckerless tube feet. Its tube feet are pointed, and instead of being super grippy, work to push sand around so the animal can sort of bulldoze its way along. As always, form follows function!

Olivella biplicata at Whaler's Cove
3 January 2019
© Allison J. Gong

In the wild, A. armatus lives on sandy flats, rarely exposed even at low tide. One of its favorite prey items is the olive snail, Olivella biplicata. Imagine this life-and-death encounter taking place below the surface of the sand: Olivella is burrowing through the sand, minding its own business and unaware that Astropecten is following the slime trail it (Olivella) left behind. Astropecten catches up to Olivella, shoves a couple of arms into the sand around Olivella, engulfs the snail, and swallows it whole. Eventually an empty Olivella shell is spat out. Incidentally, many small hermit crabs, especially Pagurus hirsutiusculus and juveniles of other Pagurus species, live in Olivella shells. I've often wondered why there are so many empty but intact olive snail shells for the hermit crabs to find, and now suppose that Astropecten's method of feeding might have something to do with it.

Interesting star, this Astropecten. I'm really happy that it is on exhibit again, because even most visitors will never see it, watching it come out to feed is always fun.

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A few weeks ago I was contacted by a woman named Kathleen, who reads this blog and is herself a student of the seaweeds. She said that she studies a site up at Pescadero, about an hour up the coast from me. We decided to meet up during the series of low tides around the Fourth of July so we could explore the area together, and she could help me with my algal IDs. My friend and former student, Lisa, joined us for the fun.

Map of the Pescadero Point region
05 July 2019
© Google

The most prominent landmark along the coastline in this region is Bird Island, which is accessible only at minus tides, when it is revealed to be a peninsula. It smells pretty much as you probably imagine, especially if you happen to be downwind. Given the prevailing wind direction, that means that the closer you get to Bird Island from the south, the stronger the smell. Kathleen's site is the south side of Pescadero Point, fortunately far enough south of Bird Island that the smell isn't noticeable from that distance. She has a permanent transect that she surveys regularly, taking note of algal abundances and distributions.

South side of Pescadero Point
05 July 2019
© Allison J. Gong

One of the notable things we all noticed was the conspicuous presence of big, healthy ochre stars (Pisaster ochraceus)--many hand-sized or larger. I also saw many smaller stars, in the 2 cm size range, but these were hidden in crevices or under algae. The big guys and gals, were out there in plain sight.

Constellation of ochre stars (Pisaster ochraceus) at Pescadero Point
05 July 2019
© Allison J. Gong

However, not all was perfect for the sea stars at Pescadero Point. One of the ochre stars showed symptoms of sea star wasting syndrome (SSWS). It had autotomized two of its arms and had a sloppy, goopy open wound that extended into the oral disc. It was also mushy when I touched it and didn't firm up the way healthy stars do. This star is a goner, even though it doesn't know it yet. That's the beauty (and in this case, tragedy) of an entirely decentralized nervous system.

Sick ochre star (Pisaster ochraceus) at Pescadero Point
05 July 2019
© Allison J. Gong

After I mentioned having seen a sick sea star we compared notes on the current status of SSWS. What more do we know about the syndrome, and any recovery of stars? We came to the consensus that the oubreak was probably caused by a perfect storm of ecological conditions--an opportunistically pathogenic virus that is ubiquitous in the environment, environmental stresses, and high population densities both intertidally and subtidally. Kathleen asked me what I had been seeing recently. I told her that Pisaster ochraceus, one of the species that melted away in spectacular fashion, seems to be making a strong comeback in the places where I used to see it in large numbers. Even though every once in a while i see a sick star, places like Natural Bridges and Davenport Landing are again populated by lots of hand-sized-or-bigger ochre stars. Which of course brings up the question of where these large stars suddenly came from. I think they were tiny stars when the outbreak occurred, hiding in the mussel beds. Many of them died, but as with any plague there are always some survivors. Those lucky few managed to hang on and creep into the niches that opened up when so many adults died. But would little juveniles only a few millimeters in diameter be able to grow to the sizes that we're seeing now, in ~5 years? I suppose that's not out of the question, and we know that when fed well in the lab they grow very quickly, but individual growth rates in the field are difficult to measure.

Another animal goody that we saw were clusters of the bryozoan, Flustrellidra corniculata. Unlike most bryozoans, which are calcified and crunchy, Flustrellidra colonies are soft and flexible. They look more like strange, thick pieces of brown algae than anything recognizable as a bryozoan.

Flustrellidra corniculata at Pescadero Point
05 July 2019
© Allison J. Gong

We were there to do some basic marine botany, and although I kept getting distracted by the invertebrates I did also pay attention to the floral aspect of Kathleen's site. She pointed out that Laminaria sinclairii, one of the small low-intertidal kelps, was always abundant. It's true, there were rocks that were entirely covered with L. sinclairii, like this one:

Laminaria sinclairii at Pescadero Point
05 July 2019
© Allison J. Gong

Laminaria sinclairii and L. setchellii are the most common intertidal species of the genus on our coast. They are easily distinguishable because L. sinclairii has a single undivided blade arising from the stipe, and L. setchellii has a blade that is subdivided into fingerlike sections; in fact, the former species epithet for L. setchellii was dentigera, referring to 'finger'.

Laminaria setchellii at Franklin Point
15 June 2018
© Allison J. Gong

See the difference?

There is a third species of Laminaria on our coast, that I knew only by reputation. What I'd heard is that Laminaria ephemera resembles L. sinclairii except for the morphology of the holdfast: L. ephemera has a discoid, suction-cup holdfast while L. sinclairii has the more typical hapterous holdfast (made of intertwined cylindrical projections). I think I might have seen a few L. ephemera at Pescadero. These thalli appear to have suction-cup holdfasts, don't they?

Laminaria ephemera(?) at Pescadero Point
05 July 2019
© Allison J. Gong
North side of Pescadero Point
05 July 2019
© Allison J. Gong

We didn't spend much time on the south side of the point, but scrambled over the rocks to the north side, where there are stretches of sandy beach between rocky outcrops. Bird Island is that peninsula in the top of the picture. As I mentioned above, it is connected to the beach only at low tide, so while I think of it as a peninsula, it really is an island most of the time.

Once on the north side of the point we slowed down and made some more attentive observations of the flora. It turns out that this portion of our intertidal visit was sponsored by the letter 'P'. One of the things we all noticed was the prevalence of Pyropia, the filmy red alga that is common in the high-mid intertidal. The thallus of Pyropia consists of a single layer of cells connected to form a very thin elastic tissue. It dries to a crisp in the sun, but rehydrates when the tide returns. You've probably encountered Pyropia before without realizing it: nori is made of Pyropia that has been shredded and processed into paper-like sheets, used for things like sushi rolls.

Although it looks uniformly blackish-green when packaged for human consumption, Pyropia's color in life is a glorious iridescent mixture of greens, olives, and purples. It is another of those easily overlooked denizens of the intertidal that deserves a much closer look than it usually gets.

Pyropia sp. at Pescadero Point
05 July 2019
© Allison J. Gong
Plocamium cartilagineum at Pescadero Point
05 July 2019
© Allison J. Gong

Another common red alga at Pescadero Point is the delicate and lacy Plocamium cartilagineum. This is one of the hobbyist phycologist's favorite species because it presses like a dream and makes great gifts or wall decorations. As I wrote about here, Plocamium has a doppelganger: Microcladia coulteri. These algae share a similar morphology, but as I mentioned in the previous post, natural history makes it easy to distinguish between the two in the field. Microcladia is epiphytic, growing on other algae, and Plocamium is not.

Plocamium grows on rock surfaces in the mid-to-low tide regions. It sometimes gets surrounded or even buried in sand, but if you dig down far enough you'll always find the holdfast attached to a rock (or shell or other hard object).

Last month I wrote about Postelsia palmaeformis, the sea palm. We found a most handsome specimen washed up on the beach. Note that, as per usual, it wasn't the holdfast of the kelp that failed. The holdfast did its job perfectly well, and it was the mussel it was attached to that broke free of the rock.

Postelsia palmaeformis at Pescadero Point
05 July 2019
© Allison J. Gong

The sad thing about finding great specimens like this on the beach is the realization that it will soon be dead. In fact, so will the mussel. Such is the price organisms pay for failing to hang onto their substrate (or for their substrate's failure to hang on). The rocky intertidal is a harsh place to live, and can be unforgiving of mistakes and bad decisions.

That's part of the reason I find it so fascinating. Most wild organisms live on the knife-edge of survival, with only the thinnest margin between life and death. Every organism has its predators, pathogens, and parasites to deal with on a daily basis, in addition to the physical stresses of its habitat. All of the organisms that I study in the intertidal are marine--not freshwater or even brackish, although some can tolerate reduced salinity (and on the other extreme, some tolerate very high salinity). They evolved to live in the ocean, in a habitat where the ocean abandons them for a few hours twice a day. Yet as improbable as that sounds, the diversity in the intertidal is astonishingly high. Obviously, for those that can live there, the trade-off between stability and safety is worthwhile. Nature will always find a way.

Be honest now. When you think of clams, what comes to mind? If you're like most people, visions of clams steamed in white wine, garlic, and butter might dance in your head. Or perhaps clams in cioppino or a hearty chowder would be your go-to. In any case, I doubt that clams, as actual living creatures, occupy much of your brain. Because let's face it, at first glance even living clams aren't the most energetic and charismatic animals. Most of the really cool things that they do, like suck water through their shells for filter feeding and gas exchange, they do while buried in the mud.

When you think about it, though, just the fact that clams live in the sand or mud while depending on water that may be quite far from them is rather amazing. All animals require oxygen, and for marine animals that oxygen comes from seawater. Animals that move freely through the water have access to a ready supply of oxygen. But clams live more or less fixed lives encased in sediment, and water can be quite far from their bodies. How, then, do they pull water into their shells and across their gills? They use siphons, which can reach up to the surface of the sediment into the water column.

Siphons of a pholad bivalve at Pleasure Point
16 July 2018
© Allison J. Gong

A clam has two siphons--one pulls clean water into the shells and the other expels water from the shells. This arrangement allows for one-way flow across the gills, which serve double duty as both feeding and gas exchange organs. The siphons themselves are somewhat muscular and can open and close, but it's the ciliary action of the gills that create the actual water current. In a living clam the only visible body parts are the siphons, which in some species (e.g., geoducks) are so large that they cannot be entirely withdrawn into the shells.

Of the two siphons in the picture above, can you tell which is the incurrent and which is the excurrent? What do you think is the functional significance of that network of white structures that cover the opening of one of the siphons?

Not only do clams live buried in sediment, but some of them can actually bore into rocks. These boring clams, the pholads, have shells that are morphologically and functionally different from the typical clams you've encountered in cioppino. They are elongated on the anterior-posterior axis and the anterior ends are heavily sculpted and fortified to grind into rock. Of course, they can do this only in areas where the rock is soft--you don't see pholads burrowed into granite, for example.

Pholad siphons at Pleasure Point
16 July 2018
© Allison J. Gong

Fortunately for the pholads, much of the rock in the Santa Cruz area is a soft mudstone, easily eroded and burrowed into. I've seen pholads at intertidal sites from Capitola to Davenport. Both dead pholads and live pholads can be seen, but it takes a careful eye to spot the live ones. Of course, all you'd ever see of a living pholad is the siphons. When the animal dies, though, the shells are left behind. As the mudstone continues to erode the shells can be exposed, just like fossils. And as a matter of fact, the mudstone formations around here are known for their fossil contents. I think, but am not certain, that these empty shells in holes belong to Parapholus californica.

Pholad shells at Natural Bridges
8 June 2019
© Allison J. Gong

How does a clam burrow into even soft rock? A description of burrowing activity of Parapholus californica can be read here. As you can imagine, it's a slow and continuous process. Fortunately, these clams don't have much else going on and can take their time. In some ways, their lifestyle sounds pretty ideal: hang out in a snug burrow where predators can't get at your soft body and extend your siphons out to bring in clean water for food and oxygen. Sure, when it comes to reproduction the only option available is free-spawning and hoping for the best, but that has proven to be a successful strategy for countless generations of your kind. Aside from the cost of making gametes, it's a pretty low-energy way to produce offspring. Maybe the old saying "happy as a clam" isn't that far off the truth.

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