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4

On the afternoon of July 31, 2020 the world of invertebrate biology and marine ecology in California lost a giant in our field. Professor Emeritus John S. Pearse died after battling cancer and the aftereffects of a stroke.

John Pearse in the intertidal at Soquel Point
2017-05-28
© Allison J. Gong

John was one of the very first people I met when I came to UC Santa Cruz. Before we moved here, my husband and I came and met with John, who was not my official faculty sponsor but agreed to show us around so we could check out different areas for a place to live. In fact, I had applied to the department to do my graduate work in John's lab, but because he was considering retirement the department wouldn't let him take on a new Ph.D. student. But when we needed some help getting acquainted with Santa Cruz, John and his wife, Vicki Buchsbaum Pearse, graciously let us stay at their house and spent a day driving us around town and showing us eateries as well as potential neighborhoods.

By happenstance we ended up living down the hill from John and Vicki. We had met their blue duck, Lily, and I used to fill spaghetti sauce jars with snails from our tiny yard and trudge up the hill to feed them to her. She gobbled them up like they were her favorite treat.

As one of the regional experts in invertebrate biology, John was on all of my graduate committees. There were always a half-dozen or so of us grad students working with invertebrates, and we all tended to hang out together. John was one of the things we shared in common. And even if he wasn't technically on one's committee, he would always be available for consultation or advice as needed.

When John retired, he didn't leave the campus. He remained a presence at the marine lab, and still did field work. He started incorporating young students in his long-term intertidal monitoring research, which morphed into the LiMPETS project. The combination of working with students while producing robust scientific data was the perfect distillation of John's legacy. He said this about LiMPETS:

This is one of the best things I could ever do to enhance science education and conservation of our spectacular coastline. Working with teachers and their students is a wonderful and fulfilling experience.

John S. Pearse, Professor Emeritus
UC Santa Cruz

The last time I saw John was in the summer of 2019, during his annual Critter Count. He started these Critter Counts back in the 1970s, monitoring biota at two intertidal sites in Santa Cruz. These sites have since been incorporated into the LiMPETS program. I'm sure it made John smile whenever he thought of generation after generation of schoolkids traipsing down to the intertidal with their quadrats and transect lines, counting organisms the way he had for so many years.

When I started teaching my Ecology class, John suggested that I take the students out to Davenport Landing to monitor at the LiMPETS site there. That is another of his long-term sites, and he was worried about losing information if it were not sampled at least once a year. My students have done LiMPETS monitoring three years now, and John accompanied us on at least two of those visits. I tried to impress upon the students that having John Pearse himself come out with us was a Big Deal, but am not sure I was able to convince them of how fortunate they were. I bet there are a lot of marine biologists in California who would dearly love to go tidepooling with John. And now no one else will.

John Pearse and Todd Newberry, the other professor who gets the blame for how I think about biology, taught an Intertidal Biology class. I came along on many of the field trips the last year they taught it. I remember getting up before dawn to drive down to Carmel, park in the posh neighborhood streets, and walk down to meet John and Todd in the intertidal. I remember slogging through the sticky mud at Elkhorn Slough, digging for Urechis and hearing John shout "It's a goddamned brachiopod!" from across the flat. I remember bringing phoronid worms back to the lab, looking at them under the scope, and watching blood flow into and out of their tentacles. I remember John taking an undergraduate, Jen, and me out to Franklin Point, and showing me my very first staurozoans. That was probably around 1996, and I'm still in love with those animals.

I'm no John Pearse or Todd Newberry, but I'm a small part of their giant legacy in this part of the world. I strive to instill in my students the joy and intellectual pleasure in studying the natural world that I inherited from John and Todd. Partly to honor them, but mostly because it suits my own inclinations, I'm on a one-woman crusade to bring natural history back into modern science and science education.

I've spent the last two mornings in the intertidal at two of the LiMPETS sites, as part of a personal tribute to John. I thought there would be no greater way to memorialize John than by spending some quality time in the intertidal, where he trained so many young minds. I was thinking of him as I took photos, and thought he would be pleased if I shared them.

Natural Bridges—4 August 2020

Shore crab (Pachygrapsus crassipes)
2020-08-04
© Allison J. Gong

And because, like me, John had a special affinity for the anemones:

Sunburst anemone (Anthopleura sola)
2020-08-04
© Allison J. Gong
Sunburst anemone (Anthopleura sola)
2020-08-04
© Allison J. Gong

And he would have loved this. What is going on here? How did this pattern come to be?

Anemones (Anthopleura elegantissima and possibly A. sola)
2020-08-04
© Allison J. Gong

And look at this, three species of Anthopleura in one tidepool! Can you identify them?

Tidepool at Natural Bridges
2020-08-04
© Allison J. Gong

Davenport Landing—5 August 2020

It was windy and drizzly this morning. I ran into a friend, Rani, and her family out on the flats; they were leaving as I arrived. I hadn't seen her since before the COVID-19 lockdown began back in March. She was also visiting the tidepools to honor John Pearse. We chatted from a distance and exchanged virtual hugs before heading our separate ways.

It felt like a John Pearse kind of morning. I recorded the video clip I needed for class, collected some algae and mussels for a video shoot tomorrow, and took a few photos.

A typical intertidal assemblage (sea stars, sea anemones, and algae) at Daveport Landing
2020-08-05
© Allison J. Gong

And even though I'm not very good at finding nudibranchs, even I couldn't miss this one. It was almost 4 cm long!

Nudibranch (Triopha maculata) at Davenport Landing
2020-08-05
© Allison J. Gong

The ultimate prize for any tidepool explorer is always an octopus. When I take newbies into the field that's what they always want to see. I have to explain that while octopuses are undoubtedly there and common, they are very difficult to find. You can't be looking for them, unless you really like being frustrated.

But John must have been with me in spirit this morning, because I found this:

Red octopus (Octopus rubescens) at Davenport Landing
2020-08-05
© Allison J. Gong

It was just a small one, with the mantle about as long as my thumb. I found it because I spotted something strange poking out from a piece of algae. It was the arm curled with the suckers facing outward. I touched it, and the arm retracted. It didn't seem to like how I tasted.

And lastly, for me this is the epitome of John Pearse's legacy: Working in the intertidal, showing students how to identify owl limpets. I hope they never forget what it was like to learn from the man who with his wife, literally wrote the book about invertebrates and founded LiMPETS.

John Pearse in the intertidal with my students
2016-04-29
© Allison J. Gong

RIP, John S. Pearse. You left behind some enormous shoes to fill and a legacy that will stretch down through generations. I count myself lucky to have spent time with you in the field and in the lab. While I will miss you sorely, it is my privilege to pass on your lessons. Thank you for all you have taught me.

Every summer, like clockwork, my big female whelk lays eggs. She is one of a pair of Kellett's whelks (Kellettia kellettii) that I inherited from a labmate many years ago now. True whelks of the family Buccinidae are predatory or scavenging snails, and can get pretty big. The female, the larger of the two I have, is almost the length of my hand; her mate is a little bit smaller.

Many marine snails (e.g., abalones, limpets, and turban snails) are broadcast spawners, spewing large numbers of gametes into the ocean and hoping for the best. These spawners have high fecundity, but very few, if any, of the thousands of eggs shed will survive to adulthood. We say that in these species, parental investment in offspring extends only as far as gamete production. Fertilization and larval development occur in the water column, and embryos and larvae are left to fend for themselves.

The whelks, on the other hand, are more involved parents. They maximize the probability of fertilization by copulating, and the female produces yolky eggs that provide energy for the developing embryos and larvae. Rather than throw her eggs to the outside world and hoping for the best, the female whelk deposits dozens of egg capsules, each of which contains a few hundred fertilized eggs.

Over a period of about three weeks I shot several time-lapse video clips of the mama whelk laying eggs. Due to the pandemic we need to work in shifts at the lab. Fortunately I have the morning shift, which means I can start as early as I want as long as I leave before 11:00 when the next person comes in. Each 2.5-hr stint at the lab yielded about 30 seconds of video, not all of which was interesting; even in time-lapse, whelks operate at a snail's pace. Still, I was surprised at how active the female could be while she was apparently doing nothing.

The freshly deposited capsules are a creamy white color, as are the embryos inside them. As the embryos and then larvae grow, they get darker. Each of the fertilized eggs develops through the first molluscan larval stage, called a trochophore larva, within its own egg membrane. The embryo, and then the trochophore, survives on energy reserves provided by the mother snail when she produced the egg. These larvae don't hatch from their egg membrane until they've reached the veliger stage.

Pumpkin seed-shaped egg capsule of the whelk Kellettia kellettii, 13 mm tall.
Egg capsule of Kellettia kellettii
2020-06-20
© Allison J. Gong
Veliger larva of Kellettia kellettii
2020-07-25
© Allison J. Gong

The veliger larva gets its name from a lobed ciliated structure called a velum. Gastropods and bivalves have veliger larvae. As you might expect, the bivalve veliger has two shells, and the gastropod veliger has a coiled snail shell. These Kellettia veligers have dark opaque patches on the foot and some of the internal organs. That coloration is what you see in the photo of the egg capsule. You can see below which of the egg capsules are the oldest, right?

Mated pair of Kellettia kellettii and their egg capsules
2020-07-25
© Allison J. Gong

By the time the veligers emerge from the egg capsule, they have burned through almost all of the energy packaged in the yolk of the egg. They need to begin feeding very soon. The current generated by the beating cilia on the velum both propels the larva through the water and brings food particles to the larva's mouth. The velum can be pulled into the shell, and, as in any snail the opening to the shell can be shut by a little operculum on the veliger's foot. As is the case with most bodies, the veliger is slightly negatively buoyant, so as soon as it withdraws into the shell it begins to sink. However, once the velum pops back out the larva can swim rapidly.

Watch how the veliger swims. You can also see the heart beat!

So now the egg capsules are being emptied as the larvae emerge. I'm not keeping the veligers, so they are making their way through the drainage system back out to the ocean. As of now there are no iNaturalist observations of Kellettia kellettii in the northern half of Monterey Bay, so it appears that for whatever reason the whelks have not been able to establish viable populations here. Or it might be that the whelks are here but there aren't enough SCUBA divers in the water to see them.

These little veligers will be very lucky if any of them happen to encounter a subtidal habitat where they can take up residence as juvenile whelks. Even for animals that show a relatively high degree of parental care, the chances of any individual larva surviving to adulthood are exceedingly small. However, for the reproductive strategy of Kellettia to have evolved and persisted, there must be a payoff. In this case, the reward is an equal or greater reproductive success compared to snails that simply broadcast thousands of unprotected eggs into the water. Some gastropods such as the slipper shell Crepidula adunca, take parental care even further than Kellettia; in this species the mother broods her young under her shell until they've become tiny miniatures of herself, then she pushes them out to face the world and find a turban snail to live on. Crepidula adunca does not have a swimming larval stage at all. The fact that we see a variety of strategies—many eggs with little care, fewer eggs with more care, and brooding—indicates that there's more than one way to be successful.

I've written before about the rocky intertidal as a habitat where livable space is in short supply. Even areas of apparently bare rock prove to be, upon closer inspection, "owned" by some inhabitant or inhabitants. That cleared area in the mussel bed? Look closely, and you'll likely find an owl limpet lurking on the edge of her farm.

See?

Owl limpet at edge of her territory, a clear area surrounded by mussels.
Owl limpet (Lottia gigantea) on her farm at Natural Bridges
2017-04-01
© Allison J. Gong

And of course algae are often the dominant inhabitants in the intertidal.

Assemblage of algae in the intertidal
Assemblage of algae north of Waddell Creek
2020-06-09
© Allison J. Gong

When bare rock isn't available, intertidal creatures need other surfaces to live on. To many small organisms, another living thing may be the ideal surface on which to make a home. For example, the beautiful red alga Microcladia coulteri is an epiphyte that grows only on other algae. Smithora naiadum is another epiphytic red alga that grows on surfgrass leaves.

We describe algae that grow on other algae (or plants) as being epiphytic (Gk: epi "on" + phyte "plant"). Using the same logic, epizooic algae are those that live on animals. In the intertidal we see both epiphytic and epizooic algae. For many of them, the epizooic lifestyle is one of opportunism--the algae may not care which animal they live on, or even whether they live on an animal or a rock. Some of the epiphytes, such as Microcladia coulteri, grow on several species of algae; I've seen it on a variety of other reds as well as on a brown or two (feather boa kelp, Egregia menziesii, immediately comes to mind). Smithora naiadum, on the other hand, seems to live almost exclusively on the surfgrass Phyllospadix torreyi.

Animals can also live as epiphytes. The bryozoan that I mentioned last time is an epiphyte on giant kelp. Bryozoans, of course, cannot move once established. Other animals, such as snails, can be quite mobile. But even so, some of them are restricted to certain host organisms.

The aptly called kelp limpet (Discurria insessa), lives only on the stipe of E. menziesii, the feather boa kelp. Its shell is the exact same color as the kelp where it spends its entire post-larval life. Larvae looking for a place to take up a benthic lifestyle settle preferentially on Egregia where adult limpets already live. It's a classic case of "If my parents grew up there it's probably a good place for me."

Limpet on stipe of feather boa kelp
Discurria insessa on stipe of Egregia menziesii
2020-06-07
© Allison J. Gong

The limpets cruise up and down the stipe, grazing on both the epiphytic diatoms and the kelp itself. They can make deep scars in the stipe and even cause breakage. Which makes me wonder: What happens to the limpet if it ends up on the wrong end of the break? Does it die as the broken piece of kelp gets washed away? Can it release its hold and find another bit of Egregia to live on? Somehow I doubt it.

Discurria insessa on stipe of Egregia menziesii
2018-05-16
© Allison J. Gong

The last time I was in the intertidal I encountered another epiphytic limpet. Like the red alga Smithora naiadum, this snail one lives on the narrow leaves of surfgrass. It's a tiny thing, about 6 mm long, and totally easy to overlook, given all the other stuff going on in the tidepools. But here it is, Tectura paleacea. Its common name is the surfgrass limpet, which actually makes sense.

Top view of surfgrass limpet on leaf of surfgrass
Surfgrass limpet (Tectura paleacea) on surfgrass (Phyllospadix torreyi) at Davenport Landing
2020-07-07
© Allison J. Gong

Tectura palacea feeds on the microalgae that grow on the leaves of the surfgrass, and on the outer tissue layer of the plant. They can obviously grow no larger than their home, so they are narrow, about 3 mm wide. But they are kind of tall, although not as tall as D. insessa.

Lateral view of surfgrass limpet on leaf of surfgrass
Surfgrass limpet (Tectura paleacea) on surfgrass (Phyllospadix torreyi) at Davenport Landing
2020-07-07
© Allison J. Gong

Cute little thing, isn't it? Tectura palacea seems to have avoided being the focus of study, as there isn't much known about it. Ricketts, Calvin, and Hedgpeth write in Between Pacific Tides:

A variety of surfgrass (Phyllospadix) grows in this habitat on the protected outer coast; on its delicate stalks occurs a limpet, ill adapted as limpets would seem to be to such an attachment site. Even in the face of considerable surf, [Tectura] palacea, . . . , clings to its blade of surfgrass. Perhaps the feat is not as difficult as might be supposed, since the flexible grass streams out in the water, offering a minimum of resistance. . . The surfgrass provides not only a home but also food for this limpet, which feeds on the microalgae coating the blades and on the epithelial layers of the host plant. Indeed, some of the plant's unique chemicals find their way into the limpet's shell, where they may possibly serve to camouflage the limpet against predators such as the seastar Leptasterias hexactis, which frequents surfgrass beds and hunts by means of chemical senses.

And that seems to sum up what is known about Tectura palacea. There has been some work on its genetic population structure, but very little about the limpet's natural history. The intertidal is full of organisms like this, which are noticed and generally known about, but not well studied. Perhaps this is where naturalists can contribute valuable information. I would be interested in knowing how closely the populations of T. palacea and Phyllospadix are linked. Does the limpet occur throughout the surfgrass's range? Does the limpet live on both species of surfgrass on our coast? In the meantime, I've now got something else to keep my eye on when I get stranded on a surfgrass bed.

Every year, in June, my big whelk lays eggs. I have a mated pair of Kellettia kellettii living in a big tub at the marine lab. I inherited them from a lab mate many years ago now, and they've been nice pets. They've lived together forever, and make babies reliably. As June rolls around I start looking for eggs. This year I want to document the entire process, from egg-laying to larval development. Fortunately, I had the foresight to photograph the parents in May, as I didn't want to disturb the female once she began laying.

The female is significantly larger than the male. I know the big one is the female because that's the one that lays the eggs. I've never managed to catch the whelks copulating, but given the female's track record they either copulate regularly or she is able to store sperm for a long period of time.

In any case, she started laying eggs today. I went in to check on them and there she was!

Female whelk laying eggs
Female whelk (Kellettia kellettii) laying eggs
2020-06-12
© Allison J. Gong

I know from previous years that it can take over a week for the female to lay her entire clutch of eggs. Each of those pumpkin seed-shaped objects is an egg capsule, containing a few dozen embryos. The newly lain capsules are white, as you see above, and will gradually get darker as the embryos develop into larvae. The mother will lay the eggs and then depart. When the larvae are ready to leave the capsule, a small hole will wear through in the top of the capsule and the larvae will swim out. More on that later, hopefully.

I took some time-lapse video of the female, and was able to record her moving over the egg capsules and then leaving. I'd also put some food in the tub, and I think she got distracted.

I think it's really cool to see how well the snail can swivel around on her foot. Snails are attached to their shell at only a single point called the columella, the central axis around which the shell coils. Some snails can extend quite far outside the shell, and they can all pull inside for safety. The dark disc on the back of the foot is the operculum that closes up the shell when the snail withdraws into it.

Tomorrow when I check on things at the lab I'll see if she has resumed laying.

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

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.

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.

Combine the words "gold" and "California" and you automatically come up with the Gold Rush, don't you? After all, California is the Golden State. And while that nickname may be to honor the golden hills of summer or the poppies that are the state flower, it may also be a tribute to the discovery of gold in 1848. For better or worse, the Gold Rush initiated rapid development of this area, and California eventually became the 31st state in 1850.

For me, and I suspect for many people, gold is one of the quintessential colors of autumn. Yet here we are in the middle of winter heading towards spring, and I saw a lot of gold in the forest the other day. I had taken my Ecology students to Rancho del Oso for the first field trip of the semester and set them loose to saunter through the woods and practice noticing (and recording) patterns in nature. Incidentally, I have adopted the word 'saunter' as a replacement for 'hike' for most of my own outdoor adventures. I have always been a slow hiker, and felt that in order to keep up with other people I had to miss seeing what was going on around me. Not to mention the fact that I'm always stopping to take pictures or examine some weird thing on the ground, or in the trees, or wherever. By giving myself permission to saunter along at the pace at which nature occurs, I have time to slow down and observe more carefully, and come away with a much better understanding of the world I've passed through. It certainly doesn't work for everybody, but I've learned that the journey is as important as the final destination, and that has made hiking sauntering much more enjoyable for me.

So, back to the gold. One of the very first thing I noticed when we hit the trail was this brilliant yellow-orange slime mold growing on twigs on the forest floor. This area is a mixed forest of hardwoods (mostly oaks) and various pines. I can't be certain what these sticks hosting the slime mold are, but they may be some kind of pine.

Slime mold
Slime mold, possibly Leocarpus fragilis, at Rancho del Oso
2019-02-01
© Allison J. Gong

Slime molds are very strange organisms that don't fit into any of the major eukaryotic kingdoms of life (Animalia, Plantae, or Fungi). The current taxonomic position of slime molds is up for debate and far from settled, so I won't go into it here. Like fungi, slime molds feed on dead and decaying plant matter and are part of the decomposer niche of organisms. Also like fungi, most of a slime mold's life is microscopic. In the case of fungi most of the body, called a mycelium, is a network of extremely thin threads called hyphae. The mycelium for most fungi is underground and thus invisible to the casual observer. What we call a mushroom is only the reproductive fruiting body, which pushes to the surface so that spores can be released into the air.

For most of the time, or at least as long as food is plentiful, a slime mold exists as single amoeba-like or flagellated cells that feed on bacteria. These cells are haploid, containing only one set of chromosomes. Sexual reproduction (labelled SYNGAMY in the figure below) occurs when an amoeba-like cell encounters a compatible flagellated cell. I would also be willing to bet that the amoeboid and flagellated cells are triggered to find each other and initiate syngamy when food is scarce, as is the case with many animals.

Life cycle of a slime mold
© Pearson Education, Inc.

The result of syngamy in a slime mold is a zygote which develops into a macroscopic stage called the plasmodium. The plasmodium undergoes nuclear division multiple times but cytokinesis doesn't occur, resulting in a large cell bounded by a single plasma membrane and containing many nuclei. In animal tissues we describe this condition as syncytial; I don't know if the same word is used by slime mold specialists, but the concept applies.

One of the things that makes slime molds truly bizarre is their method of locomotion. Using time-lapse videography, you can actually see how the contents of the cell swash back and forth in a process called cytoplasmic streaming. The net result of all this cytoplasmic streaming is the physical movement of the plasmodium into new territory. It's a process much easier to understand if you can see it, so here's a video from KQED's Deep Look series:

As with many fungi, slime molds are difficult to identify if you don't see the fruiting body. The slime mold that we encountered the other day was an immature plasmodium that hadn't yet produced fruiting bodies. The experts who took a look at my observation on iNaturalist agreed that it is likely Leocarpus fragilis, based on location and time of year, but they cannot be certain.

Continuing with our theme of gold, we saw several small blotches of golden jelly growing on tree trunks. These were the Tremella fungi. There are two species of golden Tremella in our region, T. mesenterica and T. aurantia. It seems that differentiation between the species depends on examination of microscopic structures, so I am unable to tell which species this little blob is. However, I will point out that the species epithet aurantia means 'gold', so I really hope that's the name for this blob.

One of the golden jelly fungi (Tremella sp.) at Rancho del Oso
2019-02-01
© Allison J. Gong

Saving the best for last! Moving away from the creek and into the more enclosed forest we entered the realm of everybody's favorite terrestrial pulmonate gastropod, the banana slug. They were out in full force, chowing down on mushrooms and sliming up the foliage. One of my students picked up a banana slug and let it crawl on her hand for a while, but to my knowledge nobody licked one. All of the banana slugs that I saw were bright yellow with no brown or gray blotches, so I conclude that they were either Ariolimax californicus (the so-called Peninsula banana slug) or A. dolichophallus (the Santa Cruz banana slug, also the school mascot for UC Santa Cruz).

Banana slug (Ariolimax sp.) at Rancho del Oso
2019-02-01
© Allison J. Gong

But this is where things get interesting. According to their mitochondrial DNA these two species, A. californicus and A. dolichophallus, do not have overlapping ranges. And the dividing line between them is Rancho del Oso, with A. californicus occurring to the north and A. dolichophallus occurring to the south. So, if Rancho del Oso is the magic line defining the ranges of these two species, what species are the slugs at Rancho del Oso? I think that answering this question will require a much finer scale study. For now, I'm just going to call them Ariolimax sp., because that seems to be the safest option until things get sorted out.

I've written about banana slugs before, but I've never had a chance to photograph them doing the actual nasty. Luckily for me and the students, banana slugs have no shame. I think the entire class got to get a close look and photos of this copulating pair:

Copulating banana slugs (Ariolimax sp.) at Rancho del Oso
2019-02-01
© Allison J. Gong

This perfect yin-yang symbol is the result of how banana slugs align themselves during copulation. Each hermaphroditic slug has a genital open behind the head on the right side of the body. There's a lot of kinky stuff that happens during banana slug sex, including the chewing off of one partner's penis, but suffice to say that one animal's penis is inserted into the vagina of the other and, well, we don't know how quickly sperm is transferred, but the animals remain locked together for several hours. Yes, HOURS. Ahem. The penis chewing thing doesn't happen every time slugs mate, and biologists are still trying to figure out the function for this unusual behavior.

We have another several weeks (hopefully!) of rainy weather, so there will be lots of time to explore the world of fungi, slime molds, and banana slugs. The combination of rain and lengthening days creates great conditions to revel in the gold of a California winter in the forest.

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