It never really feels like springtime until the swallows have returned to the marine lab. This year the barn swallows (Hirundo rustica) came back right on schedule in the last week of March. They have been flying around ever since. I've seen them gathering mud on the banks of Younger Lagoon, but they haven't been very serious about nest building. The cliff swallows (Petrochelidon pyrrhonota), on the other hand, were late arrivals, and I was beginning to worry about them. I didn't see the first cliff swallow until the 1st of May.
Unlike the barn swallows, the cliff swallows immediately started spackling mud on the buildings. There have been a few aborted attempts, but overall they have been very busy little birds. I've been keeping an eye on one nest in particular, as it seems to be the one progressing most rapidly. This nest is located on a building that has been used every year, by both barn swallows and cliff swallows. In previous years I've seen and photographed the nests side-by-side. There isn't any reason to expect them not to nest together again.
So here's what has been going on so far.
The first stage is a simple shelf of mud. The birds are building on vertical walls, but corners where they can use two walls are prime locations. Even a tiny ledge can be used to support those first splats of mud, and once the mud dries it becomes foundation for additional layers. See the mud in the swallow's beak?
Sometimes the birds get this far and then decide to abandon the nest. And sometimes they keep going. Here's what's going on less than a meter away from this nest.
Notice how the birds use their tails as braces so they can cling onto vertical walls. Woodpeckers do the same thing. Of the four birds in the photo above, the two outer ones look like they're just starting to build. I've seen two birds flying in and out of that larger nest, but have no idea whether or not they're the same two birds every time. I suspect they are a mated pair.
The cliff swallow's nest is the gourd-shaped one, with a body that narrows to a much smaller opening just big enough for one bird to pass through. When the nest I was watching got to the stage in the photo above, I thought the opening would be on the left, since there's already a nice curve along that side of the front edge.
I might be kind of right. Today the opening is much narrower, and located off-center towards the left.
This nest may be finished tomorrow or the next day. It will have taken the birds about a week from start to finish.
Why are there no birds near the nest, you may ask? Well, when I walked out of the building I noticed that all of the swallows were in the air, and nobody was at the nest site. The birds were making alarm calls and flying around, but it didn't look like they were feeding, and none were returning to the nests even for short visits. I looked around and up and found the reason for the swallows' unrest.
The hawk was perched directly across from the swallows' nest site, and the swallows were not happy about it.
(For some reason the embedded video isn't working right now. You can watch it from YouTube directly by clicking on that link in the lower left corner.)
Some of the swallows flying around the hawk were barn swallows. I think they are nesting in the breezeway of the building under the hawk. Anyway, the hawk's presence was obviously upsetting to all of the swallows. It hung out for about 10 minutes and then flew away over the field. And immediately, the cliff swallows got right back down to the important business of building a home for the next generation.
A few weeks ago I went out to Franklin Point and saw that the sea lettuces (Ulva sp.) were spawning in the high pools. I revisited the site today, with a more lower tide to work with, and spent a considerable amount of time looking for and photographing the staurozoans. I did find some, too! But they are not the focus of this post.
As the tide came back in, I spent more time working my way through the higher pools. At Franklin Point there are very few places where the water is still. Even in the high regions the intertidal terrain is more surge channels than pools. But if you go high enough up the beach there are some quiet areas where the water, if it moves, does so very slowly. It is in these areas where the algal spawn forms those beautiful patterns that I photographed at the beginning of the month. Today there was much less algal spawn accumulating in the calm areas. It was also windy (and cold) this morning, so the patterns were not as crisp as they had been in early April. Still pretty, though!
On my way back up the beach I saw something that looked like an iceberg viewed from the air.
This is an accumulation of foam being pushed ashore. I didn't have any way to collect a sample to bring back to the lab for closer observation, but foams like this are usually due to algal particulates. Surface agitation whips up the organic matter, which act as surfactants and produce tiny bubbles. I'd be willing to bet that the Ulva spawn is at least partly responsible for this foam.
I watched the foam for several minutes, and was rewarded for my vigilance. I found an area where the highest reach of the incoming tide was gently washing back and forth.
I found the slow swirling to be rather mesmerizing. Maybe that was due to the early morning, the brisk sea air, or hunger pangs. But when I saw this I thought to myself, "I've seen that somewhere before." You might be able to guess where.
To validate my intuition, when I got home I looked up some images and found that I was sort of right after all.
Okay, so maybe the resemblance isn't as strong as all that. But I can still imagine the streams in van Gogh's painting swirling and flowing the way the algal foam does. What do you think?
For animals that do essentially nothing when you see them where they live, chitons have a lot of charm. They are the kind of animal that, once you develop the search image for them, you start seeing everywhere. It helps that they are easily recognized as being chitons because of their eight dorsal shell plates—nothing else looks like them. Depending on species, those shell plates can be smooth or sculpted, and pigmented or not. Patterns of sculpting and pigmentation (or lack thereof) are diagnostic features used to distinguish different species. Some species are reliably consistent in appearance and look the same wherever you happen to see them. Other species show a lot of phenotypic variation, often even at a single site.
One of my favorite chitons is Mopalia muscosa, the mossy chiton. It's one of the easiest of our chitons to identify, because its girdle (the layer of tough tissue in which the shell plates are embedded) is densely covered by long, curved spines. They're called spines, but they're quite soft and flexible. Your basic Mopalia muscosa looks like this:
Mopalia muscosa is one of the species whose appearance is quite variable. Many of them wear algae, usually reds but occasionally greens or browns, on their shell plates. Not all species of chiton do this. I've often wondered why some chiton species wear algae and others do not. This individual is probably fairly old, judging by the worn condition of the shell plates. The plates show signs of erosion, but are not decorated. There are some small pieces of coralline algae amongst the spines of the girdle, though, which I always associate with age. Smaller, and presumably younger, M. muscosa tend not to have algae on the girdle even if they are wearing some on the shell plates.
The degree of shell decoration in M. muscosa varies from none, as above, to heavy encrustation. This individual below has been colonized by only a small bit of coralline algae and perhaps some brown diatom-ish film on the edges of the shell plates:
This next one has only a small bit of coralline alga, but sports a jaunty sprig of something quite a bit larger.
This season's fashionable chiton will go all out with the coralline algae, wearing both encrusting and upright branching forms. Look at this:
Sometimes the chitons wear the larger leafy red algae, in addition to or in place of the coralline algae. I always think that these individuals must be very old, by chiton standards.
And sometimes the chitons are so covered with algae that they blend in perfectly with the surrounding environment.
These chitons can get very heavily fouled by algae. Is there any benefit to the chiton, to carry around a load of red algae? And if wearing algae is for some reason advantageous, is there a way for a chiton to attract algae to settle on their shell plates? Well, let's think about that. Chitons' main predators would be sea stars, crabs, and birds. Sea stars do not locate prey visually, so camouflage would not be very helpful in avoiding them. Birds such as oystercatchers and surfbirds certainly do pry up chitons and limpets, and blending in with the background just might help a chiton go unnoticed by an avian hunter.
Regarding the matter of how the algae end up living on chitons' bodies, I want to start with the question of how prevalent algal fouling is on Mopalia muscosa, and the extent of fouling on the chitons that are wearing algae. A little research study might be a fun way to spend my time in the intertidal. Pigeon Point is a lovely site on a foggy summer morning, and many of the most heavily decorated M. muscosa in my photo library are from there. Yes, I can foresee several visits up the coast over the next few months. Laissez les bons temps rouler!
This morning I went to Pigeon Point to poke around and do some collecting. It's a favorite site of mine, as it's exposed and dynamic, with the diversity you'd expect. Of the sea stars, the most common by far are the six-armed stars in the genus Leptasterias. They are small (less than 8 cm in diameter, often smaller than 1 cm), somewhat drably colored, and sometimes on the underside of rocks, all of which means that they are not always conspicuous. But once you get the right search image, you see them everywhere.
Six arms, see?
Sea stars are well known for their ability to regenerate lost arms. It is not uncommon to see a star that looks healthy in every way except that one of its arms is shorter than the others. This must happen in Leptasterias, too. Searching through my library of pictures of Leptasterias, I did find a couple of examples of regeneration.
When these stars finish regrowing those arms, they will have the typical number of arms for the genus, which is six.
Today I saw something that I'd never seen before. It was a Leptasterias that was regenerating arms. Only this was weird. It had three full-size arms and was growing four!
When (if) this star survives, it will eventually have seven arms. And that's strange. I asked my friend Chris Mah, who is the sea star systematist at the Smithsonian, if it was common for Leptasterias to do this. He said he'd never seen it, either. So it is indeed a rare phenomenon.
Now, there are stars in the genus Linckia that actually reproduce by deliberately leaving behind an arm, which then goes on to regenerate the rest of the body. While they do so they look like comets:
My regenerating Leptasterias isn't quite a comet, but it is doing something equally strange and wonderful. I really wished I could bring it to the lab and keep an eye on it over the next several months. However, Leptasterias are on the no-take list, I think because their populations are so patchy. It is extremely unlikely that I will ever see that same individual again, so we will never know what happens to it. Unless, of course, I happen to come across a 7-armed Leptasterias at Pigeon Point sometime in the future. If you see it, take a photo and let me know!
I had seen the sea lettuces (Ulva spp.) spawning in these high pools at Franklin Point before, and usually cursed the murkiness of the water. But today the water was dead calm, with the tide low enough that there were no waves to slosh into the pools. The result was a gorgeous marbled swirl in the water. The patterns were stunning.
What these photos show is the Ulva releasing either spores or gametes. Without microscopic examination it's impossible for me to know whether these tiny cells are spores or gametes. What I can say is that the spawn is released from the distal ends of the thallus, making the body of the alga look ragged.
The parts of the thallus that have already spawned are now clear. The tissue itself will soon disintegrate, leaving behind only the healthy green parts, which should be able to regrow.
All of these photos were taken in pools where the spawning itself had either completely or mostly stopped. Obviously when the tide comes back all of this yellow spooge will get mixed up. It's only when the water is perfectly still that these streams would form. It was hard stepping around the pools to take the photos, as the last thing I wanted to do was stomp my big booted foot into a pool and disrupt the beautiful patterns. Fortunately the sun angle was a little cooperative this morning, and I was able to find a pool where active spawning was happening.
What appears to be an act of destruction—the alga's brilliant green thallus being reduced to yellow streaks that drift away with the tide—is really an act of procreation. This is terminal reproduction, literally the last thing an organism does before it dies. Salmon do this, as do annual plants. The sheer amount of algal spawn in these tidepools is astounding. Imagine the number of 2-micron cells needed to color the water to this degree. But if reproducing is the last thing you're going to do in your life, you might as well go all in on your way out, right?
The other day I was on a field trip with a couple of students in the Natural History Club, at Younger Lagoon. We had permission to go down into the lagoon itself, where we chased tiny red mites around rocks in the intertidal without getting caught by waves, observed a very interesting interaction between a coyote and assorted water fowl, and witnessed killdeer mating. Did you know that in killdeer the actual copulation is preceded by about half a minute of massage? Neither did we! The purpose of the field trip, other than merely to be outdoors looking at cool stuff, was to spend some time doing focused nature journaling. As a result I didn't have my big camera with me. But I did have the good binoculars, and got to watch all of the action closely.
Nature journaling should be part of any natural history club. Over the years I have seen an increase in the tendency to equate nature journaling with science illustration or other types of art. This conflation is what causes people to believe that they can't keep a nature journal because they don't produce museum-quality works of art. While I appreciate a beautiful science illustration as much as anybody else, a nature journal serves a completely different purpose. A nature journal's job isn't to be beautiful. Its job is to be informative.
If you were to compare my nature journal entry with a photograph of the site, you would see that my sketch is nowhere near realistic in the sense of looking exactly like the real thing. I've compressed the entire lagoon into a short stretch that I could fit in these two pages. But I think the sketches and notes do convey the fascinating things what we saw that day. And even if I were not familiar with Younger Lagoon, I would be able to look at these pages and remember them. That's the job of a nature journal.
I returned to Younger Lagoon two days later with the camera in tow, hoping that some of the birds we'd seen on Monday would still be there on Wednesday. In addition to the usual Canada geese and mallards, I hoped to shoot a couple of water birds that I didn't recognize.
Let's start with the obvious:
All told, there were a couple dozen Canada geese, in the water, in the air, and on the sand. They were a noisy bunch, as usual. Except for when the coyote showed up. Read that little story in my nature journal.
Now take a look at these geese:
See the one goose that doesn't belong? That was the mystery goose I saw on Monday, and was fortunate enough to see again on Wednesday. From the photos in my bird field guide—National Geographic Field Guide to the Birds of North America—I thought it might be a greater white-fronted goose (Anser albifrons), although I couldn't be entirely certain. I knew I hadn't seen one before, but a consultation with Cornell's All About Birds verified the ID. iNaturalist shows only a handful of observations of A. albifrons in the Monterey Bay region. The greater white-fronted goose is a long-distance migrator, breeding on the tundra of the high Arctic and overwintering in California's Sacramento and San Joaquin Valleys and the Gulf coast of Texas and Louisiana.
A third goose, and another winter-only bird, is the snow goose. It is a little bigger than the greater white-fronted goose. While the word "snow" implies white plumage, snow geese also come in a blue form, which is a dark blueish gray with a white head. The blue coloration is due to a single gene, and the allele for blue is incompletely dominant over the allele for white. The blue and white morphs are the same species and interbreed freely. The offspring of a pure blue bird and a pure white bird will be dark, but may have a white belly. Goslings from pure white parents will be white, and those from pure dark parents will be mostly dark but may have some white.
Of the two snow geese in the photo, the one in the front is all white except for the black wing tips of the species, while the one in the back has more dark coloration. In the photo the beak looks dark, but in better light it's as pink as on the bird in the front.
So that's three species of geese. Now whose butts are these?
These tails belong to American wigeons (Mareca americana), a male and female pair in the background and a lone male in the foreground. As you might guess from the behavior, wigeons are dabbling ducks, foraging on aquatic vegetation. Like the greater white-fronted goose and snow goose, these are also winter visitors to California's waterways, and will soon be headed north.
In their winter plumage, the wigeons are rather dull. The breeding male has a brilliant green patch extending backwards from his eye and a broad white streak from the top of the bill over his head. During the winter the green patch becomes is much less conspicuous, although the white streak remains.
Three species of waterfowl. I couldn't get the snow geese to cooperate and make up the quartet.
Living as we do along the Pacific flyway, we find that spring and autumn are great times for watching birds as they migrate between summer breeding grounds and wherever they overwinter. Sometimes I think it's rather unfortunate that I don't get to see these birds in the glory of their breeding plumage, but that's okay because I get to see them in the winter. And the birds that left here for the winter are returning: I saw the first barn swallow of the season right after the vernal equinox! Soon they and the cliff swallows will be building their nests on the buildings at the marine lab. At home, the first of the season's hooded orioles flew past the back deck. He may have been on his way to a nesting site in a palm tree down the street. There is so much going on right now. I do love the spring!
A utility pole across the street and one house down has, for years, been an object of interest for a variety of birds. The hairy and downy woodpeckers drum on it in the spring, and various songbirds hang out and rest on the top. About a month ago now I saw a raptor up there, eating something. It was a female merlin (Falco columbarius). According to Cornell's All About Birds, merlins are in our area during the nonbreeding season, but I've never been certain about having seen one.
On the morning of Saturday 13 March I went outside to look around, and saw a bird on the pole. It appeared to be either eating or cleaning its beak. I ran inside to grab the camera, which fortunately had my longest lens and the 1.4x teleconverter attached, and snapped off a bunch of shots. The sun was rising, but I was able to get some decent photos of the bird even though from the best vantage point it was backlit.
Clearly, he's eating something:
But what is it eating? Rodent bits?
No, look at that foot. It's a bird!
Yep. Definitely a bird.
And here he is, taking a break between courses:
Merlins are members of the falcon family. Smaller birds make up the majority of a merlin's prey, but they also eat large insects such as grasshoppers. As with peregrine falcons, merlin populations were severely reduced in the years when DDT was widely used to keep insect populations down, but they have since recovered. Truly, the recovery of birds of prey after DDT was banned is one of the great successes of conservation biology.
There were feathers in the street below the pole. I assume they are from the merlin's prey, as when I looked at the top of the pole through binoculars I could see the same sort of feathers up there. I compared the feathers with photos on a few ID sites, but it's no easy identifying feathers without any additional context. Someone suggested that they might be from a male house finch. We have lots of those around all the time, so that's probably the best guess possible.
So there you have it: Saturday brunch with songbird on the menu!
It has taken me months to gather all the photos and videos I needed for this post. I could blame it on the stress of teaching online for the first time, the COVID-19 pandemic itself, or residual malaise from the dumpster fire that was 2020. But really, it's the animal's fault.
In this case the animal is the orange cup coral, Balanophyllia elegans. I've written about this beast before, and lately I've been paying more attention to the corals that we have in the lab. In many ways it is the typical anthozoan—its life cycle consists of only a polyp stage (i.e., no medusa), it is benthic, and its body is vaguely anemone-like. Like the reef-building corals of the tropics, Balanophyllia is a scleractinian coral. This means that it secretes a calcareous base, or exoskeleton, upon which sits the living tissue of the polyp. But unlike the reef-building corals of the tropics, Balanophyllia is solitary, which means that it does not clone or form colonies. Nor does it contain photosynthetic zooxanthellae in its cells, as the reef-builders do. This means that Balanophyllia is a strict carnivore: unable to rely on photosynthetic symbionts to do the hard work of fixing carbon, Balanophyllia has to catch its own prey.
Balanophyllia has separate sexes and reproduces sexually. Males release sperm that are ingested by the female, and she then broods larvae for several months. What is strange about this is that, when you consider the anatomy of Balanophyllia (or any cnidarian polyp, for that matter) the question that comes to mind is, "Where are the larvae brooded?" So let's take a look at the basic cnidarian body plan.
The professor who taught my undergraduate course in invertebrate zoology described a cnidarian's body as a baggie inside a baggie, with a layer of jelly between the two baggies. The inner and outer baggies represent the tissue layers that developmental biologists refer to as endoderm and ectoderm, respectively. The jelly between the two tissue layers is called mesoglea. For the sake of this discussion, it's the endoderm that interests us. In a cnidarian, the endoderm is also called gastrodermis, because it is, literally, the skin of the digestive system. The cnidarian gut is a cavity that opens to the outside via a single opening. We could call that opening either a mouth or an anus, since both food and wastes pass through it, but for politeness' sake we call it a mouth. The gut itself does double duty as both the digestive system and the circulatory system, so its formal name is gastrovascular cavity (GVC).
It's easy to imagine the GVC as being essentially a tubular vase, but it's more complex than that. The GVC extends into each of the tentacles, so the tentacles are actually hollow. The main cavity of the GVC, in the stalk of the polyp, is partitioned by thin layers of endoderm. The partitions are called septa, and serve to increase the surface area of the endoderm for digestion. Think of a large office building, divided into small cubicles by movable partial walls—there's much more total wall space for hanging things such as calendars than there would be in the big room with only four walls. In anthozoans, gonad tissue is associated with the septa.
Now back to how Balanophyllia broods its larvae. As we've seen before, Balanophyllia's planula larva is an orange-reddish wormlike blob about 2 mm long, which is ciliated all over. It doesn't feed, but survives on energy reserves supplied by the mother in the egg; it may also be able to take up dissolved organic material directly from the seawater. After brooding larvae for some period of time in their GVC, Balanophyllia females release larvae. Or rather, the larvae ooze out of their mother's mouth and crawl around to settle and metamorphose nearby.
The planulation season, when larvae show up amongst the corals we keep in the lab, begins in the fall and runs through winter into the spring. Sometimes the larvae metamorphose right away—one day there are no larvae and the next day there are baby corals. Other larvae squirm around for days or weeks, not getting any smaller but not metamorphosing, either. This past season (Fall 2020-Spring 2021) we saw both extremes: planulae that settled and metamorphosed right away, and others that I collected weeks ago and are still worming around.
I peered into a bunch of corals, hoping to see what the larvae are doing inside the GVC, with no luck. Then I decided to try some time-lapse video under the dissecting scope, and had a bit of success with that. In the video below you'll see about two hours of action compressed into about 1 minute. Watch for a small red ball squirming around inside the GVC. And remember that the GVC extends into the tentacles, so the larva can wiggle its way in there, too.
All of which makes me wonder if the pathway from mother's GVC to the scary world outside travels in only one direction, or if the larvae can retreat back into the safety of the mom's digestive system. How strange is it, that the safer location might be inside the mother's gut!
Eventually, whether it be a matter of hours or weeks, the planula larva settles (i.e., sticks to a surface and stops crawling) and metamorphoses (i.e., makes the anatomical and physiological changes into the juvenile body). There doesn't seem to be a clear connection between surface characteristics and whether or not larvae will settle there. You might think that they'd choose to settle nearby or maybe actually on the parent, but that's not always the case. At some point, however, the larva will have to either settle and metamorphose, or die. When they metamorphose into juveniles, they reveal other aspects of their nature as scleractinian corals.
From what I've seen, the larva begins the settlement process by sticking to a chosen spot and becoming a more circular (i.e., less elongated) blob. On the other hand, I've seen perfectly round red blobs that look for all the world as though they might be pushing out tentacles, change their minds and go vermiform again. But once they stick permanently and begin to metamorphose by forming the polyp's body, they have to continue.
The juvenile in the photo above has not yet pushed out any tentacles, but it does show signs of its scleractinian ancestry. The Scleractinia (stony corals) and Actiniaria (sea anemones) are both members of a group called the Hexacorallia. The Hexacorallia and Octocorallia are in turn grouped together as members of the class Anthozoa. As you might infer from the name 'Hexacorallia', animals in this group of a body symmetry based on the number six. And you can see in the photo above that the juvenile coral's body is divided into 12 wedges. When the juvenile begins growing tentacles and a more polyp-like body, the hexamerous symmetry becomes even more evident:
The young coral begins by forming six primary tentacles, which establishes the visible hexamerous symmetry. Then it grows the six shorter secondary tentacles. The bumps on the tentacles are cnidocyte batteries, clusters of stinging cells, indicating that the animal can already capture and kill prey. Good thing, too, because if it hasn't already then it soon will deplete the energy stores its mother partitioned into its egg.
By this stage in its development the coral will remain where it is for the rest of its life. As it grows it will deposit CaCO3 and grow taller, but the living tissue will always be restricted to a layer sitting on top of the calcareous base. Once the base grows more than a few millimeters tall, there is no living tissue in contact with the surface. Thus there is no mechanism for the animal to move to another location, or even to re-attach itself if it gets broken off its surface. There are many animals that are likewise unable to move once their larvae have attached and metamorphosed. The decision of where to put down roots becomes quite stark for them, as a bad one can result in a very short life indeed. The selective pressures that enforce a good decision are quite clear, and are important factors in the distribution patterns we observe in the intertidal.
For several weeks now I've been raising another batch of bat star (Patiria miniata) larvae, from a fortuitous spawning that occurred in early January. Since this is rather old hat by now I'm not diligently taking photos or drawing the larvae as often as I would have years ago when this kind of undertaking was new to me. But I still change the water twice a week and look at them on Fridays, and I still have the set-up that attaches my old phone to the microscope so I can take pictures of them.
Last Friday it occurred to me that: (A) my gizmo holds the camera steady over the microscope, so I can take pictures at multiple focal planes within objects under the scope; and (B) I have software that will stitch those many snapshots into a single image. Neat!
So I made this:
This larval stage is called a bipinnaria or a brachiolaria. From top (anterior end) to bottom (posterior end) the larva is about 1 mm long. It swims with the anterior end in front. In some sea stars the bipinnaria grows long arms, at which point we call it a brachiolaria ('brachio' = 'arm' in Greek). Bat stars don't grow long arms, so the distinction between bipinnaria and brachiolaria is much fuzzier.
I took 11 photos of this larva, each one focused on a different horizontal plane, and did a focus merge in my photo processing software. Crossed my fingers as the software did its magic, and then peeked at the result. It worked! When looking through the microscope I have to focus up and down through the body to get an idea of its three-dimensional structure. But if the animal holds still long enough, I can do the focus merge thing and get images like this one.
And that slight halo that you see around the exterior surfaces of the larva? That is not an artifact of the photo taking or processing. That halo is due to the cilia that cover the body. There is a ciliated band, which you can see as the dark gold ribbon that snakes along the lobes of the body, and the other body surfaces are ciliated as well. The ciliated band is what the larva uses to swim through the water. Each photo freezes the ciliary action at the moment it was shot, but stitching several photos together causes the cilia to blur into that pale halo.
Intact shells are a limited resource in the rocky intertidal. Snails, of course, build and live in their shells for the duration of their lives. A snail's body is attached to its shell, so until it dies it is the sole proprietor of the shell. Once the snail dies, though, its shell goes on the market to whoever manages to claim it. Empty shells tend not to remain on the market for long.
Hermit crabs also live inside snail shells. They are the ones that compete for empty shells when they do become available. Here in California, at least, the hermit crabs can't kill snails for their shells; they have to wait for a snail to die. And once a shell comes on the market, it will have a taker even if it's not the ideal size for the crab. It's not at all uncommon to see hermit crabs that can fit only their abdomen into the shell, leaving the head and legs exposed and vulnerable. On the other end of the spectrum, many hermit crabs are so small that they can pull into the shell and not be seen by an inquisitive tidepool visitor. Anybody taking a snail shell home as a souvenir—where such takes are allowed, of course—must be certain that there is no tiny hermit crab hiding deep in the depths.
From a hermit crab's perspective, the best shell is one that is big enough to retreat into but light enough to be carried around. Snail shells come in a variety of shapes and corresponding internal volumes. Turban snails, with their roughly spherical shape, have a large interior space and are coveted by larger hermit crabs. For example, the grainy hand hermit crab (Pagurus granosimanus) seems to really like both black and brown turban snail shells.
Original inhabitant and builder of the shell:
And opportunistic second inhabitant of the same type of shell:
Other snails are not even remotely spherical. Olivella biplicata, for example, is shaped like the pit of an olive. Unlike Tegula, of which both intertidal species are found in rocky areas, O. biplicata burrows in sand. Note the shape and habitat of this olive snail:
These olive snails have a smaller internal volume, and thus tend to house smaller hermit crabs. Young individuals of P. granosimanus can be found in olive snail shells, but they quickly outgrow the cramped quarters and need to find a larger home. Smaller hermits such as Pagurus hirsutiusculus, though, are often found in olive shells.
Any hermit crab that finds itself robbed of its snail shell has a short life expectancy. The front end of the hermit resembles the front end of any crab, with the familiar armored legs, claws, eyestalks, and antennae. But the abdomen is soft and unarmored, covered by only a thin cuticle. The abdomen is coiled to follow the coiling of the snail shell, which allows the crab's body to curl around the columella, the central axis around which the shell spirals. In this way the crab can hang onto its snail shell and resist a tug by a would-be predator. A strong enough tug, though, will rip the crab's front end (head + thorax) away from its abdomen. So if you ever find yourself with a hermit crab in hand, do not be tempted to remove it from its shell by yanking it out!
The next time you encounter gastropod shells in the tidepools and want to know whether the inhabitant is a snail or a hermit crab, watch to see how it moves. Hermit crabs scuttle, as crabs do, while snails glide along very slowly. You would also notice a difference as you pick up the shell: snails stick to the rock with their foot, which you will feel as a suction. Hermit crabs don't stick at all, so if the shell comes away easily it likely houses a crab instead of a snail. See? Easy peasy lemon squeezy!