Christie Willcox wrote a nice article this week on how one small group of organisms called "vertebrates" first evolved to live on land. Since you are a vertebrate who lives on land, you should probably go and read Christie's piece. I wouldn't want you, however, to go around thinking those first fish to leave the ocean behind were pioneers making a uniquely difficult transition. By my figuring, onycophorans (velvet worms like peripatus), tardigrades, annelids, nematodes, nemerteans (ribbon worms) and quite a few arthropod lineages have also taken up a terrestrial lifestyle. Many of those lineages were already breathing air before Tiktaalik, Ichthyostega and your other long-lost relatives came along to join them on land. But if you want to talk about transitions from marine to terrestrial lifestyles then you really want to talk about snails. You can find snails living in  almost every habitat between the deep ocean and the desert, and snails have adapted to life on land many different times. In fact, a litre of leaf litter taken from a New Zealand forest can contain snails representing three separate transitions from water to land.

Almost all the land snails I've talked about here at The Atavism are descendants from just one invasion of the land. We call these species the stylommatophorans and you can tell them from other landlubber-snails because they have eyes on stalks (as modeled here by  Thalassohelix igniflua):

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Most of the description of Cyclophoroids here is taken from:

Barker, GM (2001) Gastropods on land: phylogeny, diversity and adaptive morphology In Barker (Ed.),  The biology of terrestrial molluscs (pp 1—146) CABI Publishing.

Labels: molluscs, native snails, photos, sci-blogs, science, sunday spinelessness

Markus Pfenninger and his colleagues asked just that question by looking at snails from the Northern Hemisphere genus Trochulus (doi: 10.1186/1471-2148-5-59). This genus contains many species that sport very fine and soft hairs. Pfenninger et al.collected ecological data for each species, and used DNA sequences to estimate a the evolutionary relationships between those species. From these data, they were able to infer the common ancestor of modern Trochulus species was probably hairy, and three separate losses of hairyness can explain all the among-species variation in this trait. Moreover, it appears the loss of hairs in Trochulus is associated with a switch for wet to dry habitats. Given this finding, Pfenninger's team hypothesised that, in Trochulus at least, hirsute snails might stick to host plants more effectively than their bald brethren. Indeed, in experiments it took more force to dislodge a hairy shell from a wet leaf than non-hairy one.

Labels: molluscs, native snails, new zealand, sci-blogs, science, sunday spinelessness

When I tell people I study snails for a living I get one of two replies. There's either some version of the "joke" that goes "that must be slow-going" or "sounds action packed", or there's "oh, you mean those giant killer ones we saw when we went tramping?". I guess the joke is funny enough, but I want to make it clear that those giant killer snails from the family Rhytidae, cool as they might be, are not the most interesting land snails in New Zealand.

The local land snail fauna displays a pattern that is quite common for New Zealand animals - we have a very large number of species but those species are drawn from relatively few taxonomic families. Since taxonomic groups reflect the evolutionary history of the species they contain, that pattern most likely arises because New Zealand is (a) quite hard to get to, so few would-be colonists make it here and (b) full of ecological niches and geographic pockets that can drive the formation of new species. In total, there are are probably about 1200 native land snail species in New Zealand - about ten times the number found in Great Britain, which is approximately the same size. That diversity extends to the finest scales - individual sites in native forest might have as many as 60 species sharing the habitat. New Zealand forests probably have the most diverse land snails assemblages in the world (although tropical ecologists, who generally hold that diversity in terrestrial habitats almost invariably increases as you approach the equator, have argued against this conclusion).


You may now be asking why, if this land snail fauna is so diverse, have you never seen a native snail. Well, you've probably walked past thousands of them without noticing. Most of our native land snail species are from the families Punctidae and Charopidae, groups that are sometimes given the common name "dot snails". Meembers of these families are usually smaller than 5 mm across the shell, and are restricted to native forest and in particular to leaf litter. But in native forests, where there's leaf litter there's snails. Grab a handful of leaves, or pull up a log and you're likely to find a few tiny flat-spired snails going about their business. Hell, down here in Dunedin you can even find charopids living under tree-fuschia in a suburban garden.


Like so many native invertebrates, we know very little about our land snails. Lots of people have dedicated substantial parts of their lives to documenting and describing the diversity of these creatures, but even so we don't have a clear understanding of how the native species relate to each other or to their relatives in the rest of the world, or even where one species starts and another ends. Without such a basic understanding, its very hard to ask evolutionary and ecological questions about these species, so for now we remain largely ignorant of the forces that have created the New Zealand land snail fauna.


For the time being I can tell you that a lot of them are really quite beautiful. Since most people don't have handy access to a microscope to see these critters, I thought I would share a few photos from this largely neglected group over the next few weeks. The 2D photographs, with the relatively fine depth of field, don't quite record the beauty of these 3D shells, but I hope it's at least a window into the diversity of these snails.


 Let's start with a snail that is very common in Dunedin parks and forests. This is a species from the genus Cavellia (the strong, sine-shaped ribs being the giveaway) but I won't be able to place it to species until a new review of that genus is published. 




This particular shell is from an immature specimen, and is about 2mm across. When flipped, you can see an open umbilicus that lets you see straight through to the apex of the shell.

Labels: molluscs, native snails, photos, sci-blogs, snails, sunday spinelessness

One awesome mollusc deserves another, so let's follow up last weeks octopus post with one on that group's close relatives the cuttlefish.

Cuttlefish are relatively small (the largest grow to 50cm) squid-like cephalopods that present a nice soft and digestible meal to predatory fish and marine mammals. Having lost the shell that most molluscs use to protect themselves cuttlefish have had to develop other defences. Most strikingly, cuttlefish are masters of camouflage


Just this week, researchers have reported evidence for a other trick that cuttlefish can pull off. When males of the Austrian Mourning Cuttlefish (Sepia plangon) see a female they put on a show, producing striped patterns that evidently impress the female. But these animals form male-dominated groups, and rival males often interrupt would-be woo-ers  in mid-display. So, when they spy a receptive female, males want to put on their flamboyant show for her to judge, but also want to make sure they don't attract the attention of rival males that might want to spoil the party. The male Mourning Cuttlefish's answer to this problem? Using only half of his body to put on the female-impressing show, and throwing would-be spoilers off the scent by mimicking a female with the other half.




This gender-splitting  tactic seems to be pretty common. In aquarium experiments about 40% of males would  attempt the deceptive signal when they were displaying in the presence of a rival. Just as the cuttlefish camouflage response requires information from the physical environment, the gender-splitting trick is influenced by what the male can learn of the social environment. If more than one female is available the male will display to both  without bothering to hide his intentions for observers (probably because working out an angle from which he could excite two females while staying under the radar is just not possible). Likewise, if more than one rival male is about that don't bother with the deception - since it wouldn't be possible to maintain the illusion for two rivals viewing from different positions.

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Brown, Garwood & Williamson (In press) It pays to cheat: tactical deception in a cephalopod social signalling system. Biology Letters. http://dx.doi.org/10.1098/rsbl.2012.0435w

Labels: cephalopods, molluscs, sci-blogs, science, sunday spinelessness

I don't want to talk too much more about the purpose of the argonaut shell, partly because it has already been well covered. Ed Yong wrote a predicably clear and interesting post on the research which uncovered it (which also produced an interesting comments thread) and the lead researcher, Julian Finn from Museum Victoria in Australia, also discussed his work in a really great video.


Instead, I want to talk about the origin of the argonaut shell. Octopuses are molluscs, part of a group of soft-bodied animals that includes clams and mussels and snails. Most molluscs have shells. In fact, despite being arugably the most morphologically diverse of the 35 animal phyla, only a few small groups of molluscs don't contain at least some species that produce shells. The easiest way to explain the presence of shells in so many different molluscan groups is to hypothesize that the last common ancestor of all molluscs had a shell, and most of that ur-mollsuc's descendants have retained this organ. 

In evolutionary biology we call traits that are shared between organisms as a result of their shared evolutionary history "homologies". Homologous traits are often compared with "analgous" ones, parts of organisms that are similar as the result of independent innovations in different evolutionary lineages. We can illustrate the concept using a bat's wing as an example. The forelimbs of bats and whale are made up of the same bones, despite the fact that whales swim and bats fly. That's because bats and whales are both mammals, and they inherited their forelimb bones from a common ancestor before each group radically repurposed their limbs. On the other hand, despite the fact that both bats and stoneflies fly, the insect wing and the bat wing are separate evolutionary inventions and not something the two groups share as a result of shared evolutionary history:

Labels: argonauts, cephalopods, molluscs, sci-blogs, sunday spinelessness

It would take the most dedicated reader of The Atavism to remember the empty snail shell I wrote about last year. I'll admit even I'd mainly forgotten about myself, but this weekend I went on a little mini-field trip to collect a few samples for a colleague's ongoing project. In planning that trip I did remember the slightly mysterious shells I found last winter, and so decided to head back and see if I could get a few more to send along an an expert who might be able to put a name to them. 

Sure enough, I found plenty more empty shells in different states of aging , but deep within the leaf litter I also uncovered one shell that was still playing house to an animal. I couldn't quite be sure there was a healthy animal in the shell when I first picked it up, since the snail was already retracted inside. Thhe easiest way to encourage a sleeping snail out from its shell is to warm in up, so I clasped it in my palm for about a minute and, well, here's the result:

Labels: Dunedin, environment and ecology, molluscs, native snails, photo, sci-blogs, snails, sunday spinelessness

Labels: chemistry, chitons, molluscs, photos, sci-blogs, science, sunday spinelessness

I was going to start this post by saying taxonomy has a language of its own, but that's not really true. Taxonomy just has a whole lot of Latin. When I've given talks in schools, I've almost always been asked why scientists insist on giving creatures those strange Latinised names. The answer is, we need names that anyone anywhere can recognise and pin to particular species or group. A common name like "black bird" might make sense in conversation in New Zealand, but in other circumstances it could refer to a single species (Turdus merula), the five species that make up Turdus, or a whole bunch of species in the new world family Icteridae. When taxonomy really got started Latin was the language which scientists from different countries used to speak to each other, so modern taxonomy follows Linnaeus in using Latinised names.

But the Latin terms used by taxonomists don't stop with names, if you thought they were difficult to understand you should open a taxonomic work. The first time I did I was lost in a sea of lecto-, neo- and allo-types; homonyms junior and senior and Nomina nuda, obltum and protectum. The title of this post, Incertae sedis is another of those strange taxonomic terms and translates as "uncertain placement". Most big taxonomic reviews include a few species the author is certain are well defined, but hard to place into a higher group. Those species get placed under the heading Incertae sedis.

I used that title because today's subject is... a land snail... of some sort

I found this guy (actually, all land snails are hermaphrodites, so I need a better familiar term for them) in our garden and it's pretty tiny (this is a 50 cent coin, almost exactly the same size as a quarter for North American readers):

Labels: environment and ecology, molluscs, new zealand, photos, sci-blogs, sunday spinelessness

It seems all the cool bug bloggers have escaped to the tropics just at the time I've got back to more temperate climes. I spent a couple of weeks in Vanuatu over the Christmas and New Year break and have a couple of memory cards full of Melanesian wildlife to share here over the next few weeks. I have another post based on life in Vanuatu I really want to finish editing before I talk about those bugs, and no shortage of real work to do before I can get to that. So, let's kick of a new year of spinelessness with a lame joke:

(That's a very sick Achatina fulica, one of the villains in this story and pretty common snail in and around Efate, the most populous island in Vanuatu. You can bet this very picture is going to show up as a slide is all my talks about Pacfic Island snails.)

Labels: environment and ecology, molluscs, Pacific, sci-blogs, snails, sunday spinelessness, vanuatu

I don't use these pages to write about my own work very much, partly because it's not yet published and partly because I write about that all day as it is. The shortest answer I can provide to the question "what do you do" is "I use genetic tools to study evolution" and I guess that makes me an evolutionary geneticist. You can split the people that work in our field into two groups: there are biologists that are really interested in a group of organisms and have learned some genetics to help their study of them, and there are people who are interested in a particular question and have chosen their study organisms to suit. I'm very much of the second sort, and like most people in that group I've caught myself saying "I'm interested in the questions, not the animals". Paraphrased, that becomes something like, "Oh sure, I study Pacific land snails, but for all I care they're just little bags of genes that help me answer questions". But that's a lie. You can't work on animals without having them effect you. When I started my PhD I had no particular love of snails, but now I'm a complete snail fan-boy and I frequently find myself preaching on the wonders of life as a terrestrial mollusc to people whose only mistake was to ask me what I do for a living. Did you know most slugs retain the remnants of their shells? Or that almost all snail shells coil to the right? Or that mating in many land snail species only proceeds after one snail has stabbed the other with a "love dart"? A couple of weeks ago I was recounting the the sad tale of The Society Islands partulids to someone I'd met three minutes earlier, and today I'm going to tell you that story (though, of course you have an advantage over the first recipient of the story, since you don't have to read this crap)

Believe it or not, land snails are one of the characteristic animals of Pacific Islands. Anak krakatau is so young it's still smoldering, and it has a native land snail species and Rapa nui (Easter Island), which is arguably the most isolated island in the Pacfic, had its own land snail fauna back when it had forests. It's not entirely clear how these unlikely colonists get to islands. Darwin was so interested in the question* he, ever the experimentalist, stuck snails to ducks' feet to see if they'd survive an inter-island journey. Birds have been shown to carry snails great distances, but wind blown leaves are probably a more common mode of conveyance. We might not know exactly how snails get to islands, but we know what happens once they establish themselves. The land snails of the Pacific include some of the most outrageous explosions of diversity in the biological world. Chief among these evolutionary radiations were the partulid snails of The Society Islands (the French Polynesian archipelago that includes Tahiti). Partulids are very elegant tree snails that form part of the land snail fauna across most of Polynesia, in the Societies they made up most of the land snail fauna. In total, the tiny islands had 58 species of these snails with each of the main islands have their own endemic forms.

A plate from Crampton's monograph on the partulids of Moorea

The Society Islands' land snails were a marvel all by themselves, but they were also an extraordinary resource for scientists. The first person to seriously take up their study was the American embryologist and evolutionary biologist Henry Crampton. Crampton was working at the turn of the 20th century, a time in which the mechanisms underlying genetics and evolution were very much up for debate, and he hoped Tahitian and Moorean partulids could help set the story straight. Crampton's monogrpahs are famous (at least among people that spend thier lives thinking about snails) for their detail. He collected and measured over two hundred thousand shells, then calculated summary statistics for each species, each site and each measurement. By hand. To eight decimal places.

Those massive tables (there are more than 100 pages of them in the Moorean monograph) might seem like an old-fashioned, descriptive, way to do biology. But in many ways Crampton was ahead of his time. For one, he was a Darwinist when not every evolutionist was. By the end of the 19th century Darwin had convinced the world of the fact evolution had happened, but relatively few naturalist bought his theory of how evolutionary change happened. The anti-Darwinian theories that prospered during the so called "eclipse of Darwinism" placed very little importance on the variation within species. The orthogenesists and the lamarckians thought evolution had a driving force, pushing species towards perfection. In their scheme variation within a species was deviance from the mainstream of evolution and was quckly stamped out by natural selection (which they didn't deny, they just said it couldn't be a creative force). Similarly, saltationists thought large-scale evolutionary changes occurred in a single generation, and the small changes you see in populations were of no consequence in the grand scheme of evolution. Crampton realised that, in a Darwinian world, variation within populations was the raw material of evolution. He was obsessive about measuring his shells because he knew could use the data he was recording to understand where species came from. In particular, we was able to show that isolated populations of the same species varied from each other. That finding that makes sense in light of Darwin's theory, since species arise from populations evolving away from each other; but is harder to fit into progressive theories of evolution, in which you'd expect different populations of the same species to follow the same trajectory.

Crampton's results influenced people like Dobzhanky, Mayr and Huxley who helped to re-establish Darwinism as the principal theory of evolution in the Modern Evolutionary Synthesis. But Crampton also predicted arguably the most important development in evolutionary theory since the modern synthesis. In the middle of the 20th century evolutionary genetics was defined by a single debate. The "classical" school held that populations in the wild would have almost no genetic variation, because for every gene there would be one 'best' version and every member of the population would have two copies of that gene. Arguing against the classical school, the "balance" school argued that, quite often, there would be no single best gene and organisms would do better having two different versions of the same gene**. The ballancers thought natural selection would keep lots of different versions of maybe 10% of a species' genes. Both schools assumed natural selection was such a pervasive force that selection would dictate the way populations were made up, they just disagreed on what would result from it. Here's the funny thing, they were both spectacularly wrong. When scientists started being able to measure he genetic diversity of populations in the 1960s it became clear almost every single gene had multiple different versions. Now, in the post-genomic age there is a database with 30 million examples of one sort of genetic variant amongst humans.

Faced with the overwhelming variation he recorded in partulid shells, Crampton had argued natural selection didn't have a damn thing to do with it. Snails isolated from each other by a mountain weren't adapting to their local habitat, they just varied with respect to traits that had no influence on their survival. The fact two populations were isolated meant each would follow its own path and two populations could drift apart from each other. Faced with the overwhelming genetic variation coming from studies in the 1960s Motoo Kimura proposed the neurtral theory of molecualr evolution. Kimura's explanation was the same as Crampton's, almost all of the variation we see at genetic level has no bearing on the success or failure or organisms so the frequency of different variants drifts around at random. The neutral theory is at the heart of a lot of modern evolutionary genetics, and Crampton had understood the underlying principle 50 years before we knew we needed it!

At the end of his monograph on the partulids of Moorea, Crampton said he'd got as far as his measurements could take him, and it was time for someone to study their genetics. In took a bit longer than Crampton might have hoped, but in the 1960s two leading geneticists took up the study of his snails. James Murray from Virginia and Bryan Clarke from Nottingham spent almost 20 years working in what they called, in more than one paper, the perfect "museum and laboratory" in which to study the origin of species. Their work helped scientists understand, among other things, how ecology can contribute the formation of new species and what happens to species when they hybridise with others from time to time. Then, in 1984, Murray and Clarke had to write the most heart-breaking scientific paper I've ever read. It's written in the careful prose scientists use to talk to each other, but the message it delivered was devestating:

In an attempt to control the numbers of the giant African snail, Achatina fulica, which is an agricultural pest, a carnivorous snail, Euglandina rosea; has been introduced into Moorea. It is spreading across the island at the rate of about 1.2 km per year, eliminating the endemic Partula. One species is aiready extinct in the wild ; and extrapolating the rate of spread of Euglandina , it is expected that all the remaining taxa (possibly excepting P. exigua) will be eliminated by 1986-1987.

The bad guys: Euglandina on the left, Achatina on the right.

Euglandina rosea is better known as the Rosy Wolf Snail. It senses the mucous trails of other snails, tracks them down and eats them. It's not clear if the wolf snail had any effect on the pest species it was introduced to control, but it had huge impact on the partulids. By the time Murray and Clarke wrote their paper, E. rosea had already done for one species and it was too well established to control. All they could do was watch as human stupidity and molluscan hunger slowly (1.2 km per year) destroyed the species they'd been studying for 20 years and Crampton had dedicated 50 years of his life to. The same slow torture played itself out in Tahiti and then the rest of the Society Islands. Where there were 58 named species, there are now 5 alive in the wild. Crampton's hundreds of pages of tables should have been the starting point from which the evolution of the partulids could have been tracked. Murray and Clarke's natural laboratory should still be open and be taking advantage of a new generation of technologies that might be able to reveal the genetic and genomic changes that occur when a new species arises. Extinction is a natural part of life, and the fate of all species eventually, but when it's driven by human short-sightedness and robs us of not just a wonderful product of nature but a window through which we might have understood nature's working it's very hard to write about.

I should end by saying there is just a tiny scrap of good news in this story. The partulids are no longer an iconic species in the study of evolution, but they have become the pandas of invertebrate conservation. Murray and Clarke were able to get 15 of the species of the islands and into zoos and labs across the Northern Hemisphere. Breeding programs have been succesful, and new lab-based studies come out form time to time. The relict populations back in the Societies don't have nearly the range they used to, but it appears they've held on to most of their original genetic variation. Perhaps, one day, Eulglandina can be taken care of and some of the partulids can have their islands back.

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*Darwin had to be interested in dispersal. Before evolution was widely accepted naturalists thought creatures were created for their habitat (the modern creationist notion of a post-flood diaspora explaining the distribution of animals is almost entirely an invention of Seventh Day Adventists, no, really, it is), Darwin's theory did away with special creation but still needed to explain how life came to live everywhere

** A concept similar to "hybrid vigour", in which crosses between relatively unrelated strains/cultivar bring together different genes and do well as a result. You've seen evidence of this phenomenon any time you've eaten yellow and white "honey and pearl" corn. That corn is a hybrid between a white and yellow cultivar and if you count up the kernals you should get close to the 3:1 ratio Mendel preficts for a dihybrid cross.

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Some further reading:

Stephen Jay Gould, who was a snail man himself, wrote and essay on Crampton and the Society Island partulids in which made a humanistic argument for the importance conservation. I resisted the urge to re-read it in researching this piece so anything I stole from him I stole sub-consciously!

• Gould, S.J., 1994. Unenchanted evening in Eight Little Piggies: Reflections in Natural History

Crampton's monograph on the Mooeran partulids (from which the figures above are taken) is available online

• Crampton, H.E., 1932. Studies on the variation, distribution, and evolution of the genusPartula. The species inhabiting Moorea. Carnegie Institution of Washington, 410, 1335.

Finally, the paper in which Clarke and Murray told the world about the demise of their snails:

• Clarke, B., Murray, J. & Johnson, M.S., 1984. The extinction of endemic species by a program of biological control. Pacific Science, 38(2), 97-104.

Labels: environment and ecology, molluscs, Pacific, partulids, sci-blogs, snails, sunday spinelessness

Spineless creatures have been all over the news this week. We've had strange beasts appearing on beaches, wondrous finds from the deeps and (apparently) a new front on the war on cancer.

Let's start here in New Zealand with a surprising find on a Wellington beach.

Photo from stuff article /Emma Best

That's a giant squid (Architeuthis sp.) and at 3m long it's a baby, full grown females can reach 13m. Giant squids are really deep sea predators, so beachings are relatively rare. This one appears to have been in a fight. We don't know too much about the food webs at the bottom of the ocean, sperm whales are the only creatures known to take on adult giant squids but sharks and other whales might very well attack sub adults like this one. As an aside, studying parasites might be the best way to work out who eats who in the deeps. When you dissect a shark or a whale its stomach contents give you a picture of what it's eaten recently, when you look at its parasites you can paint a picture of what it's eaten over its lifetime. However the squid got its injuries, they proved fatal. it washed up dead and was returned to the ocean with the next high tide.

Then there was the announcement of the results of the Census of Marine Life, an inventory of the species living in the earth's oceans. The news of that story was almost universally illustrated with pretty pictures of weird animals photographed from deep sea submersibles. Here's a particularly striking example of the genre, an amphipod with a house which showed up in the Guardian and National Geographic:

Photo from Texas A&M press

Obviously, that mean-looking shrimp-like amphipod is a spineles creature, but it might suprise you that its transluscent home is also an animal. In fact, it's one of your closest relatives in the biological world. It's a salp, a relative of the sea squirts, and it's a member of the chordata - the same phylum as the all the vertebrates. Matt Cobb has all the details in his guest post at why Evolution is True.

Finally, scientists published the first complete genome sequence from a sponge. As has become customary for genome studies, the press releases and the resulting new stories all suggest that these DNA sequences will help us fight cancer. Well, perhaps. Cancer is ultimately about uncontrolled cell proliferation and sponges are the simplest animals to have to find a way of marshalling thousands of cells into a single body. But is that really the only reason we should be exciting about a tool that helps us to understand the origins of multicellularity, or being able to see where the precursors of the development programs that pattern the bodies of more complex animals came from? I'd like to think the average reader is just an excited by a little scientific awe as they are by some ill defined and far off medical benefit. (Before I sound too grumpy I should say that the press release from UC Santa Barbara was really very good, it's just a shame some of those points didn't get picked up in the reporting.)

To those three stories you can add a call for insects to replace red meat, sub zero octopus venom and a BBC tour of invertebrate collections form the Gulf of Mexico. It really was quite the week for invertebrate news.

Labels: carnivorous sponge, crustacean, genomics, molluscs, porifera, salp, sci-blogs, squid, sunday spinelessness

Stuff.co.nz has a very cool video that was shot from an underwater video camera as it was grabbed by a lurking octopus:

It's amazing to see the octopus's suckered arm shoot out from a reef and grab at the diver, and to see the octopus enjoying a ride on the blunt side of the diver's spear a little later. But for me, as someone who spends his days studying landsnails, it really set me to thinking about the remarkable diversity of form in the phylum Mollusca.

This series has focused on the other great animal phylum, the Arthropoda, and with good reason. Most animals are arthropods. But that group has achieved its dominance by finding a good basic body plan, an exoskeleton and set of jointed limbs, and making a million variations on the theme. By comparison, evolution has molded the soft mollusc body into thousands of forms. Cephalopods like the Octopus maorum featured above are perhaps the most amazing result. They are intelligent, free swimming predators with three hearts and amazing powers of disguise. Which is why I still find it amazing that cephalopods likely evolved from snail-like ancestors.

I really shouldn't be so surprised. There are plenty of snails around today that cast off that group's reputation for a sedentary lifestyle to swim the seas as predators. One example that might be familiar to readers is the violet snail, Janthina sp., whose pretty purple shells are occasionally washed up on New Zealand beaches. The violet snail can't actively swim as an adult, instead it builds a raft of air bubbles trapped in chitin and floats on the surface where it eats swimming jellyfish.

Violet snails shell from Flickr user quadprop and washed up animal with raft from wikipedian Rez242

The violet snails are pretty cool, but they aren't a massive divergence from the gastropod body plan. The snails that used to be known as pteropods are much more radically adapted to swimming. These snails are now recognised as two distinct groups, the "sea butterflies" (Thecosomata) which have retained their shells and the "sea angels" (Gymnosomata) which have lost them. As you might have guessed form their names, both of these groups have developed "wings" (derived from the foot muscle) they use to flap their way through the water column.

Sea angel from USGS and sea butterfly from NOAA

There is one group of swimming snails that are even more stunning than the sea angels and the sea butterflies. The nudibranchs (commonly referred to as "sea slugs") are pretty weird bunch of gastropods to start with. They come in a bewildering range of colours and some of them are even solar powered and there are a couple of species that have become adapted to swimming, or at least floating, on the ocean surface. Two species from the family Glaucidae, Glaucus atlanticus and Glaucilla marginata have developed swim bladders and long limb like protuberances that make the look, to me at least, a little like fish:

Glaucus atlanticus licensed CC 2.0 by flickr user tarotastic

Labels: cephalopods, environment and ecology, molluscs, octopus, photos, sci-blogs, snails, sunday spinelessness