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

It's the first week of semester two down here at Otago, which means I will helping with undergraduate labs for the first time this year. I suspect most students end up not liking me all that much, because I find my self teaching in the parts of the genetics program that undergrads like the least. Population genetics, as the name suggests, is the study of the way genes behave in populations, and in many ways its the base from which our understanding of evolution is built. So it's important, but it's also pretty mathsy.


It seems quite a few students have planned their high school and unversity careers in the hope that studying biology meant that could leave maths behind. So, when they are confronted with "p² + 2pq + q² = 1" and asked to do something with it, they are unhappy.

That particular formula is for something called the Hardy-Weinberg equilibrium and a significant proportion of students roll their eyes and slump their shoulders when you tell them they are going to need to use it for a problem. They think its arbitrary and irrelevant to anything the least bit important, and what's more it looks a little like it's already solved. So, I'm always looking for ways to convince people that Hardy-Weingberg isn't just simple, but actually intuitive and important. So here's my attempt to explain why knowing a little population genetics is helpful.

You may remember last year  a Danish sperm bank had started turning away would-be donors with red hair, since there is little demand for sperm that might contribute to the creation of a readhead. It turns out, if you know a little bit about population genetics you can see that policy will have little effect on the number of red heads the sperm bank helps to bring into the world.

Hair colour is partially controlled by a gene called MC1R. There are different versions of MC1R floating around in human populations, and one of them has a mutation that stops melanin (a dark pigment)  passing into hairs as they grow. Geneticists call different versions of a gene "alleles", so we'll call this flavour of MC1R the "red hair allele" and give it the symbol r.As I'm sure you know, you have two copies of most of your genes, one inherited from your mother and the other from your father. Red hair is a recessive trait, which means in order to have red hair both of your copies of MC1R need to be the r allele: if you have one or two copies of the "normal" MC1R allele (which we'll call "R") you have some pigment passing into your hair and it will be another colour. We call the total genetic make-up of a person their "genotype", and their physical characteristics their "phenotype", so here's a table showing the genotypes and the phenotypes we're talking about in this post:

Genotype	r/r	r/R	R/R
Phenotype (hair colour)	Red Hair	Not Red Hair	Not Red Hair

I know there are a lot of technical terms there (Carl Zimmer will not be happy...), but we do need to be precise when we talk about genetics because, strange as it may seem, there isn't a single definition of the word gene. Once you've got your head around the terms, it's all pretty straight forward: you need two copies of the r allele to have red hair. Think what this means for the Danish sperm bank though. Turning away red headed sperm donors doesn't turn away red headed sperm since there will still be "carriers" with only one copy of r (and, thus, non-red hair) donating sperm and half of those sperm will be "red headed sperm".

How big a problem is this likely to be? First we need to work how common the r allele is, and we can use the frequency of redheads to find that. By long tradition, the frequency of a recessive allele is denoted by "q", so, in a population where one quarter of the alleles are r we'd say q = 0.25. We know that in order to have red hair you need both your copies of the MC1R gene to be the r allele and that you inherit each allele separately. When probabilities are independant we can mutiply them, so the chace someone in this population is a redhead is q x q = q²  = 0.06 .Following the same logic, the frequency of the R/R genotype must be the frequency of R squared (by convention, the frequency of a dominant allele is called "p", so that's p²).

Knowing this relationship, we can work backwards and find the frequency of r if we know the proportion of redheads in a population. In most of Northern Europe, about 4% (0.04) of the populaiton are redheads so  q² = 0.04 and q = √0.04 = 0.2. As you can see, red hair genes can be a lot more common than redheads:

To understand how the sperm bank's policy will we work, we need to know about those 'carriers' with the mixed genotypes (called "heterozygotes" by genetics geeks). It doesn't matter which order your genes come in, so the probability of being a carrier in the population above will be the chance of getting an R then and r (p x q) plus the chance of getting and r followed by an R (q x p). You can simplify that to 2pq. You might recognize that term, because with it we've rediscovered the one in the first paragraph "p² + 2pq + q² = 1". The Hardy-Weinberg equation is just away of moving from allele frequencies to genotype frequencies (or the other way around) and it's based on some very simple observations about the way populations work. We saw that in a population with  4% redheads you get q = 0.2, so p=0.8 and 2pq = 2 x 0.2 x 0.8 = 0.32. Almost a third of the population are carriers, and that's eight times the number of redheads! While the frequency of the red hair allele is low, only a small proportion of the red haired alleles in a population will actually in red haired people:
That's why the sperm banks policy, while prefectly sensible if there really is no demand for sperm from redheads, will do little prevent the creation of red-headed babies.  In the typical case, where 4% of the population are redheads the probability that a donated sperm carries the r allele only moves from 20% to 17% when you exclude red haired donors

It's easy to calculate how the policy would work in populations with more or less redheads:

So, that annoying equation we make the undergrads learn actually tells us something about the world. Obviously, the example I've talked about here is a pretty silly one, but the basic ideas we've discovered above can help us understand some important ideas. Like why genes that cause debilitating diseases aren't completely removed by natural selection, and why inbreeding is a bad idea.

A lot of rare diseases are caused by recessive alleles. They remain rare for the obvious reason that people with such diseases are unlikely to pass on their genes. But they remain present in populations because, as we've found, once recessive alleles get rare the overwhelming majority of them are found in carriers. In this way, rare recessive alleles are seldom exposed to selection so they stick around for a long time.*

Because disease causing genes stick around in populations, there is a pretty good chance that you carry a few alleles that would cause a debilitating disease in someone who had two copies of them. The same applies to anyone you might be hoping to have children with. Thankfully, its very unlikely that your prospective mate with have disease-causing alleles for the same genes that you do. That is, as long as you look beyond the family tree when you look for a mate. If you have a child with someone who is closely related to you, you will have each inherited some of your genes from the same source, which increases the chances you share disease alleles.

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*In fact, they often reach a point called "mutation-selection balance" in which the frequency of the allele remains static because new mutations re-create the allele as quickly as selection removes it. JBS Haldane was the first person to notice this, and he used his theory to create a very accurate estimate of the human mutation rate well before we knew what genes were made of!

Labels: genetics, population genetics, sci-blogs, science, teaching

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