Posts Tagged 'sexual selection'

Species recognition in dinosaurs? Not so much

Those with an interest in dinosaur cranial crests and exaggerated structures (which should really be everyone since they turn up in pretty much every major lineage one way or the other) will probably be aware of the exchanges going on in the literature over these features. Although myself and colleagues have been advocating that sexual selection (and or socio-sexual signaling: the two can be hard to separate) is a likely strong candidate as the prime driver for many of these features, others have been advocating that this is not the case and instead the answer lies in species recognition. The latest to delve into this area is a paper I’ve done with Darren Naish and is the first time we’ve addressed this issue directly. While we have written or contributed to a number of efforts looking at support for sexual selection in dinosaurs, this is the first time we have tackled the other side of the problem.

The paper originally started as a long section that was included in our paper on mutual sexual selection with Innes Cuthill, but as we were later forced to cut down the length of the submission, this was a section that was relatively easy to prune as tangential to the main issue. However, we felt it needed saying and with new data coming out and the discussion ramping up, we revived and revised the work and it is now out. (Well, it has been in press and available for a while but is now properly out).

This is an important area for discussion – after all, the horns, crests, frills, plates, bosses and the rest (not least feathers) are key features and adaptations in various dinosaur lineages and trying to work out how they might have been used and what this means for evolutionary drivers and patterns is going to be a major issue. It’s hard to really understand stegosaurs or ceratopsians say if you can’t say that much with confidence about their ‘bonus’ features. While obviously each clade, or even each genus / species probably needs to be taken on a case-by-case basis when it comes to detailed analyses, some gross patterns can be seen or at least discussed. In the case of species recognition, is it even an actual ‘thing’ when it comes to exaggerated structures, and if it is, how is it supposed to work. The hypothesis has enjoyed some support in the literature for some unusual dinosaur features so it’s well worth examining.

Species recognition (in the context of exaggerated structures) for those who don’t know, is the idea that individuals of a species use these features to help them recognise cospecifics with to ensure they mate with the right species, or to maintain herd coherence. In short, carry round a key feature and you should be able to make it easier to stay in touch with the right animals and avoid the wrong ones. Various lines have been put forward to support this idea (in general and specifically towards dinosaurs) but we feel that none of them actually stack up and some have some serious problems.

First off is a pretty big issue – to our knowledge there is no evidence of any living species using some form of crest or exaggerated structure for species recognition. Individuals of species do recognise each other (not a big shock) but actually things like antlers or casques don’t seem to form part of the pattern that conspsecifics recognise. This may not be a big shock, after all, you can recognise a species by the overall appearance (size, shape, colour), their smell or specific sounds they make, behaviour, and other features. On top of this, some species are very varied in appearance for the big features (antlers of deer look very different as they grow, and are different between males and females and between juveniles and adults etc.) so relying on one feature is a bad idea at best, and a plastic one an especially bad call.

Plus of course, you often get closely related taxa that are sympatric. Is some big set of horns going to help you correctly identify conspecifics if there are half a dozen similarly-looking species also in the area? Look at things like African antelope and gazelle, or more extreme examples like tyrant flycatchers. We have trouble telling them apart sometimes based on their morphology, yet they seem to have no trouble. If this is so critical to dinosaurs, why to the iguanodonts seem relatively free of crests, but the hadrosaurs go nuts with them? And why are they all so similar in general form between species when they are supposed to help separate them out? Surely they should be divergent, not all similar in appearance. And why do we see things like Wuerhosaurus or Spinosaurus running around with all this weight to make sure they don’t mate with the wrong species when there are no other members of their clade to get confused with?

In some cases we see both issues coming together. If we look at the various small protoceratopsians of China / Mongolia, we see disagreement between researchers as to how many species (or genera) there may be. What is notable however, is that the characters being used to separate them out don’t typically involve the frill or bosses of the skull, and where they do, may be things that are not externally visible (e.g. the width of the media bar in the frill). In short – if there are multiple species here, the frills are apparently similar enough that we can’t separate them and so are unlikely to be part of the identity concept of the animals. If however, there is only one species present, then we are back to the paradox of a large frill being carried around but with no other species that could confound any signals.

On top of that, is it really worth it? After all, while you do want to stay in touch and make sure you mate with the right species, bolting on a good few kilos of bone to your head, and then the extra muscle to support it, and then lugging that around for your entire life is a lot of effort. When you can probably already identify conspecifics by their colour, patterns, scent and calls (of simply because nothing else like them at all is on the same continent) surely these would experience strong negative selective pressures if they didn’t have any other support.

Furthermore, how would such features ever evolve? If the populations / species were allopatric then we return to the situation of them not having another group to get confused with and crests are unnecessary for recognition. If they were sympatric though, how would this work? Pretty much the definition of a natural biological population is one that is breeding within itself, but here we’d have to have a population diverging because some don’t recognise each other as conspecifics even though we would expect, pretty much by definition, there not to be too much difference in structure shape between them (e.g. a tiny crest vs no crest). Now some animals might prefer each other, but that’s mate choice, not recognition, and there would have to be enough individuals for this to work – one mutant with a crest when no one else has one is not going to start forming a new species, and if there were a bunch of the with the new crest they’d also have to identify each other as different and avoid mating or hanging around with the others. So how would a large feature that’s for correct recognition allow a population to split in this way? To us at least it appears most unlikely to occur at all, let alone repeatedly.

In addition to this, there is rampant hybridization of closely related species in the natural world (and indeed in captivity). Even extravagantly ornamented species like pheasants with numerous adornments and bright colours and patterns hybridise regularly – clearly no matter how extreme the cue, at least some animals regularly have problems with them or are indiscriminate, but either way they are not that effective.

While some data like the apparent rapid growth of structures late in ontogeny has been used to support the idea that they are characteristics involved in socio-sexual signaling, it’s also a problem for the herd coherency part of the model. After all, lots of juvenile dinosaurs are known from aggregations suggesting they spent a lot of time together, even when the adults did not appear to. If these features were key, we’d expect juveniles to have them, and adult perhaps to shun them when they were no longer needed, but instead the opposite is true. In general the herd coherency argument is a bit odd anyway, again you have lots of ways of identifying and keeping in touch with conspecifics and some are clearly better than visual aids. Scent can have a temporal component, and vocalizations can be interactive beyond line of sight (especially useful in forests, or when things are behind you, or you are foraging and looking down etc.). No matter how big they are, visual structures are not always going to be that useful, even if they are unique.

In the increasingly infamous issue of Torosaurus and Triceratops, if these animals are truly conspecific then for a start we are back to the issue of ‘lone’ taxa (I don’t think Leptoceratops is going to be much of an issue here) and the pointlessness of crests where none are needed. On the other hand, this is also potentially a problem for the mate recognition idea. We know that at least some dinosaurs were sexually mature before they were osteologically mature and this could be the case for these animals too. If so, then the alleged transformation between one morph and the other would create confusion – both the Triceratops morph and the Torosaurus morph (or indeed anything in between) would be viable mates.

In short, we really have no clear evidence for species recognition in any living species, and that alone should make it unlikely to have been a key player across dinosaurs for the whole Mesozoic. Such structures would be costly, and yet not necessarily do the job it is supposed to with other signals being cheaper and just as effective, or more effective in many circumstances. It’s not clear why it should be so important for some clades and not other similar forms (iguanodotids vs hadrosaurs for example) and is clearly either redundant for some taxa, or would not actually reduce confusion. Nor is it clear quite how this would evolve in the first place, or why it would be sustained, and hybridization suggests that crests alone would not even prevent incorrect matings. Put this all together and we feel that there really is no good support for the idea of crests and other structures being primarily used in species recognition. They did of course likely have an effect – it would be odd if Stegosaurus or Corythosaurus didn’t use their respective features as part of how they identified one another. But that does not make them the prime, or only, driving force of all these different features in all these different lineages.

There was a fashion in dinosaur palaeo to write off any odd structure as simply sexual selection and leave it there. This was rightly railed against, but what was often criticised was the fact that sexual selection seemed undiagnosable in the fossil record and so the problem was that it was untestable rather than the fact that such throwaway remarks devoid of context or explanation do little for the subject. Now we are in the odd position where rarely you see very similar comments (in terms of their style) about species recognition popping up in the literature about exaggerated structures despite the lack of support for it, and the now (well, we think), strong cases made for sexual selection, or at least it’s assessment. Although previously the case for sexual selection was pretty weak, it is at least an extremely common phenomenon in living taxa and with obvious powerful effects on anatomy and behaviour. Species recognition has not yet even been shown (in relation to exaggerated structures) in any living clade, and while offhand one-line explanations are not the way to go, it seems odd that one has been replaced with the other.

 

Hone, D.W.E., & Naish, D. 2013. The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non-avialan dinosaurs. Journal of Zoology, 290: 172-180.

Sexual selection in the fossil record

Regular readers will know that for the last few years I’ve been slowly building a research profile concentrating on the behaviour and ecology of dinosaurs and pterosaurs. While the various papers on feeding behaviour, stomach contents, predation and niche partitioning in theropods has been the more high profile, I think the work on sexual selection is arguably more important as it potentially has profound implications for how we interpret all manner of fossils and how they may (or may not) relate to one another. After all, there’s a major ecological and taxonomic difference between identifying two species of a clade, and one species that exhibits major sexual dimorphism.

My colleagues and I have already looked at the idea that sauropod necks were driven by sexual selection, and after much strife, finally got a paper published discussing mutual sexual selection and the implications that has for diagnosing taxa in the fossil record and what it might mean for parental care and other aspects of behaviour. There’s more to come in these areas as I have further work planned and am involved in some other areas linked to this, so the area is growing rapidly and, I hope, ripe for a general revisit in the literature. However, while these papers have in large part being about drawing out some false assumptions in the literature and providing new hypotheses about sexual selection that could be looked at in the fossil record, they were a bit short on how this could be done, and were if anything, narrow in focus (not that Ornithodira is a small group, but its got nothing on Animalia).

So then to a paper in TREE that came out yesterday online. Led by entomologist Rob Knell, it also includes  Darren Naish and myself and attempts to provide a review of the entire question of sexual selection in the fossil record. We look at ways in which this could be diagnosed, some false dichotomies and assumptions that have been put forwards in the past, try to identify some key features that may help diagnose sexual selection and look at some of the more convincing cases for this that have been put together to date. Clearly there’s a limit to what we can get into under 10 pages for what is supposed to be a review, but I think there’s some nice synthesis in there and a bit more “we can try doing this”-type stuff, that just covering what has been said before. Anyway it’s out and available (though behind a paywall, sorry) so go take a look.

Knell, R., Naish, D., Tompkins, J.L. & Hone, D.W.E. Sexual selection in prehistoric animals: detection and implications. Trends in Ecology and Evolution, in press.

Neck multifunctionality

Following on from my post on sauropod neck lengths (and indeed that of the SV-POW! boys and Tet Zoo too), at the end I made the inevitable comment that necks (and indeed other structures) can be multifunctional. A long neck can be an indicator of sexual selection at the same time as providing increased reach for food. As noted before, analogy plays an important role in working out (or at least hypothesising) palaeo behaviour, so are there any animals out there that seem to do this. Indeed there are, so step forward, the Galapagos giant tortoise.

Tortoise neck use - above, reaching for food, below, challenging rivals. By Darren Naish, from Taylor et al., 2011.

Research shows that those animals with longer necks do gain a distinct advantage in reaching higher placed food. The neck provides a genuine basis for natural selection based on neck length. However, it has also been shown that when tortoises stand off in dominance battles, the individual with the longer neck tends to win. So, necks would also appear to be under some measure of sexual selection / dominance as well.

So what about the giraffes? They have long necks and the males fight with their heads, so what’s going on there. Well males do have longer necks than females, but there’s not really anything to say that longer necked males do better. Bigger males do better (no surprise) but not through a longer neck per se. Plus of course the males are actively fighting with their heads. (And tortoises can be quite vicious if you’ve ever seen them fight, then bite and butt with their shells).

Posturing only gets you so far in nature. Sure, there are cheats out there (false cleaner fish, milk snakes, female mimic salmon etc.), but they can only prosper as long as they are in the minority. This is because sooner or later someone is going to square up to you in one way or another and find out if you really can back the bark with bite. If you can’t, you’re going to lose. And if say most of the population were lying, once a dominant animal (or predator etc.) finds out, then that is going to take over damned fast. So lying only works when there are few liars, and most things are honest. In other words, if they are advertising that they can win a fight, it’s because they can and will.

What does this mean for sauropods? Well is has been suggested in the past that sauropods might fight one another, with their necks. Now if this was going to happen you’d expect to see some evidence of this in sauropods. Like the especially tough and thick skulls of male giraffe, or the prow-shaped rams of some tortoises, or robust necks and heads in male sauropods and you’d see injuries from some serious sauropod neck-on-neck action. Only of course there aren’t any.

Instead sauropod skulls are incredibly weak and fall apart if you look at them funny, let alone ram them into something else at speed. And while the neck as a single unit might be quite tough, it has those lovely wafer-like lamina and those oh-so-thin cervical ribs. If they were fighting we’d see breaks, pathologies, healed bones and the rest. And you can’t cheat by just having a big neck and expecting the others to back down, you have to back that up or someone will realise it’s all talk.

I’m sure sauropods did fight on occasion, sooner or later animals of pretty much any species will come into competition and of course it is members of the same species that tend towards the fiercest competition. There will come times when accessing that water hole, or harem, or territory is critical and combat becomes inevitable. But was it with the neck? No. The neck might have been a *symbol* of the power of the individual even if it wasn’t used (pheasants and cockerels show off their colours to demonstrate their fitness, but they fight with their spurs).

Sauropod necks – what were they for?

Long time readers might well recall that for a few years I was a volunteer at London Zoo where I worked on what was the Cotton Terraces (now sadly rebranded as the “Africa Zone”) with various hoofstock and ungulates. It was, generally, an absolute joy and I learned a great deal about animal management and animals in general. Among my occasional charges were the giraffe and I have a long and abiding affection for them from my time there. This has in part manifested itself by my keeping up with giraffe research in the literature and when an opportunity arose to write about giraffes and dinosaurs, well, I could hardly resist.

So what do giraffes and dinosaurs have in common? Well not much to be honest, but perhaps inevitably the long necks of giraffe have been brought up a number of times in the past as analogues for the long necks of sauropods. It’s not unreasonable really as good analogues are important for inferring the functions and behaviours of extinct taxa and there are very few terrestrial long-necked herbivores out there, and even less that look even vaguely like sauropods. So the high reach of giraffes has been used to suggest sauropods reached high into the trees to feed.

Male giraffe sparring at Beijing zoo.

Set against this background has more recently been the controversy over sauropod neck postures and quite how much (or even if) they could raise their heads and necks. But it gets more complicated still (yay!). Back in the mid 1990s it was suggested that actually giraffe necks hadn’t evolved for feeding high in trees, but instead were sexually selected structures. Perhaps not surprisingly, the same hypothesis was then extended onto sauropods too! If they couldn’t raise their necks up, and the only obvious living example (in giraffes) was only an opportunistically high browser, then maybe sauropod necks were the result of sexual selection too? (Mike Taylor has a preview of these ideas here).

In a new paper out today myself and the SV-POW! boys (Messers [or rather, Doctors] Taylor, Wedel and Naish) take on this idea. The detail is of course in the paper but there are several interlinked threads to this paper that affect different lines of sauropod research and ecology. Perhaps most interestingly for me is the work on giraffes though, since actually the idea of sexual selection in these beautiful artiodactyls is far weaker than originally proposed and actually high browsing does seem to be the primary function of the neck.

Sauropod neck lengths. From Taylor et al. in press

Thus the original analogy can be restored and the analogy for sauropods having sexually selected necks rather falls by the wayside. Couple that with the existing work by my colleagues on the potential vertical reach of sauropods and another barrier falls by the wayside. Even if sauropods couldn’t reach high into trees though, that’s not necessarily a barrier to their long necks being adapted for feeding efficiency. Anyone who as seen a grazing goose will recognise their feeding pattern with the body of the bird only plodding along slowly but the long neck sweeping from side to side so that for every step forward (a big deal of effort for a 50 ton sauropod say) a fair area of new grass can be covered by the head. In short, a long neck can be efficient whether you are reaching up or not.

All together (and there is obviously more to this than I’m covering briefly here) we find no convincing evidence of sexual selection going on in sauropod necks and are satisfied that the long necks would have provided a significant horizontal and vertical reach and thus did afford a significant part of their feeding ecology. There’s no good evidence (at the moment, at least) that any sauropod necks were under sexual selection or indeed, that those of giraffes were. That doesn’t mean it wasn’t happening, but there is nothing to suggest it was.

This must, of course be hedged with a couple of most important and basic caveats to these kinds of papers. Critically is that of multifunctionality – that structures can have more than one function, so there could be some cryptic sexual selection ongoing alongside feeding advantages. Though as it happens in this case, it’s this very issue that helps us deal with some of the evidence for sexual selection. Secondly of course this paper is a review – we talk about general patterns and trends and evidence. We do not look at all sauropods in great detail and while we have great confidence in the findings of our paper we are talking about generalities – there is of course a reasonable chance that something was an exception.

Taylor, M.T., Hone, D.W.E., Wedel, M.J. & Naish, D. The long necks of sauropods did not evolve primarily through sexual selection. Journal of Zoology, in press.


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