Monday, 30 September 2013

Theropoda Sympatrico Pt.2

In the previous post we had started to look at the importance of theropods within the ecosystem and I suggested that the possible extinction of large carnivorous dinosaurs may have been the catalyst for selective extinction for some animals and also the consequential acceleration and proliferation of others. In other words, large theropod extinction may have had direct implications for faunal turnover within ecosystems.
Although I have essentially been looking at large theropods it is worth remembering that the diversity of theropods in most ecosystems was generally very complex. Regardless of size and/or niche even the disappearance of a small theropod may have caused limited environmental problems (more of this later). In simple terms, and if we use a large predator as an example, if a particular carnivore was in decline or on the road to extinction, for whatever reason, then it is likely that the herbivorous animals that they preyed upon would increase in numbers. This, in turn, would have added pressure to the plants, trees and other flora since, as the herbivores increased in number, and then there would be less chance of a successful germination by those plants affected.
When plants are affected then, not only non-avian dinosaurs, but  everything becomes affected including, in Mesozoic times, insects, pterosaurs, mammals and birds and these effects would have changed their distribution, relative abundance  and, most importantly of all, their intraspecific behaviour. The loss of a single large theropod therefore would have had enormous effects on an ecosystem.
Another interesting thought is whether theropods displayed variable feeding tendencies like we see in carnivores today. It is certain that the majority of carnivorous theropod dinosaurs could at least be termed as hypercarnivorous – that is their diet would have consisted of at least 70% meat and it is fair to say that the majority of stereotypical theropod teeth are blade like and serrated for exactly this kind of diet.
However, it is difficult to comprehend that theropods, by their very nature, would have been able to utilise the other 30% worth of plant matter and fungi to be genuinely referred to as hypercarnivores and the vast majority of theropods should, indeed, be known as obligate carnivores since it is certain that they had neither the dentition or, even more importantly, the digestive capability to assimilate such matter.
But theropods were still exceptionally diverse and we have discovered many weird and wonderful animals now that were indeed cosmopolitan in their tastes. Ornithomimids appear to be clearly omnivorous, therizinosaurs were certainly herbivorous and the peculiar oviraptorids appear to be capable of consuming a great variety of foodstuffs. The fascinating Alvarezsaurids seem ideally suited to be insectivores whilst, at the other end of the scale, we have the great baryonychines and spinosaurs that appear to be both great fish and flesh eaters.
Regardless, the largest carnivorous theropod in any given ecosystem would still have been the most important ecological factor in determining the health and prosperity of the system. They were at the very top of the trophic cascade and would exert a level of influence on those species at the next level down – their prey. These animals too exert influence on those species below them and so on until we reach the very bottom of the trophic ladder. This is known as top-down regulation.
Large theropod dinosaurs, although relatively few in number, would have exerted considerable pressure on species below them. Fear would have been a strong driving force and their presence alone would have indirect effects on prey animals and, of course, predation would have very direct effects indeed. These effects would have had a ripple action throughout an entire community of animals so that the entire ecosystem was affected.   
And yet, as we have already observed, palaeocommunities were also inhabited by smaller theropods and these were much more diverse and common than their larger cousins. Although not at the apex of the trophic ladder they would have filled many varied predatory niches and it was this adaptability that enabled them to diversify and, as a consequence, their numbers would have been far greater than their giant cousins.
However it does seem likely that a small theropod’s disappearance would have only had a moderate effect on the ecosystem. Many of the smaller carnivores’ predatory niches would have almost certainly overlapped thus the effect on the environment would have been reduced as opposed to those effects caused by the loss of an apex carnivore.
Large apex theropods would have needed large territories to fulfil their nutritional requirements especially as it is now generally accepted that they were active fast growing animals - if not fully endothermic. In an extant mammalian ecosystem, the rule of ecological efficiency suggests that there needs to be ten times the amount of prey animals to support a modest carnivorous population. Even if dinosaurs were not fully endothermic we do understand their physiology enough to realise that this general rule of thumb would probably apply to large theropods but perhaps on a slightly lesser scale.
Allowing for sampling bias and other taphonomic distortions the fossil record for large theropods in various ecosystems tend to support this equation and that they are generally found in low population densities and would therefore have required large territories. So what were the implications for ecosystems if a top carnivore declined in numbers on the road to extinction?

Because of their limited numbers, any decline in large sized theropod populations would have been keenly felt – sooner rather than later. Their numbers would have become increasingly scattered thereby decreasing the likelihood of intraspecific encounters with others of their own species which, in turn, greatly decreased their chances of a successful mating. This leads to an accelerated rate of decline as the animals that remain may inbreed leading to further genetic complications and disease. This whole process would likely have been extremely rapid leading only to the one conclusion – extinction.  
As the large theropod population declined it is certain that the population of their main prey animals would boom and the effects on the flora and fauna would be dramatic. Plants and trees would be seriously compromised and degraded due to increased and unregulated grazing. The unchecked herbivorous dinosaurs, without fear of predation, would have also probably increased their own ranges which, in turn, would have enabled them to perhaps exploit other food sources leading to yet more environmental consequences for other species. So many things would have changed including those that would even seem to be relatively mundane such as longer feeding spells at different feeding times and, of course, the sizes of the groups and herds as they got bigger.
Would the disappearance of the giant apex theropods affect their smaller brethren?   The answer to this is an uncompromising yes. Not only did the giants keep the herbivorous population in check but it is very likely that their presence kept the numbers of their smaller carnivorous cousins under control. Released of these shackles then small carnivorous theropods may have also gone through something of a population boom which, naturally enough, would have increased predation of other taxa thus increasing further the pressure on the ecosystem. These smaller prey animals are often essential parts of ecological stability that filter right through to the bottom of the trophic ladder.
And just as the herbivorous dinosaurs behaviour would have changed so the behaviour of the smaller theropods would have changed too. It is fairly certain that the small theropod population would have been fearful in the presence of their larger cousins and very likely kept their distance. Freed of this fear factor small theropods would also probably have changed their behavioural patterns and this would also have added yet more pressure to an already distressed ecosystem.
So we can see how the sudden loss of a group of large apex theropods could possibly be the spark that leads to the degradation of an ecosystem which affects every kind of plant and animal on the trophic ladder. Understanding the trophic ladder is key to understanding ancient ecosystems and it is this realisation that everything is linked together from top to bottom that helps us see life for what it was – and is.
So perhaps you may be thinking that there may be something to all this - and I believe there is. Certainly there is every reason to believe that localised extinction events and regular faunal turnover were almost certainly driven when large theropod carnivores came under pressure and started to disappear. There are a number of factors that may have led to large theropod extinction and these include, for example, disease and environmental stress.    
And yet, as I have alluded to before, dinosaurian ecosystems represent something of a conundrum in as much that a quite a few of them now seem almost impossible to have functioned efficiently – and yet they obviously did. In the final part of this mini-series we will try and work out some of the permutations that led to the success of these ecosystems, try to understand the apparent high faunal turnover levels and why sympatry in large theropods is still kind of weird.

Tuesday, 10 September 2013

Theropoda Sympatrico Pt.1

When the Dinosaur Renaissance gathered momentum during the Seventies, its biggest proponent and champion was one Robert T. Bakker – a palaeontologist who needs no introduction. Bakker was instrumental in displaying dinosaurs in a new light. Gone were the old depictions of cold blooded, slow moving, dim witted reptiles of the primeval swamps and in their place came this new dynamic clade of animals – animals that were fast, agile, rapidly evolving and, above all, warm blooded. Indeed, things would never be the same again and we all got swept away in this new glorification of the dinosaurian race.
Chief amongst Bakker’s evidence for proclaiming the existence of endothermian dinosaurs were the use of predator/prey ratios (PPR’s).  Simply put, in any specific ecosystem, the amount of carnivores that can be supported by any given population of herbivores can be measured as a ratio. Because cold blooded carnivores generate their body heat from the sun they can eat much less frequently and, as a result, larger numbers of carnivores can exist contemporarily. But because warm blooded carnivores need to generate their heat internally they must eat much more frequently so the amount of carnivores that can be supported in the ecosystem is much much lower.
Bakker argued at the time that by comparing, not only extant mammalian communities, but also Permian reptilian faunas with dinosaurian faunas, that because PPR’s in dinosaur communities were so low, and those in the Permian were so high, that dinosaurs must have been warm blooded. At the time this was seemingly ground breaking research and there were many believers but this was clearly wrong and we now know that there are many variables that rendered the hypothesis redundant.  Not only is it flawed to compare extant mammalian communities with extinct ones but sampling in the different formations is highly variable and problematic and thus it is difficult to formulate any ratio with any confidence.
Dinosaurs may very well have been warm blooded and there is a weight of evidence now that supports that they were, at the very least, more than ectothermic if not fully endothermic. However, there was a curious by product of the predator/prey hypothesis and that was that it was commonly accepted that you would generally expect to find, in any sampled dinosaurian fauna, only the one large predator in residence. This persisted for some years probably perpetuated by the fact that Tyrannosaurus rex, as one of the best known and well researched theropods, is, indeed, the only large carnivore in the Late Maastrichtian of North America.
It was strange too that the revelation that Daspletosaurus and Albertosaurus were contemporary large tyrannosaurs in both the Dinosaur Park and Two Medicine Formations was simply accepted as unusual. The most popular explanation was that this was evidence of niche partitioning with Albertosaurus primarily feeding on hadrosaurs while the more heavily built Daspletosaurus preyed upon ceratopsids (Russell 1970).
But eventually, as more and more formations became better sampled, it became apparent that, actually, sympatric large theropods are the norm with, perhaps, the most extreme example being the population of the Late Cretaceous Kem Kem Formation in Morocco which has produced Carcharodontosaurus, Deltadromeus, and Spinosaurus amongst others.
These co-habiting theropod populations bring their own unique problems to the table and continue to fascinate palaeontologist’s in trying to work out how these large carnivores managed to share out the various resources available. These issues are amplified when we look at the amazing populations of Laramidia which displays several much smaller pocket environments where it appears virtually impossible for there to be enough resources to go around for all dinosaurs – let alone the carnivores.
Dinosaurs are freaks then and this is precisely why you cannot simply compare extant faunal communities with dinosaurian faunas. It appears almost impossible in some cases for dinosaurs to be able to coexist at all – but they obviously did and what a magnificent race of creatures they are. So how can we explain how large theropods co-existed successfully in what would appear to be extraordinary circumstances? There are many things to consider but perhaps we should look at the nuts and bolts of the carnivorous dinosaur – what is a carnivorous dinosaur, what were their functions and, most importantly of all, how important were they to the ecosystem?
Theropod dinosaurs are spectacular – from the smallest raptorial forms right through to the largest tyrannosaurs. And it is safe to assume that, when compared to their herbivorous contemporaries, that they were relatively more “intelligent”. I have put the word intelligent in quotation marks because it is important to make the distinction between intelligence, as displayed by extant mammals, and that hypothesised for dinosaurs for it is unlikely that any dinosaur was capable of intelligent reasoning in comparison, for example, to a dolphin.
But it is obvious that they would have been smarter than their prey because their predatory nature demanded it and, I believe, it is safe to assume that this would also entail a degree of complexity. How complex, again, remains a matter of conjecture but it is a possibility (some of us believe likely) that there was a degree of social complexity within Theropoda whether they were solitary hunters or hunted in groups. This is highly suggestive that there would have been a significant variety in their behavioural patterns which would have included their hunting techniques, their courtship and mating rituals and even how they may have raised their young. We only have to look at birds to get, perhaps, a hint of what may have been a variety of different behaviours displayed by their dinosaurian ancestors.
Large theropods must truly have been a sight to behold. The largest of them would certainly have nothing to fear in their ecosystem aside, perhaps, from another of their species. Other dinosaurs may have been as equally impressive and many were indeed far larger than even the biggest theropod but for me they have no equal. And despite the fact that we know so much about them it is still the vast amount of information that we don’t know about them which is so fascinating. But the one thing we can be sure of, as with any ecosystem, theropods, and large theropods in particular were at the top of the food chain - the very peak of the trophic pyramid.
Carnivorous dinosaurs were essential components of their ecosystems and it is easy to think that all they did was eat prey animals – but this is far from the truth. Without the carnivores then chaos would soon ensue. Theropods evolved to eat their contemporaries -not withstanding therizinosaurs which we will come to a little later – and they were obviously very successful at it since they were dominant throughout the dinosaurs reign but we often forget, that throughout the Mesozoic, there were a multitude of different environments. Theropods evolved in tandem with deserts, forests, coastal plains, uplands, lowlands and everything else in between and every single ecosystem depended on them to function correctly.
Theropods were the major controllers of herbivorous dinosaur populations and it is safe to assume that the rules that govern the trophic pyramids of today would have applied to those of the past. I find it particularly interesting that we consider many aspects of dinosaur extinction, not only at the K-T boundary, but also the various phases of dinosaur extinction throughout the Mesozoic without often considering environmental disruption because a species, particularly a carnivore, became extinct. We all consider environmental change due to volcanoes, earthquakes, mountain building and even the occasional meteor strike, and no doubt these were considerable influences on the environment, but extinction due to the disappearance of species is well documented – even today.
In the next part of this mini-series we will look at the cause and effects of theropod extinction on an ecosystem and consider whether this too was a major driver in dinosaurian evolution and species turnover.
Russell, Dale A. (1970). "Tyrannosaurs from the Late Cretaceous of western Canada". National Museum of Natural Sciences Publications in Paleontology 1: 1–34.