The Largest Living Systems

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For anyone who studies evolution, it is important to realize that there are characteristic evolutionary patterns. For example, evolution tends towards greater complexity (although not always). Evolution also has a variable speed (which is often contingent on the environment). And a study recently published in PNAS indicates that evolutionary processes generally select for species-level living systems with universal size distribution. Science Daily summarized the importance of this universal size distribution well:

Flocks of birds, schools of fish, and groups of any other living organisms might have a mathematical function in common [… researchers] showed that for each species studied, body sizes were distributed according to the same mathematical expression, where the only unknown is the average size of the species in an ecosystem.

For the researchers of this study, these apparent universal size distribution may be useful for understanding how systems of living matter operate. However, this study made me think of the role of size in evolutionary processes. Specifically, what causes different living systems to evolve different sizes? And what living system has evolved the largest overall size?

The role of size in evolutionary processes has always been a contentious issue for evolutionary theorists. Central to the issue of size has been the idea that natural selection tends to drive the evolution of larger and larger overall size, regardless of whether the living system is a bacterium, a hydra, or a chimp. This observed trend has been labeled Cope’s rule after Edward Cope, a 19th century paleontologist who first proposed the trend. The late evolutionary theorist Stephen J. Gould disregarded Cope’s rule as a “psychological artifact”, however recent studies have provided empirical evidence to support the general pattern.

Paleontologist Joel Kingsolver supports the idea that evolution tends to favour large body size, stating that:

In 80 percent of the studies, there’s consistent selection favouring larger size.

Disappointingly, the theory to explain this pattern is still underdeveloped. In fact, Kingsolver contends that there may not be any universal driver of larger body size:

My guess is that it’s a mix of particular reasons for particular speices. You may be able to make through lean times better than someone who’s smaller. Females that are larger are able to produce more eggs. If males are competing for females, larger size is often favoured.

Paleontologist and science blogger Brian Switek echoed a similar perspective recently in an article about large dinosaur body size:

The evolutionary driving forces behind the evolution of truly huge body size are not clear, and likely differed from one group to the next.

Although evolutionary theory explaining the drive behind selection for larger body is underdeveloped, we do have a better idea of proximate determinants of body size. For example, many theorists have demonstrated that mode of locomotion and reproduction are both important factors either constraining or enabling large body size.

As Brian Switek discussed at length recently, the monstrous sauropod infraorder was able to “sidestep” the costs and risks that constrain mammalian size by “externalizing birth and development.” The size distribution of sauropods dwarfed the size distribution of all other known terrestrial organisms to ever exist.

So of these supermassive sauropods, what species holds the title of largest? The answer to this question was far more difficult to find than I originally thought. Michael Stevens from VSauce recently claimed that Giraffatitan was the largest known “with certainty of a complete skeleton”. Estimates of Giraffatitan come from one skeletal sample, and was thought to be 72-74 feet in length and weigh ~30-40 tons. Compare that to the largest known African elephant which weighed ~12 tons.

However, there is general consensus in the paleontological community that there were larger sauropods than Giraffatitan. Thankfully, I had some help from Brian Switek to better understand the contemporary debate:

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According to Switek Argentinosaurus and Supersaurus are the leading contenders for heavyweights in the dinosaur world. The longest known of these giants was a Supersaurus that is estimated to be 108-111 feet long. The heaviest was a Argentinosaurus estimated to have weighed 73 tons. They were the giants of the gigantic sauropoda order.

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But we can’t forget about a living clade of animals that has experienced an explosive increase in size distribution: cetaceans. The largest (by far) of our mammalian cousins is the blue whale. And the blue whale is not just a contender for largest living animal, they are also contenders for largest animal of all time. In fact, in terms of absolute weight, it doesn’t appear to be close at all. Whereas Argentinosaurus weighed 73 tons, the largest known blue whale weighed over 200 tons! More than double the weight of the largest known dinosaur! But to be fair, blue whales don’t have to worry as much about the crushing weight of Earth’s gravity. The battle is much closer when we compare length: Supersaurus was between 108-111 feet and the largest known blue whale was ~110 feet.

blue_whale_1273905983_770.jpg Blue whale Balaenoptera musculus = heaviest of all time?

The SV-POW paleontology blogger team made a brilliant point that we should suspect that Supersaurus was on average longer than blue whales because we are comparing with biased sample sizes:

A huge sample of blue whales included none longer than 110 feet, while our comparatively pathetic sample of sauropods has already turned in one animal (Supersaurus) that may have just edged that out, and another (A. fragillimus) that - assuming it was really as big as we think - blows it out of the water.

In case you were wondering, A. fragillimus is estimated to have been between 130-200 feet long! It completely blows my mind that a terrestrial organism can reach those sizes on our planet (just imagine how big sauropods would have been if they had evolved on planet the size of Mars!).

Longest_dinosaurs1.png The red image represents A. fragillimus, potentially the longest organism ever

In case you were wondering, no primate species has ever been a contender for largest living system. The primate order is comparatively small, with the largest contemporary species (gorillas) weighing between 300-400 lbs (or about 0.15-0.2 tons!). Even if we consider extinct species, no primate has ever even been a contender for largest land mammal. The largest, Gigantopithecus, weighed approximately 1,200 lbs (or about 0.6 tons). Of course, I think Gigantopithecus is aptly named (and I think sympatric populations of Homo erectus would agree); but they are only aptly named in comparison to our relatively puny order. Primate size has probably always been constrained by underdeveloped quadrupedalism and selection for long-term infant dependence.

2007-06-28_133054.png Reconstruction of Homo erectus and Gigantopithecus in Southeast Asia

However, it is interesting to know that all species body sizes (from prokaryotes to sauropods) are distributed according to a potentially universal power law. This universal describes how ecology influences average species size, while genetics contains variability around that average. In the future, I’ll be interested to see whether evolutionary theorists can better describe the adaptive pressures selecting for larger size. It is useful to have a grasp on the proximate causes of body size, but the ultimate causes will be necessary to better describe how living systems develop over time.

Discuss this on Hubski or let me know what you think on Twitter!

 
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