Arthropleura: The Titan of the Carboniferous

Artists reconstruction of the Carboniferous Titan Arthropleura
Image Credit: Nobu Tamara, http://spinops.blogspot.com/

One day, two executives at Warner Brothers film studio are sitting in their office with their heads in their hands. The last film has massively under-performed at the box office, the writers are struggling for ideas, A-List actors are turning their noses up at working for them and their film sets are so quiet you can hear the crickets chirping. However there is a knock at the door. “Come in!” shouts one of the executives. A nerdy looking man; wearing a shirt with an otter on the front, sport shorts that don’t match his top and sandals of a hideous brown and orange colour, steps in to the room. “Hello Brothers Warner! I have a script I’ve written for a potential film, its a mix of Sci-Fi and horror” he exclaims confidently. One of the executives sighs “alright, lets see it”. “Can’t possibly be worse than what we’ve just put out” the other remarks. The executives flick through the script, reading the tale of an adventurer hacking her way through the lush forest of a strange, almost alien world. The adventurer encounters plenty of dangers; ducking a dragonfly the size of an eagle dive-bombing for her head, narrowly dodging the thrusting stinger of an ambushing scorpion the size of a large house-cat and throughout the film she is pursued by a large, unknown monster. At the films climax the adventurer, remarking that she had seen it all now, stumbles on what seems like a large log. She turns round and watches in horror as the log rises and squirms. The monster is revealed! It is a millipede, which rears up high enough to meet her eyes. It hisses, ready to lunge!

“Wow! This script has serious promise! even if it is a little cheesy” One of the executives gasps. “But where is it set?” The other enquires. “Oh turn to the last page!” The geek squeals excitedly. The executives do so, and see the words that would mark the twist ending to this film.

Earth. 300 Million Years Ago.

It sounds absurd, but believe it or not 300 Million years ago planet earth was home to these enormous arthropods. Meganuera and Pulmonoscorpius, the dragonfly and scorpion in our geek’s story, are both fascinating animals. However the most striking of these arthropods, and the one that I have a personal story of, is our monstrous millipede, which has the scientific name Arthropleura, meaning “jointed ribs”.

First discovered in 1854 by Jordan & Meyer, there are two known species of Arthropleura. While the smaller species was only a quarter of a metre in length the larger species was the biggest terrestrial arthropod of all time. Measuring just over two and a half metres long and nearly half a metre wide it was a size that modern millipedes could only dream of. It had a body plan of 30 jointed segments and each segment was covered by a relatively thin armour plating. Its large size meant that it would have had few natural predators (unless one of the large amphibians of the time got in a lucky shot). Since its discovery the main controversy surrounding Arthropleura was whether it was related to millipedes or centipedes, and as a result whether it was a herbivore or a carnivore. This is due to no mouth-parts having been found as of yet. While remains of giant club mosses had been found in some fossil remains a relatively recent study by Kraus showed that this was actually the shed skin of an Arthropleura that had been deposited on top of these club moss fragments. However despite this the common consensus at the moment is that it is a millipede relative and as such a herbivore, feeding on dead plant matter like modern millipedes do today (although its jaws would have given you a bad bite!). Arthropleura first evolved around 315 million years ago and went extinct 299 million years ago. The main reasons for its extinction being a combination of the disappearance of the coal swamps that it resided in and reduction in oxygen levels.

A fossil of the armour plates of Arthropleura from the Senckenberg Museum of Frankfurt in Germany. Image Credit: Ghedoghedo, https://commons.m.wikimedia.org/wiki/File:Arthropleura_armata.jpg

Unfortunately such an animal really was a product of its time. Arthropleura and other arthropods could only grow to such a size because of the special environment conditions present during the Carboniferous period. The earth had a much higher concentration of oxygen compared to modern times (almost 35%, compared to about 21% today). This suited arthropods in particular as it meant that they could take in more oxygen and therefore have more energy via respiration to use in growth. Arthropods take in oxygen via tiny tubes on the side of their bodies, known as trachea, or even directly through their skin. This system is nowhere near as efficient as true lungs are in terrestrial vertebrates. As a result the lower oxygen levels of today means that arthropods simply can’t get enough energy from respiration to maintain such large sizes. If Arthropleura was alive today it would not be able to survive in such a low oxygen environment. However a large concentration of oxygen isn’t the sole reason for their size, as a lack of competition and predation could also have helped.

Fossilised tracks of Arthropleura such as these are found in places in Scotland (e.g. the Island of Arran) and the USA. Image Credit: Ashley Dace, https://www.geograph.org.uk/photo/1994274

To finish this blog i’m going to share my personal encounter with an Arthropleura! Or rather, its fossilised tracks. When I was around the age of 11, during a family holiday in Scotland, I went to see the Arthropleura tracks at Crail. My young palaeontology mad self loved seeing these tracks, and there is a photo that proves it. So as it turns out this animal is indeed Scottish! (well half-Scottish technically, as fossils and trackways have also been found in the USA, but I like to think its Scottish). Arthropleura tracks are also a famous attraction of the island of Arran in Scotland. Whilst I did go to Arran during the first year of my university degree sadly my group didn’t have time to go see them.

So Arthropleura is yet another example of the weird creatures that evolution has produced, and a true marvel of the distant past. In my humble opinion, it would make a fantastic movie monster!

References/Further Reading

A great blog on Arthropleura written by Hans-Dieter Sues, a Paleontologist based at the National Museum of Natural History in Washington DC. The blog was published on National Geographic

Sues, Hans-Dieter,”Largest Land-Dwelling “Bug” of All Time”, National Geographic, Jan. 15, 2011, blog.nationalgeographic.org/2011/01/15/largest-land-dwelling-bug-of-all-time/

A blog, published on National Geographic, which gives futher background on the Carboniferous Period

National Geographic, “Carboniferous Period”, National Geographic, nationalgeographic.com/science/prehistoric-world/carboniferous/

A paper from 1992, written by Paul Pearson (Pearson 1992) and published in the Scottish Journal of Geology describing Arthropleura tracks (named Diplichnites cuithensis) from Fife, Scotland

Pearson, P. N. (1992). “Walking traces of the giant myriapod Arthropleura from the Strathclyde Group (Lower Carboniferous) of Fife.” Scottish Journal of Geology 28(2): 127-133.

Another paper, written by Ronald Martino & Stephen Greb (Martino & Greb 2009) and published in Journal of Paleontology, describing a set of Arthropleura tracks, this time from Kentucky, USA

Martino, R., & Greb, S. (2009). Walking trails of the giant terrestrial arthropod Arthropleura from the Upper Carboniferous of Kentucky. Journal of Paleontology, 83(1), 140-146. doi:10.1017/S0022336000058200

Is it a shrimp?! Is it a jellyfish?! No its Anomalocaris!

Science is not always static. Like any living species that has, currently or will exist it is constantly changing over time, with scientific theories evolving to fit the best available evidence. This phenomenon is prevalent throughout the scientific world but one area where it can be very clearly seen is in Palaeontology, where there are many instances of reconstructions of extinct life being very different in the past than they are today. One example of this is Megalosaurus, one of the first dinosaurs to be properly described by science. The original Victorian interpretation can be seen in a full scale model at the Crystal Palace in London. It is an impressive sculpture of a big hulking four legged lizard, portrayed as the Victorian scientists interpreted it, however it is nothing like the more graceful reconstruction nowadays. Another example, whose outdated model can be seen in the Sedgwick Museum in Cambridge, is the Carboniferous arthropod Megarachne (“Great Spider”). Once thought to be the largest spider that ever lived new research in 2005 found it to instead be a small to medium sized species of freshwater sea scorpion (though in my opinion this doesn’t make Megarachne any less unsettling to look at!)

However one of the most interesting cases of an extinct animal whose palaeontological reconstruction has changed greatly over time with new discoveries has to be Anomalocaris, whose name means “unusual shrimp”. Anomalocaris lived approximately 505 million years ago during the Early-Mid Cambrian. The Cambrian, from an evolutionary perspective, was prehistories equivalent of the European Renaissance of the 15th century. New body plans and weird evolutionary experiments were emerging during this period of earths history. Some animals from this time became extinct not long after they appeared. Some, like the trilobites, survived for an amazingly long period of time (trilobites appeared 520 million years ago in the Early Cambrian and went extinct during the Late Permian 250 million years ago – a longevity of 250 million years!) While others would eventually evolve and diversify into the main animal groups alive today; including the vertebrates which humans are a part of.

Anomalocaris is one of these weird wonders. It was discovered in 1892 by Joseph Frederick Whiteeves in the Burgess Shale formation, a fossil lagerstätten (which is a site where a high concentration of fossil material is preserved due to special environmental conditions) in Canada. The original fossil, named Anomalocaris canadensis, looked like a shrimp but with no clear headparts, hence its name of “unusual shrimp”. For a long time Anomalocaris was only known from this basic description and while it was definitely strange, its lifestyle was a complete mystery. Nineteen years later in 1911 the palaeontologist Charles Walcott, who is famous for his extensive work on the Burgess Shale, discovered a fossil of what seemed to be a primitive jellyfish. He gave it the name Peytoia nathorsti. Later, in 1928, Laggania cambria was discovered; the name given to a long bodied fossil that was interpreted as a relative of sea cucumbers.

Now you may be wondering “I thought you were going to be talking about Anomalocaris? Why have you wondered off topic to these random animals?” Well this is where the story gets interesting! In the early 1980s, nearly 90 years after Anomalocaris was first named, a palaeontologist working at the University of Cambridge by the name of Harry Whittington, an expert in Cambrian arthropods of the Burgess Shale, saw something that was truly astounding while preparing a fossil from the Burgess Shale. As he chipped away at the rock he saw two Anomalocaris “shrimps” attached to the head of a larger body not too dissimilar to Laggania. Not only that but a Peytoia fossilwas found to be attached to this same head. It soon became clear that Anomalocaris, Peytoia and Laggania were not separate species, but all part of one huge Cambrian animal, which was given the name Anomalocaris as that had been the original fossil part that had been found.

Anomalocaris was the top predator of its day. At around a metre in length it was the largest single animal the earth had ever seen at that point. After identifying its prey using large compound eyes, which gave it excellent eyesight comparable to modern day insects, it then used its prongs, once thought to be shrimps, to grab and hold its prey. Anomalocaris then held the prey close to its mouth-parts, once thought to be Peytonia, so the mouth parts could rip and break through the hard exoskeletons of trilobites and the soft bodies of other Cambrian arthropods that made up its prey. Anomalocaris swam via undulatory movements of their regularly arranged horizontal side flaps in the same manner that modern day soft bodied marine invertebrates do today. Anomalocarids as a group were widely successful, ranging across the globe from Canada to China and living from the Early to Middle Cambrian period. While most Anomalocarids were predators, another species has been described relatively recently in 2014 and named Tamisiocaris borealis (“sieve shrimp”). It had a very different lifestyle to Anomalocaris, possessing bristles on its prongs which it’s thought to have used in filter feeding, behaving rather like the baleen whales of today. This makes Tamisiocaris the earliest example of a large filter feeding animal known to science.

So Anomalocarids, the weird shrimps of the Cambrian, really are a fascinating group of arthropods. Once thought to be multiple separate animals, Anomalocaris and other Anomalocarids have instead been shown to be one of the weirdest of all Cambrian animals, and a true example of the evolutionary variety that has evolved on this planet.

The strange shrimp itself
Image credit: UNE photos, https://www.flickr.com/photos/unephotos/6786859303

EDIT: In this blog I state that Anomalocaris could “rip and break through the hard exoskeletons of trilobites”. This is actually wrong! A study in 2010, led by James Whitey Hagadorn from the the Denver Museum of Nature and Science, used 3-D models of Anomalocaris‘ mouth-parts to show that (a) it couldn’t close its mouth the whole way and (b) its mouth was too delicate to crush hard exoskeletons. As a result Anomalocaris would have fed mostly on soft bodied animals, and maybe even trilobites that had just moulted (and therefore had softer shells).

References/Further Reading

A blog, written by Ed Yong and published in Discover Magazine, about Anomalocaris

Yong, Ed, “The sharp eyes of Anomalocaris, a top predator that lived half a billion years ago”, Discover, Dec. 7, 2011, discovermagazine.com/the-sciences/the-sharp-eyes-of-anomalocaris-a-top-predator-that-lived-half-a-billion-years-ago#.XW6b8S5KjIU

The official Burgess Shale website page about Anomalocaris

“Anomalocaris canadensis”, burgess-shale.rom.on.ca/en/fossil-gallery/view-species.php?id=1

Vinther et. al. 2014 paper, published in the journal Nature, on the filter feeding Anomalocarid Tamisiocaris

Vinther, J., Stein, M., Longrich, N. et al. A suspension-feeding anomalocarid from the Early Cambrian. Nature 507, 496–499 (2014). https://doi.org/10.1038/nature13010

A wired article talking about the Hagadorn 2010 study that showed that Anomalocaris couldn’t eat hard bodied prey (USED FOR THE EDIT)

Mosher, Dave, “Giant Vicious-Looking Ancient Shrimp Was a Disappointing Wimp”, Wired, Mar. 11, 2010, wired.com/2010/11/anomalocaris-trilobite-bite/

Original Hagadorn study: Hagadorn, J. (2010). Putting Anomalocaris on a soft-food diet. 2010 GSA Denver Annual Meeting.