Cambrian Explosion Month #09: Phylum Hemichordata – Enteropneusta

Enteropneusts, commonly known as acorn worms, are the most numerous group of modern hemichordates with over 100 known species. Most of them burrow in sediment eating organic detritus, but a few are filter-feeders and some deep-sea species crawl and drift around over the sea floor.

Their fossil record is poor due to their soft bodies, but the transitional form Gyaltsenglossus has recently given us a glimpse at acorn worms’ ancestral links with their cousins the tube-dwelling pterobranchs.

But that’s not the only fossil hemichordate with surprising traits from both lineages. It turns out the characteristic tubes of pterobranchs may actually have been ancestral to all modern hemichordates – with the acorn worms later secondarily losing the ability to make them.

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Cambrian Explosion Month #08: Phylum Hemichordata – Pterobranchia

Now we get to the bilaterians, all the animals with bilateral symmetry as an ancestral feature.

The hypothetical common ancestor of all bilaterians (the “urbilateria“) was probably a tiny worm-like species, and likely originated sometime in the early Ediacaran Period. The earliest definite body fossils of bilaterians come from about 558 million years ago, and possible burrow traces are a little older, from about 585 million years ago – but it was during the Cambrian Explosion that this group rapidly diverged into a wide variety of forms and ecological niches.

The deuterostomes are one of the two major evolutionary branches of the bilaterians, made up of modern hemichordates, echinoderms, and chordates (including the vertebrates). They split from their last common ancestor with the protostomes sometime during the Ediacaran, but the earliest probable deuterostome fossils are weird tiny potato-like blobs from the very start of the Cambrian.

(…We’ll talk about those later in the month.)

Hemichordates are the closest living relatives to echinoderms, and are a small phylum with only around 130 known modern species (most of which are acorn worms). But in the distant past they were much more numerous, with graptolites being major components of Paleozoic planktonic ecosystems.

All living members of this lineage have a three-part body plan, but they’re otherwise very different in both appearance and ecology. Acorn worms are solitary worm-like animals living on in the sediment of the sea bed, and are mostly detritivores, while pterobranchs are tiny colonial filter-feeders that build protective tubular structures.

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The protocetids were some of the first oceanic cetaceans, occupying a transitional position in the evolution of whales, with four paddle-like limbs and nostrils only partway up their snouts.

Early members of this group swam like otters, using a combination of undulating their bodies and paddling with large hind limbs, but somewhere in the Late Eocene they switched over to propelling themselves entirely with their tails and gave rise to even more whale-like forms like the basilosaurids.

And Aegicetus gehennae was right in the middle of that switch.

Discovered in the Wadi Al-Hitan (“Valley of the Whales”) fossil site in Egypt, Aegicetus lived around 37-35 million years ago. It was similarly-sized to earlier protocetids like Georgiacetus, measuring about 3.5m long (11’6″), but its hind limbs were proportionally smaller. Its hips were also completely disconnected from its vertebrae, giving it much more flexibility to undulate its body and tail – and preventing it from supporting its weight on land, suggesting that it spent its entire life in the water.

It wasn’t a direct ancestor to more “advanced” cetaceans, since it lived alongside several species of basilosaurids. Instead it seems to represent a late-surviving example of what the earlier protocetid-basilosaurid transitional forms would have looked like.


Mixosaurus cornalianus here was a small early ichthyosaur, only about 0.8-1m long (2’7″-3’3″) and generally considered to be transitional between the eel-like swimming style of basal forms and the more dolphin-like later forms. Living during the Middle Triassic, about 242 million years ago, it inhabited a shallow tropical sea that covered what is now the modern border between Switzerland and Italy.

It was previously thought to be a slow swimmer with a low and poorly-developed tail fin, and whether it even had a dorsal fin or not was unclear. But now new specimens with soft tissue impressions have given us a big surprise.

Not only did it actually have a fairly well-developed semilunate tail fin, but it also had a dorsal fin positioned much further forward on its body than expected, giving it a shape similar to some small sharks and representing the current earliest known dorsal fin of any amniote.

Bundles of stiffening collagen fibers inside its fins were very similar to those known from later Jurassic ichthyosaur species, indicating that this adaptation evolved much earlier in the lineage than previously thought. Along with stomach contents showing it mainly ate both cephalopods and small fish – fairly fast-moving prey – this suggests it was a capable open-water swimmer. It wouldn’t have been quite as speedy as its much more specialized Jurassic relatives, but it may have still been about as efficient as the small modern sharks it resembled.


Since the last couple of weeks have featured marine mammals, let’s have one more! This time not a cetacean but a member of the other group of fully aquatic mammals still alive today: the sirenians.

Although commonly known as “sea-cows” due to their herbivorous grazing habits, sirenians’ closest living relatives are actually modern elephants. They’re thought to have originated in Africa over 50 million years ago, starting off as pig-like or hippo-like semi-aquatic animals — but they must have been good swimmers capable of crossing oceans very early in their evolutionary history, since some of the earliest known sirenian fossils actually come from the other side of the Atlantic on the Caribbean island of Jamaica.

Sobrarbesiren cardieli here extends some of our knowledge of early four-legged sirenians to Europe, dating to the mid-Eocene about 42 million years ago. Hundreds of bones were found in Northeastern Spain, representing at least six different individuals and giving us a fairly complete idea of this species’ anatomy.

It was smaller than modern sea-cows, reaching about 2m long (6’6″), and seems to represent a transitional point between the semi-aquatic ancestral sirenians and fully aquatic later forms. It had a head very similar to its modern relatives, and probably a tail fin, but also still retained small functional hind limbs.

It was initially thought to still be somewhat semi-aquatic and capable of quadrupedal locomotion on land, but a later analysis of its hind limb bones suggests that it may actually have been much more aquatic than that. Its hind legs had a wide range of motion and were probably used for otter-like swimming, undulating the body while paddling, but might not have been capable of supporting its weight on land. So if Sobrarbesiren did still haul out of the water, it may have had to move more like a seal.


We have a fairly good picture of the evolutionary origins of most groups of aquatic mammals – except for the pinnipeds. The fossil record of early seals is still rather sparse, and for a long time the earliest known species was Enaliarctos, an animal that was already very seal-like and didn’t help much in figuring out whether seals’ closest living relatives are bears or musteloids.

But then Puijila darwini was found in the late 2000s, a transitional form with a near-complete skeleton, filling in a gap in our understanding so conveniently it almost seems too good to be true.

This is the equivalent of Archaeopteryx for seals.

Discovered in Nunavut, Canada, Puijila dates to the early Miocene, about 23-20 million years ago. It was a small freshwater otter-like animal, about 1m long (3’3″), with a long tail and webbed feet adapted for paddling with all four of its limbs.

It lived at around the same time as the more specialized Enaliarctos, so it wasn’t a direct ancestor of modern seals, instead being part of an early offshoot lineage that retained more basal characteristics – but it does gives us a clue as to what the earliest pinnipeds looked like. Along with genetic studies it also helped to clarify that seals’ closest relatives are indeed the musteloids, although they’re estimated to have last shared a common ancestor around 45 million years ago so there’s still a lot of time unaccounted for in the proto-seal fossil record.

Several other fossil species that were previously thought to be musteloids have now also been recognized as close relatives of Puijila, and it seems that they were a fairly widespread group basically filling the ecological niche of otters at a time before true otters existed.

Most surprising and frustrating of all, however, is that some of these other otter-seals actually survived all the way into the Pleistocene, only going completely extinct sometime in the last 2 million years.

We barely missed having them still alive today!


Enaliarctos mealsi, an early seal from the Late Oligocene and Early Miocene of California, USA (~23-20 mya).

Measuring about 1.5m long (5′), it was a transitional form between modern seals and their more otter-like ancestors. It was well-adapted for swimming with a flexible spine and flipper-like limbs, but unlike most modern pinnipeds it probably used both its front and hind flippers for propulsion.

Its teeth also still resembled those of terrestrial carnivores, with slicing carnassials at the back of its jaws. This suggests that it had to drag larger prey items back to shore in order to tear them apart and eat them, similar to the behavior of modern otters.


Westlothiana lizziae from the Early Carboniferous of Scotland (~338 mya).

About 20cm in length (8″), this superficially lizard-like creature (nicknamed “Lizzie” by its discoverer) had a long slender body with relatively small legs, which may have been adaptations for burrowing similar to modern skinks.

Its anatomy shows a mixture of both amphibian and reptilian characteristics, suggesting it may have been a close relative of the first true amniotes. But exactly where it fits in that area of the evolutionary tree is still uncertain, with different paleontologists classifying it as either an early amniote, a reptilomorph, or a lepospondyl.