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Enigmatic Typhloesus wellsi: Now transitional between lancelets and sea cucumbers

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Today
the ‘alien goldfish’ and late-surviving Late Carboniferous, Typhloesus welsii (Melton and Scott 1972, Conway Morris 1990), goes under DGS (Figs. 1, 4). The aim is to tease out a few more details than previously traced during earlier binocular examinations fifty and thirty-two years ago… before Photoshop’s ability to highlight structures became available to workers.

Figure 1. Typhloesus as traced by Conway Morris 1990 and with details added by DGS. ” data-image-caption=”

Figure 1. Typhloesus as traced by Conway Morris 1990 and with details added by DGS.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus_wellsi_insitu_recon588.gif?w=252″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus_wellsi_insitu_recon588.gif?w=584″ src=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus_wellsi_insitu_recon588.gif” alt=”Figure 1. Typhloesus as traced by Conway Morris 1990 and with details added by DGS. ” class=”wp-image-62193″ />

Figure 1. Typhloesus as traced by Conway Morris 1990 and with details added by DGS. Length = 9cm. Here the misidentified foregut is the notochord. Other overlooked parts are shown in various colors. The pink structures over the eye could be benthic debris, until a similar structure appears on another specimen of Typhloesus. See figure 4 for an overall enlargement rotated 90º.

Based on tracings presented here
(Figs. 1, 4), Typhloesus is transitional between lancelets and sea cucumbers sharing traits once considered unique to each type.Let’s compare traditional interpretations with several novel ones offered here. Please note that several were proposed by Melton and Scott 1972, then later misinterpreted by Conway Morris 1990.

Backstory:
Earlier we looked at the origin of echinoderms from lancelet ancestors evolving first into sea cucumbers (Fig. 2), then on to other echinoderms, from starfish to crinoids. So Typhloesus merely cements and adds detail to a relationship already documented here.


Figure 2. The lancelet, Branchiostoma, and a selection of sea cucumbers, See figure 5 for a larger diagram of the lancelet.

From the Conway Morris 1990 abstract:
Typhloesus wellsi is a bizarre metazoan from the mid-Carboniferous (Namurian) of central Montana, U.S.A.

=Yet another Bear Gulch Limestone odd and unique taxon.

Typhloesus gen. nov. is erected here, because the previous generic names used to describe this animal refer to conodonts, enclosed within the alimentary canal, that were ingested as a result of predation. All these names are invalid in the present context. Its fusiform body (maximum length ca. 90 mm) bore a paired ventral keel, interrupted at about two thirds of its length by a conspicuous break. At the posterior a prominent fin was supported by two sets of orthogonally disposed fin rays in an arrangement unique to this animal.

Here (Fig. 1) that prominent posterior ‘fin’ is identified as an organ of cloacal cilia. These are similar to anal spines or respiratory surfaces, as in certain sea cucumbers (Fig. 3), which are known to ‘breathe’ through the anus by ‘inhaling’ and ‘exhaling’ water through the posterior portal driven by expansion and contraction of the entire body. Typhloesus has a large atrium, ideally suited to do this.

“The exterior lacked other appendages, although lateral fins may be inferred on hydrodynamic grounds.”

Try not to ‘infer’ on shaky grounds. Typhloesus was not a swimmer and did not need to be hydrodynamic. It was shaped like a basal sea cucumber.

The internal anatomy consisted of an elongate foregut and a voluminous midgut.

On closer examination that ‘voluminous midgut’ is an extended, curled-up intestine,
as in sea cucumbers (Fig. 3).

The mouth was ventrally directed and apparently large.

Yes, and the mouth is surrounded by buccal cilia,
as in lancelets and sea cucumbers (Fig. 2).

“The gut appears to have been blind, lacking a hind section and an anus.

On closer examination, the winding intestines largely localized the midgut actually finds a posterior exit to a large intestine oriented ventrally to the former lancelet cloaca, but then a smaller intestine continues along the ventral margin, terminating in the cloacal cilia.

Located beneath the central midgut was a remarkable organ composed of two discs, known as the ferrodiscus, and attached to the ventral margin. Its function is obscure, but its association with possible circulatory vessels suggests a role in storage or respiration.

These discs might be exactly what they seem to be: small sacs to contain bits of iron that cannot be digested from ingested benthic debris. Occasionally these discs might be emptied by expulsion through the nearby atrial pore… or not.

“Obscure traces of possible mesenteries, longitudinal muscles and a cuticle are discussed.

The swimming muscles that power lancelets are missing
here in this sedentary, sea-cumber-like taxon.

Typhloesus gen. nov. is interpreted as an active swimmer, with lateral propulsive oscillations, probably largely confined to the posterior area where the rigid fin is inferred to have played an important role.

That’s no fin, and swimming muscles are missing, so Typhloesus was sedentary.

Gut contents of conodont apparatuses, variously disarticulated, fish and worm teeth suggest this animal was a predator and scavenger, competing with other members of this trophic group in the shallow waters of the Bear Gulch ‘lagoon’.

Rather, this taxon was likely a garbage collector, sweeping up dead conodonts, fish and worms from the sea floor. Typhloesus was not built for speed or fight. It led a quiet, but important life cleaning up dead things.

Previous suggestions that Typhloesus gen. nov. is a chordate cannot be supported, but the phyletic affinities of this extraordinary creature are unresolved.

A notochord is preserved. It starts with the overhanging rostrum and continues behind the intestines (Fig. 1).

It joins a select group of late-Palaeozoic animals whose peculiar appearance prohibits convincing comparisons with known phyla.

To the contrary! Unfortunately in 1990 Conway Morris did not have access to Photoshop, which enables parts of the photo to be increased in contrast and the whole photo to be traced in various colors (= digital graphic segregation). No doubt he examined Typhloesus under a binocular microscope, resulting in the outline of the taxon (Fig. 1). Here a computer monitor does the magnifying and Photoshop applies colors to graphically segregate elements, even those lying behind others. It’s a step-by-step process, something the human eye and brain are incapable of doing. That’s why we all trace fossils. DGS simply does this with colors.

These species, none of which is closely related to any of the others, could be survivors from the ‘Cambrian explosion’ in which a multitude of bizarre body-plans evolved, or they could represent the results of adaptive radiations nearer to the Carboniferous.

Typhloesus is a late-survivor in the Middle Carboniferous
of an Ediacaran to Early Cambrian radiation.

Absence of ancestors and taphonomic bias in favour of soft-part preservation in nearshore environments makes it difficult to resolve these possibilities, but the latter option is tentatively preferred.”

There is no ‘absence of ancestors’. Extant taxa need to be employed, not just fossils. Typhloesus is clearly transitional between lancelets and sea cucumbers, sharing traits otherwise unique to each taxon.


Figure 3. Sea cucumber diagram. Note the spiky cloaca and serpentine intestine, both as in Typhloesus.

Writing online in 2016, Christopher Taylor reports,
“Typhloesus is represented in the[Bear Creek Formation] deposit by a number of individuals in varying states of preservation.”

“By the time Typhloesus was reviewed in detail by Conway Morris (1990), it was clear that the conodont fossils had been preserved within its gut, not its mouth, and Typhloesus was a conodont-eater rather than a conodont-bearer (it has since been found that conodont animals were eel-like chordates).”

“Externally, Typhloesus was a fairly simple, cigar-shaped animal, with its body laterally compressed and higher than wide. It grew to a decent size, with the largest specimens being a little under ten centimetres in length. There is no sign of eyes or any other prominent sensory structure, and so far as is known the external skin or cuticle was smooth and unornamented.”

To the contrary, DGS revealed a sclerotic ring (for eyeballs), a brain dorsal to the notochord and sensory extensions emanating from it.

“The most distinctive external feature is a large ‘tail-fin’ at the rear. This fin was supported by an arrangement of criss-crossing rods or fibres, and would have been fairly stiff in life.”

Not a ‘tail-fin’, but a sea-cucumber-like clocal structure., perhaps eversible.

“Another pair of folds or fins ran along most of the underside of the body with a noticeable gap towards the rear.”

Not paired fold or fins. These are likely the remains of gill slits as they appear to be exits for the large ventral atrium.

“Typhloesus probably swam in a not dissimilar manner to an active modern fish, using sweeps of the tail-fin to provide thrust; the ventral fins may have provided stability and steerage.’

No swimming was likely. That’s not a tail fin. Some awkward, but sudden movement was likely possible by filling the atrium and expelling a jet of water through the atrial pore.

“The visible line of the foregut comes to a halt slightly before reaching the front of the body, and it seems that the mouth would have been slightly ventral and contained within a ‘hood’.

As in lancelets.

“Though its overall conformation and known gut-contents (most commonly conodonts, but sometimes worm jaws or fish scales) suggest an active predator, I am at a loss to understand how it located its prey without eyes. Perhaps the hood contained some sort of chemical sensors in life.”

Typhloesus was a benthic debris feeder. Anything small enough to fit into its mouth.

“When it was first found, it was thought that its overall appearance suggested a relationship of Typhloesus to the chordates.”

That hypothesis of interrelationships is supported here because a notochord is present.

“However, Conway Morris (1990) saw its internal anatomy as incompatible with this view. Fossils of this animal show a narrow foregut leading into a voluminous, sack-like midgut.”

There is more to see inside Typhloesus than Conway Morris saw (Fig. 1). Let’s be scientists, not journalists. Let’s not believe everything we read. Rather, test hypotheses and observations with your own scientific observations and data.

“Below the midgut is a pair of dark, disc-shaped organs showing a concentration of iron deposits called the ferrodiscus; though a striking element of all Typhloesus fossils, the function of this structure is completely unknown.”

Why not just consider these discs just as the are: deposits of iron because they are unsuitable for the rest of the organism. They may be periodically emptied by expulsion from the nearby atrial pore. Or not.

“What Conway Morris found conspicuous by its absence, however, was an anus: there appeared to be no sign of any gut structures in the rear of the animal.”

A terminal gut and anus is found under closer examination.

“The gut was a blind sack, with the only way out being the same as the way in.”

Not true. See Fig. 1.

“The absence of a through-gut would be unprecedented in a chordate, or indeed in many animals except jellyfish or flatworms.”

So, that’s a Red Flag that indicates a closer examination is warranted.

“Conway Morris was also unable to identify other chordate-specific structures such as muscle-blocks, gill openings or a notochord; though he confessed that the first two might be obscured by the vagaries of decay, he felt that the third at least should have left more of a sign. It was this combination of an overall fish-like appearance with a very un-fish-like anatomy that led Conway Morris to later dub Typhloesus the ‘alien goldfish’.

Conway Morris did not have the post-1995 advantage of Digital Graphic Segregation, which enables one to pick apart crushd taxa, like Typhloesus without picking it apart.

“With the exclusion of a chordate connection as a possibility, Conway Morris found himself at a loss as to just where Typhloesus fitted into animal evolutionary history.”

Add taxa. Look more closely. Don’t quit. Don’t give up. I’m surprised Conway Morris has not reviewed his original observations in the past 30 years.

Quick note: An anonymous comment on Reddit reports, “There are a few distinct groups working on this animal so I assume we’ll see some new interpretations and ideas about what it might be in the next few years. There’s a bit more anatomy than Conway-Morris described, and some of his anatomy might be totally misinterpreted.”

Confirmation is coming.

Finned swimmers are also known among molluscs, nemerteans and chaetognaths, but Typhloesus is no more like any of these than it is like a chordate. Conway Morris felt himself compelled to declare the affinities of Typhloesus completely unknown.”

He gave up. Somethimes that’s what they did back then.
That leaves more for the rest of us to discover.

“Personally, though, I can’t help wondering if the ‘alien goldfish’ might not be so alien after all: maybe it is a chordate. The overall similarities of Typhloesus to a chordate are remarkable; in particular, the hooded mouth is very similar to that of a lancelet.”

That’s the spirit! Keep going!

“But what about that missing anus, you say? Where is that all-important butthole? To which I respond, is it really missing? Looking at the figures of Typhloesus fossils in Conway Morris (1990) (which is of course a poor competitor to Conway Morris’ ability to look directly at the fossils themselves), I see that directly below the midgut is the ferrodiscus. And directly below that is a streak running between the ferrodiscus and the animal’s venter. Conway Morris saw this structure (which he called the ‘midventral strand’) as some sort of connection between the ferrodiscus and the exterior, but could it in fact be the tail-end of the reargut?”

It is not. There is no body wall opening there.

“It is certainly not unknown for the anus in chordates to not be right at the very rear of the animal; in some fish (such as the scorpionfish-like Aploactinidae) it is even moved so far forward as to be almost underneath the head.”

True. In lancelets the post-anal tail is just like the one in fish. But when the tail is no longer necessary, as in sea cucumbers and Typhloesus, the tail disappears, and with it, the anus returns to the posterior tip.

“And the missing notochord? Considering that despite the presence of specimens numbering in the thousands, a notochord was only announced in Tullimonstrum within the past year, maybe on that front Typhloesus could reward a second look.”

Turns out the notochord is not missing. A second look (Fig. 1) is always warranted. This post by Christopher Taylor goes back to 2016. Unfortunately he did not follow up on his own suggestion over the course of the last five years.


Figure 4. Close-up of Typhloesus from Conway Morris 1990.

At this point in the writing of this blogpost
I became aware of Melton and Scott 1972 (1973), who wrote in their abstract:
“Eight specimens representing four complete conodont animals have been recovered from Carboniferous sediments from central Montana. These specimens have conodont assemblages identified as Lochriea and Scottognathus. The conodont animal is 70 mm long and is bilaterally symmetrical from anterior to posterior. The anterior portion contains an oral opening, a dorsal nerve cord, and notostyle. The median section contains a gut (called the deltaenteron) and circulatory system (the ferrodiscus). The conodont assemblage is in the deltaenteron and is believed to have functioned as a food-filtering system. The posterior portion is marked by an anteroventral anal pore. Also, a rudder-fin is attached to the dorsal-posterior edge. The membrane covering the body was reticulate.”

The term ‘notostyle’ is only used in this paper. It refers to the notochord. So I am not the first to observe a notochord and dorsal nerve chord. That’s good. That the ferrodiscus is related to the circulatory system makes me think of the heart, liver and spleen in addition to a trash sac just for iron. The liver is prominent in lancelets and located in the same place (Fig. 5). As Conway Morris reported in 1990, the conodont teeth are ingested, not a part of the feeding apparatus of Typhloesus. The purported ‘anteroventral anal pore‘ is the atrium, not the anus. The rudder-like posterior fin is a sea-cucumber-like cloacal structure (Fig. 3). The anus of Typhloesus is below this (Fig. 4). ‘Reticulate‘ skin is also found in sea cucumbers.

“The conodont animal was a free-swimming form which went in search of its food.”

Cononodont animals: yes. Typhloesus: no. See above.

“The concept of assemblages, where more than one form-genus of conodont occurs in the body of one animal, is correct.”

Probably not. See above.

“The presence of a postanal portion of the body, a rudder-fin, a dorsal nerve cord, a notostyle, a bilaterally symmetrical body, and a cell structure capable of producing calcium phosphate lead us to believe that the animal bearing conodonts was in the direct line as an ancestor to the vertebrates.”

Given the new interpretations presented above, Typhloesus likely took the first left turn, toward sea cucumber echinoderms. Other taxa led more directly to vertebrates.

Figure 2. Extant lancelet (genus: Amphioxus) in cross section and lateral view. The gill basket nearly fills an atrium, which intakes water + food, sends the food into the intestine and expels the rest of the water. ” data-image-caption=”

Figure 2. Extant lancelet (genus: Amphioxus) in cross section and lateral view. The gill basket nearly fills an atrium, which intakes water + food, sends the food into the intestine and expels the rest of the water.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2019/11/lancelet588.jpg?w=300″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2019/11/lancelet588.jpg?w=584″ src=”https://pterosaurheresies.files.wordpress.com/2019/11/lancelet588.jpg” alt=”" class=”wp-image-40744″ />

Figure 5. Extant lancelet (genus: Amphioxus = Branchiostoma) in cross section and lateral view. The gill basket nearly fills an atrium, which intakes water + food, sends the food into the intestine and expels the rest of the water.

This appears to be a novel hypothesis of interrelationships.
If not, please provide a prior citation so I can promote it here.

Video of a pearlfish emerging from the anus of a sea cucumber. Click to play.

PS
Nowadays, sea cucumbers can be temporary homes for pearlfish, the relative size and elongate shape of extinct conodonts. They enter and exit through ‘the back door.’ If analogous, maybe a tiny conodont was living inside Typhloesus when it was fossilized, rather than the conodont being a prey item. With sea cucumbers, perhaps uniquely, this possibility must be considered.

References
Conway Morris S 1990. Typhloesus wellsi (Melton and Scott 1973), a bizarre metazoan from the Carboniferous of Montana USA. Philosophical Transactions of the Royal Society of London Series B 327: 595–624.
Melton W and Scott HW 1972 (1973). Conodont-bearing animals from the Bear Gulch Limestone, Montana. Book chapter in GSA Special Papers: Conodont Paleozoology by Rhodes FHT. https://doi.org/10.1130/SPE141

Typhloesus: The ‘Alien Goldfish’ of Bear Gulch by Christopher Taylor at 6/20/2016.

Figure x. Still from Fossil Friday, U of Montana fossil storage. This is fossil librarian, Kallie Moore, also of PBS Eons, holding Tylophoneus upside-down. ” data-image-caption=”

Figure x. Still from Fossil Friday, U of Montana fossil storage. This is fossil librarian, Kallie Moore, also of PBS Eons, holding Tylophoneus upside-down.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=300″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=584″ src=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=584″ alt=”Figure x. Still from Fossil Friday, U of Montana fossil storage. This is fossil librarian, Kallie Moore, also of PBS Eons, holding Tylophoneus upside-down. ” class=”wp-image-62197″ srcset=”https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=584 584w, https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=150 150w, https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg?w=300 300w, https://pterosaurheresies.files.wordpress.com/2021/11/typhloesus-montana588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 2. Click to play video. Still from Let’s Tour: U U of Montana Paleontology. This is fossil librarian, Kallie Moore, also of PBS Eons, holding Tylophoneus upside-down. This video sparked today’s blogpost.



Source: https://pterosaurheresies.wordpress.com/2021/11/16/enigmatic-typhloesus-wellsi-now-transitional-between-lancelets-and-sea-cucumbers/


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