My absolute gratitude to Happy Hen Toys for generously sending this little figure of a cute fish for me to review here. You can pick up this figure, and many others like it from there as well.
While I adore a number of prehistoric fish (like that silly Sacabambaspis that became a meme a few years ago), there is but one that has captivated me more then any other since my childhood. This is none other then Dunkleosteus, an ancient predator of the Devonian seas. While Otodus megalodon is probably the most famous prehistoric fish in the world, and Dimetrodon might be the most famous Paleozoic animal, Dunkleosteus is easily the second most famous prehistoric fish, and Paleozoic animal. Or perhaps more specifically Dunkleosteus terrelli, the largest species (and taken a step further, specimen CMNH 5768 is perhaps the most well known. CMNH by the way stands for Cleveland Museum of Natural History). From casts of its various skulls (and other remains) held in museums around the world, numerous books on prehistoric life, or media like documentaries (BBC’s 2003 Sea Monsters, etc), video games (Frontier’s 2021 Jurassic World Evolution 2, Studio Wildcard’s 2015 Ark: Survival Evolved, etc), and even movies like the recent Jurassic World Rebirth in the form of a cameo, this ancient predator continues to fascinate those who enjoy prehistoric life. My first introduction to Dunkleosteus was however in the 1996 book “Life Starts in the Sea“, followed by Pennsylvania Fishes, a guide to fish within the state, which I was gifted a copy of when winning first place in PA’s farm show aquaria contest sometime back in the mid 2000’s. In addition to all those, there’s also the toy market as well. Nearly every prehistoric toy company in recent years has produced a Dunkleosteus, or at least a good number of them (see the reviews on the blog here for a portion of them.
Back in 2021 PNSO joined those when they released a large variety of marine taxa spanning from the Devonian to the Miocene, including Dunkleosteus. They’ve given it the name Zaha. I won’t pretend that I know what the name means, or its origins, but a quick web search suggests it is of Arabic origin, typically for a girl, and means “brilliant”, “shiny”, etc. I think that would be fitting then with the iridescent coloration of the model, assuming it is correct anyways. It’s number 47 in the Prehistoric Animal Models series, and like other models in this series includes a booklet with images of the figure, a skeletal reconstruction, and a little story based on the animal. It also includes a fantastic poster of the Dunkleosteus breaching out of the water with a shark or the like. It might be in reference to potential stomach contents of a specimen of Dunkleosteus terreli (CMNH 8735), in the form of teeth from a chondrichthyan (the group of cartilaginous fishes that sharks, and rays belong to), potentially from the genus Orodus (Carr and Jackson, 2008, and Ferrón et al, 2017), or artwork of Dunkleosteus depicted hunting Cladoselache as seen in this paper (Carr, 2010). Before addressing the model itself though, let’s delve into the history on this ancient animal.
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The genus Dunkleosteus, like many fossil animals found prior to the advances in modern paleontology, was a wastebasket taxon (and potentially still is). In short, much fossil material had been assigned to the genus early on (if not much later as well), which might not actually belong to this genus. This also carries over into the genus Dinichthys that Dunkleosteus terreli was originally assigned to. It was originally named by John Strong Newberry as “Dinichthys terreli” in a footnote in the 1873 “Geological Survey of Ohio. Volume I. Part II. Paleontology.” on page 322. The genus Dinichthys itself was also only named a few years earlier in 1868, in Newberry’s paper “On some remarkable fossil fishes discovered by Rev. H. Herzer in the black shale (Devonian) at Delaware, Ohio”, published in the Proceedings of the American Association for the Advancement of Science. “Dinichthys terreli” was named after Jay Terrell who found well preserved material belonging to it in Sheffield, Ohio, and took great care with it. On the prior page, it is written in another footnote that some of this material was destroyed in a fire that happened the same year. The material was at least photographed before its destruction, as seen on plate 32 and 33 (material in plate 34 was also attributed to this species, but only those in 32 and 33 were mentioned to be destroyed). Though it wasn’t until a mandible was discovered by a “Prof. A. A. Wright” that shed light on the differences between the Dinichthys material obtained in Delaware, and from that obtained in Sheffield, Ohio, and what ultimately led to Newberry creating the new species. The newly named species wasn’t described in further detail until the 1875 “Geological Survey of Ohio. Volume II. Part II. Paleontology.“, along with more material discovered by Jay Terrell since.
There were a number of publications following in the years since (such as Branson, 1908, Heintz, 1932, etc), but it wouldn’t be until 1956 when the genus name “Dunkleosteus” would be created and assigned, nearly a century after “Dinichthys terreli” was first named prior. In the article “Les arthrodires du Dévonien supérieur du Tafilalet (Sud Marocain)” published in the French “Notes et Mémoires. Service Géologique du Maroc. 129”, it was named by Jean-Pierre Lehman to honor paleontologist David Hosbrook Dunkle, who at the time was curator of vertebrate paleontology in the Cleveland Museum of Natural History. He also researched and published on fossil fish quite a bit. The name “Dunkleosteus” is thus a combination of his last name, and the Greek ὀστέον/ostéon meaning “Dunkle’s bone”. Nearly 60 years later after the genus Dunkleosteus was named, it was also determined that Dinichthys was not closely related to Dunkleosteus either (Carr and Hlavin, 2010). So with “Dunkleosteus terreli“, the name honors not one, but two men that have made significant contributions to paleoichthyology.
As mentioned earlier, the genus Dunkleosteus terreli was from the Late Devonian period in what is now the Cleveland Shale, in Ohio (and potentially elsewhere). It specifically lived during the Famennian, the final stage of the Devonian period (if not earlier during the Frasnian stage in other locations). Historically it was considered to belong to the Arthrodira, and in turn Placodermi, and that’s how I’ll be treating Dunkleosteus for the purposes of this review. Placoderms were jawed fishes, with the front halves covered in bony armor, and the latter half made of cartilage (mostly), and arthrodires were just one branch of those (with many different sub branches, and so on). The Cleveland Shale was home to not only Dunkleosteus terreli, but a number of other placoderms, such as Bungartius, Heintzichthys, and the large filter feeding Titanichthys, in addition to many chondrichthyans like Cladoselache, Ctenacanthus, Stethacanthus, and Orodus as mentioned earlier. There were also a few bony fishes found (Carr and Jackson, 2008, and Carr, 2010). In addition to potential predation on Orodus, there appears to be damage on both a specimen of Dunkleosteus terreli and Titanichthys (per Anderson and Westneat’s 2009 paper).
Despite its popularity, the bulk of the fossil material we have for Dunkleosteus terreli is of its bony armor, consisting of the skull, and some of the thoracic parts of its body. However in addition, there’s also parts of its pectoral (shoulder) fin (Carr, Lelièvre, Jackson, 2010), part of the vertebral column (from a subadult; Johanson et al, 2019), and even some pelvic material (Engelman, 2024, linked below). The rest of its body consisted of a cartilaginous skeleton, which can be inferred from relatives with soft tissue preservation such as Coccosteus cuspidatus, Incisoscutum ritchiei, or the recently named Amazichthys trinajsticae (Jobbins et al, 2022), among others. Most reconstructions of Dunkleosteus in particular were based on the earlier reconstructions of Coccosteus cuspidatus (such as seen in Miles and Westoll, 1968), which were depicted with an eel-like (in the casual sense) tail fin, including estimates for its general body size (at times). This goes for many of the toys produced of it, and media appearances as well. Published in early 2023 was a paper by Russell Engelman that revolutionized our understanding (at least for the time being) of the Dunkleosteus terrelli body size based on numerous factors including its fossil remains, remains of relatives, and the new metric he created called the orbit-opercular length (using an insanely large dataset, with over 3000 specimens, and over 900 species). To quote from the paper, the orbit-opercular length is described as being “the length from the anterior margin of the orbit to the posterior margin of the head”. There’s a more detailed breakdown later on in the paper describing it, but I urge folks to check it out for themselves. As being one of the most popular prehistoric fish out there, and with most prior estimates having it range from over 4 to even 10 meters in length (Anderson and Westneat, 2009), it was initially met with skepticism at times from folks in the paleo community (including from myself), as the infamous “Dunk”, thought to be one of the earliest vertebrate apex predators, was now shrunk. Russel’s paper gives an estimate ranging from 3.1 to 3.5 for CMNH 5768, up to 4.1 for the largest specimen CMNH 5936 (which only consists of part of the lower jaw armor, or more specifically a left inferognathal).
Not stopping there, Engelman released another paper in late 2023 following up on the earlier 2023 size paper, as well as an early 2024 paper delving into creating an in-life reconstruction of this ancient animal (internal anatomy, and ontogeny/life stages as well). There’s also yet another 2023 paper from Engelman, and it is mostly in regards to the method used to estimate Dunkleosteus body size in the 2017 paper by Ferrón et al. The last paper focused on giving us any reconstruction of sorts prior to Engelman’s work was the 2017 paper by Ferrón et al. (as far as I’m aware) which gave support to Dunkleosteus terreli being a pelagic predator, likely having a fluked tail like seen in extant sharks. This contrasts with the earlier reconstructions of the eel-like tail based on Coccosteus cuspidatus (Miles and Westoll, 1968), which is also now outdated thanks to Engelman’s 2024 paper at the very least, which depicts it with a fluked tail based on specimen ROM VP 52664 (if not others). The short of it is that the PNSO figure is too long, although it was clearly sculpted before these papers dropped. I don’t fault it for that, but I did kind of suspect way back in 2021 that the figure was too long (if in part from the position of the dorsal fin, a bit more of my prior thoughts here as well). In fact the PNSO making of Zaha video even mentions the older 6 meter plus size estimate. The figure itself is roughly about 9 inches/22.86 cm long (which is nearly PNSO’s advertised length of 23 cm). Using my rough measurement of the figure, with the 6.88 meter total body length estimate of CMNH 5768 from Ferrón et al’s 2017 paper with the figure in an online scale calculator puts it around 1:30 scale roughly. With their measurement of CMNH 6090 at 5.34 meters, gives it an approximate scale of 1:24 (there’s a reason this one was included, see the anatomy of the head section of the review below). With their estimate of the largest specimen CMNH 5936 at 8.79 meters, gives a scale of 1:38. Using it with the 10 meter estimate (Anderson and Westneat, 2009) would put it around 1:44 scale. Using a similar method of measuring the head as Engelman did for the initial 2023 paper (see this post), gives the model a length of about 1.25 inches/3.175 cm give or take (due to using a stiff metal ruler). Comparing it to Engelman’s updated head length of 61.3 cm for the adult CMNH 5768 gives a rough scale of 1:19 (see supplementary file 1 of that 2023 paper). Comparing it to the 51 cm head length of CMNH 6090, gives it a scale of about 1:16. The width of the jaw (when open), is about 1.375 inches/3.49 cm, including the edges of the suborbital plates. The height of the tail fin is about 3 inches/7.62 cm, while the height of the dorsal armor is about 1.625 inches/4.13 cm give or take. The width of the figure including its pectoral fins is about 3 inches/7.62 cm, while the width of the thoracic armor, or the upper torso is about 1.5 inches/3.81 cm. There’s far more to address here regarding the anatomy, but I’m only just getting started.
Starting with the head for the anatomy of the figure, as far as I can tell, all major plates are accounted for. As for the thoracic armor, it is hard for me to tell on the sculpt if everything is accounted for, with it intended to be covered in skin, but the major parts appear to be present. The ventral shield should probably be more curved, and appear slightly deeper, as it instead appears to be flat, based on older reconstructions. This is in part due to the nature of how the various Cleveland Shale specimens fossilized, but other specimens found elsewhere (such as CMNH 9951) do preserve the curvature of the ventral shield. As far as I can tell, the shape of the suborbital plates appears to be like CMNH 6090 the most, give or take, maybe even a bit like CMNH 7424 in regards to the roundness of them (more so on the right side from what I can tell). This is comparing it both to the 2D reconstructions from Engelman’s 2024 paper, and with 3D models of the material (such as those on sketchfab). Worth noting that CMNH 6090 is a subadult specimen, and not fully grown like CMNH 5768, and CMNH 7424 is a juvenile, younger then CMNH 6090. Kind of peculiar they didn’t just go with the shape from CMNH 5768 in this regard, considering how common the mount of it is in museums and the like, but eh. This is why I included the earlier scale estimates with it. Granted the skull could have been based on another specimen entirely, and not any of these. The eyes appear to be small in proportion to the head’s armor, similar to the subadult CMNH 6090 and adult CMNH 5768, but do appear to be as large the eye’s orbit (as far as I can tell). The 2024 paper mentions that the sclerotic ring found within the eyes would have nearly filled the orbit, and the rings in the mounted CMNH 5768 use plaster replicas, although the rings were found preserved on the head shield. The eyes on the figure additionally do not have the bony sclerotic ring visible. While many of us on the corresponding forum here have been in favor of this for years now, the 2024 paper does open the doors to them being visible, if at least in a subtle manner. Furthermore the schlerotic rings were not flat, and did have a bulge to them. There is at least one pair of nostrils sculpted on the figure, and while the 2024 paper does mention there should be a secondary pair like on most other jawed fishes (which I’ve only just learned of when writing this review), I’m not sure if that pair would have been too small to include due to the size of the figure, or if it is included, and I’m just not picking up on it.
Now for perhaps the most exciting aspect of the figure, which is the jaw articulation. PNSO has pulled off some incredible engineering to get this to work. Most other figures of Dunkleosteus only have an articulated lower jaw like in theropod dinosaur figures, etc. As far as I know this PNSO figure is still the only one to have done this so far. When the lower jaw is opened, the upper jaw articulates with it, or vice versa. This appears to be based on work from Anderson and Westneat (2006, and 2009) that tests a four-bar linkage system hypothesis (along with estimating bite force) that Dunkleosteus may have used to open its jaws. The 2009 paper mentions that 45 degrees is the maximum the jaws can open, and I think that applies to the figure as well, more or less. As far as the placement of the gills goes, it looks like they may be correct per the 2024 paper, as they are described to be located between the head and thoracic armor in Dunkleosteus terreli, in addition to arthrodires typically having them located under the bony plates of the cheek and jaws. They also only become visible when the mouth is open on the figure, another aspect the paper mentions. Though perhaps they could extend a bit more near the lower jaw (when the mouth is open). In PNSO’s video, it’s mentioned that the mouth parts were not given extensive extraoral tissue (aka “lips”). While speculative, I do like how the main body coloration fades in on the more bone like coloration of the gnathal plates.
Moving on to the body itself, aside from overall body length being too long as addressed earlier, the body should not be so curved, particularly near the anterior, or front half, of the animal. While it does give the figure some life to it, it would have been implausible based on what we have regarding its spine. The portion of vertebrae preserved in specimen CMNH 50322 (per Johanson et al, 2019) was largely preserved in articulation (kept together like when the animal was still alive), and would have been located within the trunk (or between the dorsal and ventral shields). There was 19 vertebrae preserved, with 18 in articulation. The first 16 of these was preserved in a structure called the synarcual, and were fused together, and would have hindered it from being flexible in this region of the body. One can only wonder if the figure was in production or designed prior to this paper being released, or if it was missed. The skin of the figure largely appears to be smooth, with no obvious scales, which seems to be correct more or less, judging from Engelman’s 2024 paper, which explains most of its relatives had similar skin (and referencing a number of papers).
The shape of the dorsal fin is an interesting one. The PNSO making of Zaha video mentions Dunkleosteus likely having a triangular shark-like fin, but they opted for something shorter along the lines of a swordfish or sailfish instead, while still keeping a similar shape. Engelman’s 2023 and 2024 reconstructions both go the shark-like route however. The video doesn’t go into much detail about why they placed the dorsal fin so far back, other then the previous estimated length they used. The 2024 paper has it placed closer to the dorsal shield (but not directly over it), based upon material from both Amazichthys and Heintzichthys, in addition to Dunkleosteus own anatomy. The same paper explains that there is no evidence for actinopterygian-like (bony fishes) fin rays or spines in placoderms, or in arthrodires more specifically, so the shark-like dorsal fin would have been preferred, as it would have been more likely then what was chosen for the figure.
Zhao Chuang explains in the making of video that the pectoral fins were designed to be shark-like, and they do mention the research supporting it (likely referring to the 2010 by paper by Carr et al. describing the internal structure of the fins, or ceratotrichia, such as in CMC VP8294 and CMNH 8982, along with a soft tissue outline with specimen CMNH 8982), so this is a plus. The position of the pectoral fins seems to be more or less correct, as far as I can tell. There are tiny striations on the edges of the pectoral fins. Don’t know if sharks have these or not, or if it would have been on Dunkleosteus itself, but it gives some subtle texture to the area. The pectoral fins are also large in size, which seems to be supported in both Carr et al’s 2010 paper, and Engelman’s 2024 paper (at least to my understanding of it), though maybe they could be a tiny bit bigger too.
Focusing on the underside of the figure now, there is a very tiny cloaca present on the model, but it might be placed just a tad too far back, judging from a general arthrodire reconstruction based upon material from numerous specimens of both Incisoscutum ritchiei, and Compagopiscis croucheri in a recent paper describing well preserved internal organ material (Trinajstic et al, 2022). Additionally Incisoscutum ritchiei also preserves reproductive anatomy for both the female and male sexes, showing they had anatomy similar to chondrichthyans, with the females being viviparous (able to give live birth; Long et al, 2009), and the males having pelvic claspers (Ahlberg et al, 2009). As Zaha was intended to be female (according to PNSO’s video), the figure lacks the pelvic claspers. The shape of the figure’s pelvic fins were designed to be shark-like in response to this research. This seems to be in line with the Engelman’s later reconstructions (including the limited pelvic fin material), but they have been placed much farther back from the ventral armor. Instead they should be much closer, as many arthrodires have been preserved with evidence of such (see the pelvic fin section of Engelman’s 2024 paper if you want the specifics). The anal fin was mentioned to be designed based after a relative of Dunkleosteus with a square like shape, but they don’t specify which.
The tail, has been given a large fluked shape to it, similar to a shark, supported both by the 2017 paper by Ferrón et al. and Engelman’s work, as mentioned earlier, due to its pelagic lifestyle. However there does seem to be actinopterygian-like fin rays on the tail, much like with the dorsal fin, which is unlikely. Moving on to the coloration, Zhao Chuang explains in the video that it was based on the tiger shark, in addition to a close relative of Dunkleosteus. That relative is described as a bottom dweller that had preserved a silver-like coloration on the belly, and a red color on the top. A quick internet search shows they likely meant Groenlandaspis antarctica (which appears to have been published in a 1997 Science article called Coloring Fossils), another arthrodire placoderm. It is also worth noting the whole figure is covered in a gorgeous metallic iridescence of sorts. The gills and interior of the mouth are colored in pink, and the eyes are a silver color, while the mouth plates have a sort of bone color to them (as mentioned earlier). Without doing more digging into it, I don’t know how plausible it would be for this animal, but it is certainly pretty to look at.
Overall, despite the anatomical issues present, some of which that can’t be faulted from the papers that came after it, it is still a lovely figure and one I enjoy having, especially in hand now. Between the eye catching metallic sheen of the figure, and exciting jaw articulation, which has been a blast for me to toy around with, it’s grown on me more then it did at its launch. The head when viewed from the front (especially with the mouth wide open) is also just adorable. That said I wouldn’t be entirely opposed to PNSO redoing this one eventually, if they were to have any interest in doing so. I’ll also be looking forward to future models of it from other companies. I’d also like to see PNSO do some of its contemporaries like Cladoselache and Titanichthys someday, if not others (aside from Kaiyodo and Safari who did make figures of the former). In the meantime though, this one will be another fine addition to my collection from them. Lastly, if I’ve missed or misunderstood something, please feel free to comment and correct me. This took over a week to write (and mostly late at night, past 12am), a bunch of research, and it is now my longest review to date (exceeding 4,100 words, captions included).
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