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Sunday, 24 September 2017

The horns of Arsinoitherium: covered in skin or augmented with keratin sheaths?

1.5 Arsinoitherium zitteli trotting about Eocene Egypt, looking a bit like they could be advertising farm products. But what's with those more elaborate than usual horns?
The horns of the giant, Egyptian, Oligocene afrotherian Arsinoitherium zitteli are probably a key factor in its status as one of the better known fossil mammals. Though perhaps not quite as popular as mammoths or sabre-toothed cats, this 3 m long, four-horned species has enough osteological charisma to warrant display in many museums as well as starring roles in books and films (including, cinema fans, narrowly missing out on an appearance in the 1933 King Kong). And unlike a fossil rhinocerotid (to which it is not at all related), Arsinoitherium doesn't need us to imagine the shape of its ornament in life: two enormous horns project over the end of the snout and another pair of smaller, sub-vertical horns grew above the eyes.

Recently, I painted a portrait of Arsinoitherium for an upcoming book project and, based on my understanding of epidermal osteological correlates, I threw a keratinous sheath over the entire horn set (below). This is not a typical reconstruction - Arsinoitherium has been reconstructed with 'regular' mammalian skin (perhaps better termed 'villose skin' - Hieronymus et al. 2009) on its horns for decades but, as we all know, popularity and longevity don't always equal 'credibility' when it comes to fossil animal reconstructions.

Arsinoitherium zitteli, sporting antelope-like horn sheaths.
Shortly after this image was shared online, Darren Naish, he of Tetrapod Zoology (and the upcoming TetZooCon meeting, which you should definitely attend if you're in the UK and reading this article), had a question: had I checked horns without keratinous sheaths, like deer antlers or giraffe ossicones? It turns out that these are the more typical artistic analogues for Arsinoitherium horns, and their reconstruction without a keratinous sheath reflects this interpretation. It wasn't a question I could easily answer because I'd zeroed in on a keratinous sheath quickly in my research for the image and, in a major palaeoart faux pas, hadn't given due consideration to other options. Simultaneously, neither of us could argue for any model of Arsinoitherium horn coverage confidently because no-one has looked into this in any detail. There are some ideas in the literature, but they are fleeting and conflicting (keratin sheaths - Anonymous 1903; Andrews 1906; Osborn 1907; or skin, Prothero and Schoch 2002; Rose 2006).

It's difficult to turn away a good palaeobiological mystery, and because I like to make sure my work is as credible as it can be, I followed this question up with more research. I reasoned that the structure, development and surface texture of the three major types of mammalian headgear - horns, ossicones and antlers - could be compared to Arsinoitherium horns to see which, if any, is the best match and indicator of life appearance. Looking into this has been very informative and might be of interest to fellow palaeoartists as well as those interested in cool fossil animals, so I thought I'd share my thoughts and process here. We'll start by looking at Arsinoitherium horns themselves, then move through modern potential analogues, and finally compare them at the end to see which model seems most apt.

Arsinoitherium horns: growth, structure and surface texture

PV M 8463, the most famous of all Arsinoitherium skulls, as illustrated in Andrews (1906). Note the dotted lines across the horns - they mark the end of the preserved skull and the start of reconstructed elements.
As noted above, Arsinoitherium has two pairs of horns: a larger anterior set, which grows out of the nasal bones and over the snout, and a smaller, second pair formed from the frontal bones, above the eyes. Both sets are highly conspicuous and dominate the skull, the weight of the anterior pair presumably accounting for the development of a bony bar between the nostrils in mature animals (Andrews 1906; Court 1992). Note that the Arsinoitherium horns we're used to seeing in museums are partly reconstructed and thus of limited use as reference material. Most exhibited skulls are based on NHMUK specimen PV M 8463 (above), a 'moderately sized' adult specimen (Osborn 1907) in which neither horn is complete (Anonymous 1903; Andrews 1906). This skull was among the earliest Arsinoitherium skulls collected from Egypt but was restored rapidly once it arrived back in London. A 1903 report describes how the skull was:

"...brought home by Dr. Andrews from Egypt, and after cleaning, strengthening, and the restoration of parts deficient on the left side by modelling from the right side, is now exhibited in the central hall of the Natural History Museum in Cromwell Road."
Anonymous, 1903, p. 530

The fact that some parts of the skull were in less than stellar shape is evident from this photo of PV M 8463 (from the NHM's data portal): note the variation in colour and texture, reflecting places of reconstruction against real bone. Thus, while the familiar Arsinoitherium museum skull is a useful reference for morphology, illustrations and descriptions in technical literature will be more informative for reconstructing their integument. I've based my assessment mostly on Charles Andrews (1906) monograph, as well as that of Court (1992).

Structure. Both horn pairs of Arsinoitherium are relatively simple in gross shape and maintain the same basic morphology throughout their lives (below), though the horns of mature animals are wider, taller and more pointed than those of juveniles. The figures presented in Andrews (1906) show an increase in anterior horn base length from 41.6% in the smallest specimen to over 56% in the largest. Both horn sets are hollow, with vast internal cavities being supported by sheets of trabecular bone. In some places the exterior bone walls are surprisingly thin, only 5-10 mm (Andrews 1906).

Arsinoitherium zitteli skull ontogeny. I wonder if the horns of the largest skull should be reconstructed as longer and taller, given their arcs in the completely known skulls and gentler tapering of other nasal horn specimens (e.g. Sanders et al. 2004). Skull drawn from Andrews (1906), skull measurements by me.
Surface texture. The base of the horns are marked by deep, broad and branching neurovascular channels running from the facial region onto the horns themselves. The horn shafts are rugose on account of many deep pits, grooves and branching channels aligned along their long axes (Andrews 1906; Sanders et al. 2004). The horn tips of young animals have an especially spongy texture at the tip, presumably reflecting growth of the horn core (Andrews 1906). These textures are not typical of the rest of the skull, which are of a more typical, smooth mammalian variety even in regions where skin was probably in close proximity to the bone (e.g. the zygomatic arch, over the braincase). This is an important distinction, implying that a different epidermal configuration - different skin types, in other words - was present on the horns compared to the rest of the skull.

Having learned something of the Arsinoitherium condition, let's take a look at how modern horns, antlers and ossicones compare...

Analogue 1. Bovid-style horns (keratinous sheaths over a bone core)

Bovid horns typify a widely used approach to cranial ornamentation and weaponry across Tetrapoda. They are perhaps the simplest approach to producing a sturdy cranial projection, being little more than a bony horn core covered in a hard keratinous sheath and are permanent feature in almost animals that bear them. The one exception is the pronghorn, which sheds its horn sheath annually (it also isn't a bovid). Biology, eh: can't we have one rule without an exception?

Bovid (bighorn sheep, Ovis canadensis) horn anatomy. From Drake et al. 2016.
Structure. Bovid-style horns are composed of a hollow bony core lined with trabeculae that strengthens an otherwise thin-walled structure (Drake et al. 2016). The bone portion only occupies the basal portion of the horn, anchoring ever-growing bands of keratin that grow from the bone-keratin interface, not at the horn tip (below). This means that the tip of the horn sheath is the oldest part of the structure and that the base of the sheath is the youngest. Because keratin sheaths are inert, dead and tough tissue, they cannot be remodelled once they are formed. This dictates that the growing bony core has to forever comply with the shape of the horn sheath and cannot change shape much over time. Size changes can be accommodated as wider and longer sheath layers can cover expanding horn cores, but it is not possible to form a more complex shape - say a branch or spur - at the tip of the horn. And before anyone mentions pronghorns: their horn branches are entirely soft-tissue: the bony core retains a simple shape.

Schematic bovid horn growth, adapted from an illustration in Goss (2012).
Surface texture. Deep, oblique foramina and branching neurovascular canals characterise the surface texture of bovid horn cores. This rugosity profile is most pronounced in younger animals, but is maintained to a lesser extent in adults - in many bovids, the horns never stop growing, they just slow down a great deal. This texture is not unique to horns but accompanies many structures with keratinous sheathing, including claws and beaks (e.g. Heironymus et al. 2009). A sharp lip and particularly deep rugosity can mark the transition from horn to facial skin.

Analogue 2. Giraffe ossicones (skin over ossified dermis)

Giraffes have awesome skulls with two - and often more - ossicones that are covered in the same skin as the rest of their faces (Davis 2011). Their approach to cranial ornamentation seems unique to giraffes and their fossil relatives but might be an apt model for aberrant extinct forms, so is worth reviewing here. Clive Spinage (1968) provides an excellent overview of ossicone structure and development: the following is taken from his work.

Structure. Ossicones are low humps or columnar protuberances, continuous with the surrounding skull anatomy but formed from dermal ossifications, not outgrowths of skull bones. They eventually fuse with the skull in adult life but, unlike the underlying skull bones, ossicones are solid and very dense - they are described as having 'ivory-like' in compactness and hardness by Spinage (1968). Mature specimens show increasingly complex shapes including development of swollen tips on the frontoparietal 'horns', as well as hornlets and bosses across the major 'forehead' ossicone. Having an adaptable, living integument is essential to this process, as the ossicone covering needs to change shape to reflect the changing size and complexity of the underlying bone.

Giraffe skulls are full of sinuses, but they do not extend to their ossicones, which are extremely dense. From Spinage (1968).
Surface texture. Generally smooth with oblique foramina in juveniles and young adults, but increasingly gnarly in mature animals (more so in males). The continued ossification of dermal tissues produces a conspicuous pitting and 'flaky' rugosity profile that overgrows the surrounding skull bones and obscures the textures from earlier growth stages. In mature males, this rugosity can overgrow the entire upper surface of the skull and enhance the height and ornamentation of the ossicones considerably.

Young adult male giraffe skull by Wikimedia user Nikkimaria, CC BY-SA 3.0. Note the flaky, irregular textures of the ossicones and their complex shape: they are much more intricate and developed than those of less mature animals. There's room for more irregularity and texture on this skull, too: the skulls of old males look like they have cathedral spires growing from their faces.

Analogue 3. Deer antlers (bony projections atop cranial pedicles)

The familiarity of deer antlers allows us to forget what remarkable and unusual structures they are. Present almost universally in male deer (and in female reindeer), these elaborate, sometimes enormous structures are cast and regrown each year using a regenerative process that is the source of much anatomical and medical interest - no other mammal can regenerate such a complex appendage in this way, and the speed of the regeneration process is remarkable. Antlers are so unusual that they are only partly useful to our discussion here: we are primarily interested in antlers when they are covered in their velvet (specialised antler skin), as this is most comparable to the likely Arsinoitherium condition. Antler skin itself is interesting as, although it is continuous with the skin of the underlying pedicle, it lacks sweat glands and arrector pili (the tiny muscles that pull hair up or give us goosebumps) (Li and Suttie 2000). The antler pedicle (the permanent bony base) in contrast, is covered in the same type of skin as elsewhere on the body (Li and Suttie 2000).

A happy-looking moose (Alces alces) with his fuzzy antlers. Note the visible blood vessels on the underside of each palm. Photo by AlbertHerring, in public domain.
Structure. Both antlers and pedicles are solid, and antlers can - by virtue of growing at their tips - become more complex as they grow, developing from single spurs into networks of brows, tines and palms. As with giraffes, antler skin needs to be living and adaptable to facilitate this: a covering of inert keratin would preclude this form of growth.

More Alces antlers, this time without velvet. Note the long, branching channels. By Wikimedia user Nkansahrexford, CC-BY-4.0.

Surface texture. Antlers have variably developed rugosities consisting of conspicuous, long and branching channels impressed into smooth bone or around prominences and tubercles. These grooves are the impressions of blood and nervous networks that facilitated rapid antler growth. These textures are easily discerned even from a distance, and thus contrast with the texture of the pedicles, which are smoother and lined with relatively shallow, narrow and long impressions of vascular networks. It is unusual for hairy skin to leave such a significant osteological scar on underlying bone: typically, this form of epidermis leaves little to no remnant on skull bones (Hieronymus et al. 2009).

Arsinoitherium vs the analogues

Having looked at three major types of cranial projection in living animals, which - if any - best match the condition in Arsinoitherium? Giraffe ossicones are incomparable to Arsinoitherium horns in several aspects, perhaps the most significant being their increasing complexity and development of flaking bone textures in later life. Furthermore, the development of giraffe ossicones from bony growths in dermal tissues suggests a fundamentally different relationship between skull and dermis than of Arsinoitherium, where the bony horn component represents skull bones alone. There's enough differences here to question whether giraffe ossicones are a good model for the life appearance of Arsinoitherium horns.

In being formed of polished, deeply vascularised bone, deer antlers are closer approximations of Arsinoitherium horns. However, there is so much weirdness associated with deer antler formation and tissues that they almost remove themselves from meaningful comparison to permanent skull horn cores. The fact that antler velvet, as hairy skin, is (to my knowledge) unique in leaving deep vascular channel impressions is a major issue here, implying that either antler bone is unusually susceptible to neurovascular imprinting (do they grow so fast that they grow around their blood vessels?) or that velvet is better at altering bone textures than other skin types. Both scenarios point to antlers having some endemic oddness about them, which complicates their use as a model for life appearance of non-antlered species.

All is not lost with the cervid data, however: antler pedicles are comparable to Arsinoitherium horns in being permanent outgrowths of bone, and they also have neurovascular impressions. However, these shallow grooves compare poorly to the deeper channels and pitting of Arsinoitherium horns. Indeed, there is little about antler pedicle texture to distinguish them from the surrounding skull bones, whereas the opposite is true for Arsinoitherium.

Our comparisons improve with the bovid horn condition, which seems to chime with the Arsinoitherium skull in many regards. Both are hollow outgrowths of skull bones supported by internal trabeculae; both have bone textures characterised by deep, bifurcating neurovascular channels as well as conspicuous longitudinal grooves and oblique foramina; and both maintain the same basic shape throughout growth - excepting some basic changes in base width and horn length. Further similarities include the development of particularly deep rugosties at the base of the horn cores, which is evident in at least large Arsinoitherium skulls (Andrews 1906). This interpretation is consistent with one of the longer (but still rather short, if we're honest) interpretations of the blood vessel impressions in Arsinoitherium:

"These channels evidently lodged blood-vessels which served for the conveyance of blood to or from the covering of the horn, and judging from the marked way in which both these vessels and those on the anterior face of the horns impress the bone, it seems probable that the covering was hard and of much, the same nature as that clothing the horn-cores of the cavicorn ruminants."
C. Andrews (1906), p. 7

So...

Of the three models looked at here, it seems the basic structure and textural package of bovid-like horns best matches what we see in Arsinoitherium. Moreover, unlike the antler or ossicone models, there's no obvious mismatches with this configuration: pretty much everything we would correlate to a bovid-like horn anatomy seems present on or in the Arsinoitherium skull. The idea that a keratinous sheath might have existed in Arsinoitherium might seem odd, but it is not that outlandish given the apparent ease through which keratinous sheaths evolve. This is, after all, the tissue which has covered just about every claw, hoof, nail, horn, cranial dome and beak that has ever existed, whereas ossicones and antlers seem like specialised, clade-restricted approaches to cranial projections. The functionality of hollow Arsinoitherium horns is further reason to suspect a horn sheath. Studies of bovid horns suggest hollow cores and keratin sheaths compliment each other biomechanically, optimising the horns for for impact dissipation (Drake et al. 2016 and references therein). Stripped of a keratinous sheath, we find that hollow horn cores are great at transmitting energy but are brittle and prone to buckling and fracturing under heavy loading. It's only with a tough, fracture resistant keratin sheath that these structures can avoid breaking under heavy use so, if Arsinoitherium employed its horns for anything vaguely physically demanding, they probably needed a keratinous sheath.

It's possible, of course, that these structures were just for show, but they do look like they had a function beyond display. It occurs to me as I write this that this scene recalls the painting from Ghostbusters II. I guess we'll call this guy 'Vigo'. 
Putting all this together, I feel the case for a keratinous sheath over the Arsinoitherium horn sets is reasonable, at least so far as it can be made with publicly available data. Aspects of morphology, growth, surface texture and - perhaps - functionality seem fully consistent with a bovid-like horn configuration, whereas other potential models are less comparable. From an artistic perspective, this is exciting: horn sheaths can be extremely elaborate structures and exaggerate the size of the horn core considerably, so Arsinoitherium might have been far more extravagant in life than we have previously imagined. I've tried to hint at this with my reconstructions - remember, this animal wasn't just a funny-faced rhinoceros!

But - before we go crazy with this - do remember that the core of this analysis - the interpretation of Arsinoitherium headgear - is entirely literature based. I've not seen original specimens nor even modern, high-res imagery of an unreconstructed skull (this wasn't for lack of trying - the literature on these animals needs updating). Thus, while I've tried to be as thorough as I can with my observations, and as cautious as I can with my interpretations, I might be ignorant of some important detail. Take everything here with an appropriate pinch of salt, and please chime in below if you can provide superior insight. There's clearly scope for a more detailed study on this topic and, given how unique the horns of Arsinoitherium are, there might be some interesting functional findings to emerge from further investigation.

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References

  • Andrews, C. W. (1906). A descriptive catalogue of the Tertiary Vertebrata of the Fayum. Publ. Brit. Mus. Nat. Hist. Land. XXXVII.
  • Anonymous. (1903). A New Egyptian Mammal (Arsinoitherium) from the Fayûm. (1903). Geological Magazine, 10(12), 529-532.
  • Court, N. (1992). The skull of Arsinoitherium (Mammalia, Embrithopoda) and the higher order interrelationships of ungulates. Palaeovertebrata, 22(1), 1-43.
  • Davis, E. B., Brakora, K. A., & Lee, A. H. (2011). Evolution of ruminant headgear: a review. Proceedings of the Royal Society of London B: Biological Sciences, 278(1720), 2857-2865.
  • Drake, A., Donahue, T. L. H., Stansloski, M., Fox, K., Wheatley, B. B., & Donahue, S. W. (2016). Horn and horn core trabecular bone of bighorn sheep rams absorbs impact energy and reduces brain cavity accelerations during high impact ramming of the skull. Acta biomaterialia, 44, 41-50.
  • Goss, R. J. (2012). Deer antlers: regeneration, function and evolution. Academic Press.
  • Hieronymus, T. L., Witmer, L. M., Tanke, D. H., & Currie, P. J. (2009). The facial integument of centrosaurine ceratopsids: morphological and histological correlates of novel skin structures. The Anatomical Record, 292(9), 1370-1396.
  • Li, C., & Suttie, J. M. (2000). Histological studies of pedicle skin formation and its transformation to antler velvet in red deer (Cervus elaphus). The Anatomical Record, 260(1), 62-71.
  • Osborn, H. F. (1907). Hunting the Ancestral Elephant in the Fayûm Desert: Discoveries of the Recent African Expeditions of the American Museum of Natural History. Century Company.
  • Prothero, D. R., & Schoch, R. M. (2002). Horns, tusks, and flippers: the evolution of hoofed mammals. JHU press.
  • Rose, K. D. (2006). The beginning of the age of mammals. JHU Press.
  • Sanders, W. J., Kappelman, J., & Rasmussen, D. T. (2004). New large-bodied mammals from the late Oligocene site of Chilga, Ethiopia. Acta Palaeontologica Polonica, 49(3), 365-392.
  • Spinage, C. A. (1968). Horns and other bony structures of the skull of the giraffe, and their functional significance. African Journal of Ecology, 6(1), 53-61.

18 comments:

  1. Those Arsinoitherium reconstructions are so fresh and fantastic!
    Since the matter of Arsinoitherium horns is covered, Im now curious about how would you interpret the weird headgear of the other bizarre horned giant Uintatherium.

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    1. I'm not an expert on Uintatherium at all, and the worst thing to do here would be to make a snap suggestion based on a rapid Google image search.

      ...but, based on a rapid Google image search, the textures on the Uintatherium skull horns, bosses etc. seem less differentiated from the rest of the skull than they are in Arsinoitherium. If - and that's the sort of 'if' that should be in red, in size 72 font and underlined - that's an accurate assessment, they might just be covered in facial skin.

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    2. "Since the matter of Arsinoitherium horns is covered"

      I see what you did there.

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    3. Also makes me wonder about the nasal projections of brontotheres, specifically embolotheres.

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  2. Toys seem to be the one area where the keratinous covering is preferred to the skin covering.

    http://www.thebigtheater.com/images/products/thebigzoo/400/072830-29.jpg

    http://dinotoyblog.com/2015/10/27/arsinoitherium-collecta/

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  3. "Because keratin sheaths are inert, dead and tough tissue, they cannot be remodelled once they are formed. This dictates that the growing bony core has to forever comply with the shape of the horn sheath and cannot change shape much over time. Size changes can be accommodated as wider and longer sheath layers can cover expanding horn cores, but it is not possible to form a more complex shape - say a branch or spur - at the tip of the horn. And before anyone mentions pronghorns: their horn branches are entirely soft-tissue: the bony core retains a simple shape."

    How does this then apply to the development of the shapes and textures we do see on bovid horns, such as the furrowed or "ribbed" look (for lack of a better term), or even the bumpy projections on your Arsinotherium reconstructions?

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    1. I'm guessing the textures you mention presumably form as layers of keratin are laid down. Presumably the horizontal ribs are similar to growth rings? We could assume that those bumps in the illustration form at the base of the horn as it grows and are pushed toward the tip of the horn as the animal matures.

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    2. Yep - all those structures are produced with soft-tissue, not altering bone shape (illustrative example: http://animaldiversity.org/collections/contributors/phil_myers/horns_antlers/horn_and_core/medium.jpg).

      Keratinous sheaths grow like mollusc shells do whereby altering the material thickness (either around the entire structure or just in one region) produces all manner of shapes as the horn or shell grows. This can be deliberate or influenced by other factors. Temperate-living bovids, for instance, tend to have banding instilled by their ability to produce more horn in summer than the winter. In all likelihood, we're not representing the true diversity of shape in the keratinous horns of fossil animals.

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    3. I would love to see images of different stages of growth as the horn grows longer over time. I wonder if the horn core grows at the same rate as the keratin or if it is outpaced by the keratin and if that makes a difference in how the ends up looking. Of course whenever you use google to search for horn growth it inevitably throws in deer because people can't tell the difference between antlers and horns. However, when it happened I stumbled on "cactus bucks" in deer which are not very similar but have very striking antlers. Weird stuff going on there for sure.

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    4. Whoa! Have a look at this page about annual sheep horn growth.
      http://priscilla.saltmarshranch.com/2014/01/19/sheep-annual-horn-growth/
      There are some great images of sheep with white spotting genes and where the white is over the horns they appear translucent making the increased blood flow during growth appear pink.

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  4. There's something particularly gratifying about the bovid argument here...

    Although:

    "This is not a typical reconstruction - Arsinoitherium has been reconstructed with 'regular' mammalian skin (perhaps better termed 'villose skin' - Hieronymus et al. 2009) on its horns for decades..."

    "The idea that a keratinous sheath might have existed in Arsinoitherium might seem odd..."

    I get a strange feeling that I've been living under a palaeontological rock - or just didn't pay Arsinoitherium due attention - because I was completely unaware of the 'skin-covered horns' consensus, and don't think the keratin sheath seems odd at all. Quite the reverse.

    But that's why I like visiting this blog. Always learning something!

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    1. I'm basing this mostly on artwork, but must admit that I've not performed an exhaustive search. I'm mostly thinking of works by classic palaeoartists - maybe there's more variation out there that I realised.

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  5. One niggle. There are extinct antilocaprids (pronghorn relatives) in which the bony part of the horn is forked. Suggesting that bovid style cranial ornament isn't necessarily incompatible with complex shapes at the tip.

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    1. That's interesting - I didn't know that. But if these extinct antilocaprids used a pronghorn-like mechanic of shedding their sheath annually, that might give the horn core time to change shape when it's only covered in skin. Certainly among living species, I don't think the ability to change the shape of a horn core within a permanent keratinous sheath is known.

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  6. As good time as any to point out that embrythopods might not have been afrotheres after all.

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  7. I don't want to be dismissive of molecular concensus, but this is about a fossil taxon and there is no way to prove what is or isn't an Afrothere.

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    1. Recent studies on embrythopod biogeography and rate of expansion imply that they are laurasian, possibly related to perissodactyls as many other Paleogene ungulate groups.

      Remember, paenungulates and prissodactyls converged closely on many anatomical details.

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