Saturday, 27 December 2014

Cats and humans: the cultural evolution through history

The idea for this blog post came from wandering around the archaeological museum in Istanbul, Turkey. I do no pretend to be an archaeologist or cultural historian, but I have always been fascinated by how humans interact with the animals they regularly see, and based on my postdoc research I've focussed on cats. Cats have always held a certain place in our hearts, whether it is the big wild cats or the small domesticated (sort-of) cats that run around our houses today.

Undoubtedly, ever since the first hominids there have been interactions with the felids. Most likely these would probably have been lions, leopards and probably some extinct species chasing our ancestors. However, it is possible this wasn't just early hominins getting munched by the big cats in Africa. A modern tribe in Africa (the Dorobo), are known to go and chase lions from their kills. There is obviously lots of room for debate on how successful our smaller ancestors would have been at this against predators that haven't been under pressure from humans for many millenia, but there is no reason to believe it's not a viable option.

All of this is lovely and speculative, and its not until about 30-32,000 years ago that the first evidence of humans appreciating the cats in their environment appears. The cave paintings in the decorated cave of Pont d'Arc (known as Grotte Chauvet-Pont d'Arc, Ardèche), in South France show many images of cats, particularly those of of lions (probably cave lions: Panthera leo spelaea) and one described in all the images and text I could find as a panther (which as a species doesn't actually exist, so is probably a leopard).

Selection of images from Chauvet cave (from Top left is a small spotted "panther" beneath the bear, top right is lion panel, bottom is an unknown felid, probably lion.

Undoubtedly there will be several more examples of humans showing cats off in their culture before the next major transition I'm including. We will jump forward to around 7500BC, and not long after the first permanent settlements started to occur across the Middle East and Egypt. It's about this time that humans start domesticating small cats. Derived from wild cats of North Africa, Felis silvestris libyca, it is likely that their domestication was similar to that of dogs from wolves. The presence of permanent settlements and agriculture likely attracted wild cats looking for food (either as scraps, or in the form of rodents eating the crops). Undoubtedly some of these which were more bold came into contact with humans, and through selective choosing of offspring cats were to become domesticated. However, cat domestication is by no means as advanced as dogs, with many cats even to this day being feral or semi-feral for pest control on farms. Indeed most cats can revert within one generation back to being feral, whilst many dog breeds would struggle without human intervention.

With the domestication of the cat beginning, its perhaps unsurprising that they soon entered into mythical status too. Egypt was amongst the first places to raise cats to god status. Cats in Egypt were known in ancient times as Mau (I can't find out if that's because of the meow noise they make though, although there is now a breed known as an Egyptian Mau), and the first god feline (a lion) was Mafdet. Another called Bastet started off lioness-like, but through time became associated with domestic cats. It was this goddess that lead to the mummification of cats that were included in human burials. Indeed the Egyptians worshipped cats to such a level that when one died it was mourned much as the loss of a human relative within the household. However, from 390A.D. when their worship was banned, cats fell from such revered status within Egypt but are still tolerated to present, and remain admired due to their significance in Islam (particularly for their cleanliness).

The lion appears over and over again as a symbol of gods, and thus equally royalty (who were god kings/queens). This is probably due to their large size, fearsome reputation and historical presence across all of the regions where early civilisations emerged. Think back to Hercules slaying the Nemean lion (which was in Greece). Thus statues of lions appear guarding entrance ways to cities and palaces across the ancient world.
Lion statues. Left Portal lions from the Late Hittite period (8th Century BC), Right lion from Bucaleon (Boukoleon) palace, 5th century AD Istanbul.

The Greek and Romans are probably the most responsible for bringing most of the felids of the Old World to the knowledge of the general public. The Greeks with the expansion of various empires across the world, and the Romans followed suit. The Romans went a step further by bringing animals from across the empire for various spectacles and games in the arenas (whether small amphitheatres, or the Coliseum). Because of the array of them known to the Roman world, they appear throughout their culture, whether in statues or in mosaics, or even in sarcophagi.

Selection of Roman and Greek imagery of cats. Top are mosaics, left showing cheetah and lion, right showing tiger. Bottom left statuette showing a sleeping Eros on a lion skin; bottom right is Dionysus leaning on a pillar with a panther.

Indeed the Romans and Greeks collected many animals and kept them in zoos/collections for their personal enjoyment, study, and for supplying others around the world (whether for games or celebrations). This private collection would mostly die out with the collapse of the Western Roman Empire, and the rise of Christianity limiting the number of exhibitions where wild animals were killed. However, many royal families maintained collections through the Middles Ages. Charlemagne is known to have held lions, whilst William the Conqueror is known to have held a small collection that was expanded and King Henry I is known to have held lions, leopards and lynxes. During the reign of Henry II, his son Richard, would gain such a reputation for fearlessness in battle he would go on to gain the name Richard the Lionheart. When he became King Richard I, he would also be the one that cemented the heraldry of England with three lions.

Three lion heraldry of Richard I, which would go on to evolve into the Royal Arms of England, right.

King John who would reign following Richard I death, would officially set up the royal menagerie which would go on to hold all manner of felids, and open to the public in the first zoo (of sorts). 600 years later it would be closed by the Duke of Wellington, and the animals distributed between Dublin and London Zoo. Until this day the zoo maintains several felid species, although most individuals are now located outside of the city in the larger Whipsnade zoo. Interestingly, there is currently a break from tradition in London, with the Asiatic lions being relocated temporarily whilst their enclosure is renovated and thus there are no lions being present in London for the first time in 800 years. The lions remain very much an English thing (although sometimes they were called leopards historically on the seals/heraldry), so much so that the Three Lions remains the nick-name of the football team, and the rugby team combined from the UK and Ireland is known as the British and Irish Lions.

Having focussed very much on the UK and Europe so far, its time to spread further afield. Whilst the lion has been the symbol of European royalty for millennia, it also has represented royalty in Africa, with Ethiopia probably the best example. The lion featured until 1975 on their national flag as a symbol retained from their days as an Empire. This lion is much more significant as it is the Lion of Judah and can be traced back as a symbol of the tribe of Isrealites of Judah, descended from the 4th son of Jacob in the Torah. The royal lineage was traced (supposedly) all the way from Solomon to the leaders of Ethiopia until Haile Selassie. Due to Haile Selassie being important in Rastafarianism, the lion is an important symbol to that religion too. The Lion of Judah has also been added to the official emblem of Jurasalem in 1950 with the foundation of it (or at least the western bits officially) as the capital of Israel. Whilst jumping around, but mainly in Africa, there are some interesting tribal traditions that remain. The most famous is probably the Maasai tradition where boys were supposed to kill a lion as a show of their warrior strength before becoming a man. This tradition has mainly died out as it illegal to kill lions, but there remains great prestige for those who do kill a lion in defence of livestock.

Lion of Judah featuring on the Ethiopian flag pre-1974.

So jumping from lions and Africa, to tigers in the Far East. In China, the tiger is the symbol of the monarchy much like lions in Europe. Interestingly the shape of the stripes across the tiger head look like the symbol for king (王), and therefore the tiger is known as king of the animals (interesting as lions are the king of the jungle in the West). The tiger plays continued significance as one of the 12 animals in the Chinese zodiac, and as an important figure in some martial arts. In India, the tiger is associated with both gods Shiva and Durga, whilst in Vietnam many temples are devoted to tigers. The Tamil people adopted the tiger as their symbol and namesake whilst they fought a major war for independence in Sri Lanka before being beaten a few years ago. One fact that I didn't know but particularly enjoyed whilst doing my research for this blog (particularly on wiki it must be said) is that whilst the West has myths of werewolves, the East has myths of weretigers which shift from human to animal form and back again.

In the Americas, the jaguar (as the biggest cat there) has featured in all MesoAmerican cultures. In the Olmec culture, jaguars appeared regularly as anthropomorphised, and possibly even werejaguar like. The Maya adopted the jaguar as various gods, and as with the lion in Europe, became associated with the ruling class who often wore their pelts to distinguish them, and the ruling families often incorporated the jaguar into their names. Indeed the Maya even developed a stringed musical instrument that when played makes a noise identical to a jaguar growl. The Aztecs followed the tradition of revering the jaguar, with the elite class of soldiers being Jaguar Warriors. To attain the status a soldier had to capture 12 soldiers in 2 consecutive battles (deemed more honorable than killing).

Jaguar warrior with club laced with obsidian shards, and a shield with feathers that conferred protective powers.

The Inca also have jaguars in their mythology, and at the Temple of the Jaguar in Chichen Itza as part of the coronation the kings had to walk under a frieze of many jaguars. The jaguars are not the only big cats of the Americas, with pumas (mountain lions/cougars) also present across much of North and South America. The Inca saw the puma as a good omen for wealth, and their ancestors today still believe it lucky to see one. The native tribes of North America however were divided over pumas. Those tribes on the western side thought seeing or hearing as evil and were associated with witchcraft whilst eastern tribes (e.g. the Seminoles and Shawnee) considered them noble animals and their remains were used to make powerful hunting potions.

So that roughly covers much of the older history of cats that I found. Undoubtedly I will have missed some big details and many views on cats. Whether its black cats and their bad luck, or covering humans hunting cats, and cats hunting humans (e.g. lions of Tsavo), or indeed humans using cheetahs to hunt. I feel my views on that may be very biased against humans in that respect. After all, many of the lion subspecies are endangered (or extinct); cheetahs inhabit a tiny fraction of their former habitat; leopards have had regional extinctions; tigers have had 3 subspecies go extinct and several other subspecies critically endangered (there are more tigers in Texas than there are in India!); pumas are much reduced across a lot of its former range; and the story is the same for jaguars too. And that is just the big cats, many of the smaller ones are experiencing much the same situation.

Retraction of the tiger home ranges. Historical in pale yellow, present in green. Note in the world map the presence of tigers around the Caspian and Black Seas for which no populations survive.

It would be interesting to see how each of the former cultures would react today to how we have dealt with our cat species, and indeed how we view them in modern culture (at least here in the Western world, which I know far more than for the rest of the world and for which it would be unfair of me to speak). Would they find our zoos an impressive collection of animals that inspire awe or appreciate them as conservation/reproductive tools? Would they view them as cruel enclosures for which the animals should be freed? I feel that the former is far more likely from European cultures a long way back, but maybe not for many cultures that maintained closer roots to nature who would probably view them as the latter. Would the cultures think it appropriate that many sports teams hold felid names (e.g. Jacksonville Jaguars)? How would they view our felines in the Lion King, or even Tigger in Winnie the Pooh? Would they be intrigued by how much we love and adore our cats as pets? Would they be entertained or ashamed by the amount of time we spent taking photos of our cats and putting them on the internet? How would they feel about this:

Breading. A strange craze that went around putting cats heads into a slice of bread.

Merry Christmas all, and hope you have a great 2015!

Friday, 12 December 2014

1 year on from PhD hand-in

Its been 1 year since I handed in! It was a proud moment in life, but also one of the most anticlimactic moments in it to. Explaining that to people is really difficult, but I will give it a go.

Imagine that you've spent 3 (and in my case 1/4) years working with the single minded goal of finishing a PhD. Of course you have other goals throughout that time, and even mini projects etc within the PhD, but the overarching aim is to finish the thesis. Then imagine the end being like any big project... no matter how hard you've worked in the run up to the finish, the end is always crazy. In that crazy end you sleep far less than normal and see 4am at least a few times in the last fortnight in combination to getting up as usual. And for added benefit, pack up your room to move out of where you are living in preparation to move to London after the Christmas holidays. That sums up my last month (more or less). Oh, and I forgot, being a senior resident in charge of 36 freshers all the time, with 400+ in general, and with responsibility for one of the busiest student bars out of any of the Bristol University halls of residence.

If you can imagine all of that, congratulations. Suffice to say, I was a bit of a mess by the end. However, I managed to do it all (supplied by copious quantities of Coca-Cola). My self imposed deadline for handing in was Friday 13th December (you know, for good luck and all), but I had finished everything to the level I was happy with on the 12th, which was the day I moved most of my stuff out of my flat. With that all done, I thought why wait (even though 13 is my lucky number despite not being superstitious) and did hand in on the the same day. It was almost a disaster though as typically the Exams Office wasn't open like most buildings, and wouldn't reopen until after I had left England and returned home for Christmas (and I believe a well earned break). Thankfully a lovely receptionist in the building took my bound pre-viva copies of my thesis, and let me sign away all credit (a few days later I got the confirmation so that was a relief).

I was in an exhausted delirium, but rather than happy I was just emotionally and physically drained. There was no big celebration, there was no fireworks or confetti/balloons falling down on me or banners saying well done (not that I was expecting any of that). I don't know what I was expecting, but it just felt like such an anticlimax after all that work.

So what do you do when you've done everything you've worked towards for a bit over 3 years? I was the 2nd person in my cohort to finish, so not only was I done, there was no-one really to celebrate with as they were all toiling away and a little peeved I was in work with no work to do. No work to do! Imagine that! But the result was I was finished, room packed up, office emptied, and it was midday. Thankfully I managed to go get some food with a friend, might have been BBQ pork and/or a milkshake (the usual palaeo treats). I think I went to the pub after for a beer, but to be honest you could tell me now I did this that and the other, and I wouldn't be able to tell you if I did (not through alcohol related reasons, that much I am certain). I didn't have long to linger as I was on a plane two days later and I left all my PhD thoughts in England to recharge the batteries on a beach in the Caribbean. A bit of sunshine and the break obviously did me good, as two people on my return (independently) said I looked healthy. Worrying how bad I must have looked before I left...

Hopefully I am the exception rather than the rule for people handing in their PhDs, and people found the hand in a moment of elation rather than anticlimax. To be fair, I found my viva anticlimactic too, but that's for another time. I'd love to hear how everyone else found it in the comments. Please do let me (and everyone know)!

Sunday, 7 December 2014

Ornithomimosaurs - bird mimicking dinosaurs

Ornithomimosaurs, literally bird-mimic-lizards, are a group of dinosaurs (if you hadn't guessed from the post's title) that I spent my PhD working on. They belong to the theropod lineage, the group of dinosaurs associated with meat eaters like T. rex and Velociraptor that eventually give rise to birds. Worth remembering at this point that dinosaurs are not extinct, the non-avian ones are, the avian dinosaurs (birds) are still going and remain more diverse than mammals. The ornithomimosaurs got their names when the first specimens were found back in the late 1800s around the end of the "Bone Wars" between Edward Drinker Cope and Othniel Charles Marsh. It was Marsh who got there first, describing Ornithomimus velox in 1890 based on the remains of a partial fore- and hindlimb from North America. Marsh went on to name several other species of which most are no longer considered valid and are believed to be tyrannosaurid (a common issue) or alvarezsaurid. The reason for this recurring issue is the bird-like nature of the metatarsus in many theropods which were grouped into Arctometatarsalia. This is now a defunct group as it was realised to be a convergent characteristic in theropods.

Holotype of Ornithomimus velox showing the arctometatarsalian condition of the third metatarsal being pinched between II and IV (Marsh 1890).

It was Marsh who thought that Ornithomimus were first ornithopods, until the discovery of a more complete skeleton. A new discovery by Lambe (the ornithomimosaur record is basically a who's who of palaeontologists) in 1902 lead to the naming of Ornithomimus altus which by 1916 would become Struthiomimus altus (Osborne) and the second genera in the clade. As with Ornithomimus there have been many species added into the Struthiomimus genus, although most have been grouped into S. altus. There was to be one more major addition to the Ornithomimus that would stick being Ornithomimus edmontonicus which was named in 1933 by Charles Sternberg from the famous fairly complete skeleton now found in the Canadian Museum of Nature. Between Struthiomimus and Ornithomimus there exist many complete (or as close as you can often get in palaeontology) skeletons giving a very good insight into their anatomy. There may be several additional species from North America: Dromeciomimus (which still appears in some literature, but studies suggest it is probably a junior synonym for Ornithomimus); an unnamed large ornithomimosaur which was discovered in the Royal Tyrell collections as pieces originally (and currently) within the tyrannosaur collections; and several more specimens previously described and lumped in with others that may be split back out.

In 1933, the first ornithomimosaurs from outside of North America were discovered. Archaeornithomimus (originally Ornithomimus) was described from Mongolia. It was to take many years before its reclassification as a separate genus (Russell, 1972 in fact), and until that decade no other new genera were described. The 1970s marked the reinvigoration of ornithomimosaur finds with joint Soviet expeditions to Mongolia. Deinocheirus first came to light in 1970 as a massive pair of arms (which no-one was certain what they belonged to), and with recent excavations has turned out to be one of the craziest looking dinosaurs (let alone ornithomimosaurs) found. The massive arms partner with a large body with a small sail over its pelvis, and a relatively small and very hadrosaur/duck-billed skull (Lee et al., 2014).

Deinocheirus with is massive arms, and its pelvic sail by Michael Skrepnick

Following on from Deinocheirus, Gallimimus was described in 1972 providing complete skeletal material from a range of different sized/aged animals. Following on from those taxa, the 1980s added more Mongolian taxa: Garudimimus; Harypmimus; Anserimimus.In 1990 the first European ornithomimosaur was discovered, Pelecanimimus, which wasn't only the most basal of known ornithomimosaurs, it also possessed the most teeth of any known theropod, and preserved with the remains of soft tissue including a gular (throat) pouch from which it derived its pelican-mimic name.

Map of ornithomimosaur fossil locales. Each colour represents a different taxa. The specimen in Australia is almost certainly not ornithomimosaurian, but the presence of ornithomimosaurs in South Africa suggests there should be members in the other Gondwanan continents (S. America, Australia and Antarctica). From Palaeobiology Database.

The 2000s have led to many more discoveries of ornithomimosaurs, not least driven by the opening up of many sites within China. Hexing, Shenzhousaurus, Sinornithomimus, Qiupalong, Beishanlong all quickly joined the literature, as have ornithomimosaurs from Africa (Nqwebasaurus - with the Nq actually being a click sound from the Xhosa language in South Africa and thus being one of my favourite dinosaur names) and Thailand. Other unnamed taxa have been found in Bulgaria, Japan, France and Kazakhstan and await further description/material.

Barsbold, 1976
Nqweba =Kirkwood Fm. in Xhosa
de Klerk et al., 2000
N. thwazi
Xhosa for fast funner
Choiniere et al., 2012 assigns to ornithomimosaurs
Pérez-Mereno et al.,1994
P. polyodon
Many teeth
Pérez-Mereno et al.,1994
Ancient name for China-lizard
Ji et al., 2003
S. orientalis
Of the orient
Ji et al., 2003
“Like a crane” in Chinese
Liyong et al., 2012
H. qingyi
“with slender wings” in Chinese
Liyong et al., 2012
Harpy – Greek mythology
Barsbold and Perle, 1984
H. okladnikovi
Named after AP Okladnikov
Barsbold and Perle, 1984
Being from Thai mythology
Buffetaut et al., 2009
K. khonkaenensis
From Khon Kaen
Buffetaut et al., 2009


Lee et al. 2014
White Mountain Dragon
Makovicky et al., 2010
B. grandis
Makovicky et al., 2010
Garuda (Mongolian mythology)-mimic
Barsbold, 1981
G. brevipes
Short foot
Barsbold, 1981
Osmόlska and Roniewicz, 1970
D. mirificus
Osmόlska and Roniewicz, 1970


Marsh, 1890
Marsh, 1890 (including Dromiceiomimus Russell, 1972)
O. velox
Marsh, 1890
O. edmontonicus
From Edmonton
Stemberg, 1933 (including Struthiomimus currelli Parks, 1933; S. brevitertius Parks, 1926; S. ingens Parks, 1933; S. samueli Parks, 1928)
Osborn, 1917
S. altus
Lambe, 1902 (Ornithomimus altus Lambe, 1902)
Russell, 1972
A. asiaticus
From Asia
Gilmore, 1933 (= O. asiatcus Gilmore, 1933)
Osmόlska et al., 1972
G. bullatus
From Latin bulla – ref to parasphenoid
Osmόlska et al., 1972
Kobayashi and Lü, 2003
S. dongi
Discoverer = Dong
Kobayashi and Lü, 2003
Barsbold, 1988
A. planinychus
Flat claw
Barsbold, 1988
Quipa Formation, long = dragon
Xu et al., 2011
Q. henanensis
Henan Province
Xu et al., 2011


Unnamed specimen from Alberta

Longrich, 2008
Bulgarian ornithomimid

Mateus et al., 2010
Ornithomimidae indet

Alifanov and Averianov, 2006
Ornithomimidae indet

Averianov, 2006
French ornithomimosaur

Néraudeau et al., 2012
Japanese ornithomimosaur

Hasegawa et al., 1999

Phylogeny and Phylogenetic Positions
So I've said all about ornithomimosaur discoveries, but how do they all relate?

Cuff 2014. Composite tree of Ornithomimosauria (Based on phylogenies from: Lee et al., 2013 SVP; Makovicky et al., 2004; Xu et al., 2011; Choiniere et al., 2012; Makovicky et al., 2010; Buffetaut et al., 2009. Ornithomimosauria - 1, Ornithomimidae - 2, Deinocheiridae - 3.

I based that tree on a composite of all known phylogenies up until my hand-in, and what I could remember of the Deinocheiridae from the Lee et al. 2013 SVP talk. Since then, the overall topology is roughly the same with better resolution within the Deinocheiridae:

Modified from Lee et al., 2014

The phylogeny does suggest that we have some big gaps to fill in the ornithomimosaur fossil record based on the ghost lineages (the length of time between the known appearance of taxa, and where they separate from their relatives), particularly for Harpymimus and Deinocheirus.

So that's the inter-relationships of the ornithomimosaurs, but where do they fit within the theropods as a whole?
Cuff 2014 SVP talk. Modified from Choiniere et al., 2013 and Lee et al., 2014.

The ornithomimosaurs consistently fall out close to the compsognathids (the little dinosaurs in Jurassic Park that chirp and attack the girl in the beginning of the second film) and close to the tyrannosaurs. This phylogenetic closeness, and convergence within the metatarsals (foot bones) means that the feet of small tyrannosaurs and large ornithomimosaurs look very similar and sometimes cause confusion as with the large North American ornithomimosaur. The ornithomimomsaur position in theropods is within Coelurosauria, but just below the Maniraptora. For most laypeople that means nothing, but it helps us define the position easily, and is important for studying patterns of character change (and diet which I will come back to later).

Soft tissues
So I mentioned Pelecanimimus possessed a gular pouch, but it also seems to have possessed some form of soft tissue crest too. It, however, does not seem to possess feathers (or at least they aren't preserved). This is a bit strange as it is known feathers are found throughout theropods, as well as it seems in some of the Ornithischia. It is even more strange that Ornithomimus has feathers (Zelenitsky et al., 2012). Ornithomimus feathers are interesting because they appear to change through growth (ontogenetically) with small members being fluffy, with large individuals having large pennaceous feathers. This discovery has been suggested that wings may have first developed for sexual display.

Feathered Ornithomimus reconstructions based on Zelenitsky et al. (2012) showing the ontogenetic differences in feathering. Artwork by Julius Csotoyni (a great palaeoartist - check out his other work)

Feathers had been suggested to be also sexual display within oviraptorosaurs with pygostyles, so this fits nicely (and explains the Nature paper for what otherwise is a boring story as we already knew tyrannosaurs had feathers by 2012, so expected ornithomimosaurs to have them). An interesting find on orrnithomimosaurs (or at least two specimens) is the preserved keratinous rhamphothecae (beak - although technically that is the keratin layer and bone, but I will talk beaks from now on when I mean rhamphothecae). Indeed it is these beaks (or equally lack of teeth) that has led to many discussions on what ornithomimosaurs ate.

The two ornithomimosaur specimens preserving the keratinous rhamphothecae. Top is Ornithomimus edmontonicus (RTMP 95.110.1), Middle is Gallimimus bullatus (IGM100/1133), with the bottom an enlargement of the premaxilla from the same G. bullatus specimen (Norell et al. 2001).

In ornithomimosaur crania the loss of teeth and the preservation of soft tissue remnants of rhamphothecae has drawn more specific focus (Norell et al., 2001; Barrett, 2005). The remnants of columnar structures in the rhamphotheca in a specimen of Gallimimus (GIN 100/1133) led to comparisons with lamellae in the beaks of anseriforms, particularly a filter-feeding straining species, Anas clypeata (Norell et al., 2001). A re-examination of the fossil showed that these structures are the keratinous rods vital for the structure of a rhamphotheca and found in many other taxa (Barrett, 2005). In addition, the lack of retracted nares, a small depressor mandibulae, a potentially wide gape (Barsbold and Osmólska, 1990) and a calculation of energetics required for filtering suggest this was not a possible ecological niche for ornithomimosaurs (Barrett, 2005).

Interestingly the filtering/predating on freshwater invertebrates was not a new idea (Osborn, 1917), and was one of many hypotheses for ornithomimid feeding. In addition, the following have all been suggested at some time or another: a browsing, herbivorous lifestyle (Osborn, 1917); myrmecophagous (anteater-like) habits (Osborn, 1917); omnivorous but limited to soft food – eggs, fruits, invertebrates and small vertebrates (Barsbold and Osmόlska, 1990); or predatory, but restricted to small vertebrates, insects and possibly eggs (Osmόlska et al., 1972; Russell, 1972). All of these suggestions were based upon cranial morphology, but have never been tested. Indeed, Paul (1988) noted that many of the assumptions of weak skulls with limited adductor musculature overlooked similarities to dinornithid birds (such as bracing in the quadrate and jaw joints) and ornithomimosaurs may in fact have skulls stronger than previously suggested. I spent my PhD reconstructing skulls of several taxa and carrying out functional tests upon the skulls under different feeding regimes to finally test some hypotheses.

The most widely accepted hypothesis for ornithomimosaur diets is that they were herbivorous (or at least mainly herbivorous). A series of papers by Lindsay Zanno and colleagues (Zanno et al., 2009; Zanno and Makovicky, 2011) suggested that there were a series of characters found throughout dinosaurian lineages that could be correlated with herbivory e.g. loss of teeth in upper and lower jaws, presence of gastroliths etc. etc. Zanno and colleagues posit that around the base of the mainraptoriformes there is a transititon from carnivory to herbivory based on the acquisition of these correlates within many of the dinosaurs belonging to this group. Ornithomimosaurs possess all the correlates so are expected to be herbivorous. Despite the presence of many specimens containing gastroliths (granted most are Sinornithomimus [Kobayashi et al., 2001; Kobayashi and Lü, 2003], there had been no evidence of any food remains in the stomach cavities/within the gastrolith masses until the discovery (perhaps unsurprising as plant material often doesn't preserve as well as bones) and description of Deinocheirus (Lee et al., 2014) where some fish bones were found in the stomach area with gastroliths. The authors suggest that Deinocheirus was probably omnivorous consuming fish and plant material as its diet.

Deinocheirus stomach contents (Lee et al. 2014).

Group living
Jurassic Park first showed ornithomimsaurs (or at least Gallimimus) running around in flocks like birds. To give Jurassic Park credit on this, there is a fair amount of evidence that ornithomimosaurs were indeed group living (or at least several species were).

Two clips from Jurassic Park (1993) showing Gallimimus flocking. The bottom clip is entertaining as one of the other Gallimimus in the background (arrowed) actually stops to watch the unlucky one get munched by the T. rex. I can't decide if it's supposed to show some level of bonding/care between individuals, but do look out for it next time you watch!

There have been findings of ornithomimosaurs skeletons in relatively close association with each other relatively early on, however very little is written about their group living until the discovery of Sinornithomimus (Kobayashi et al., 2001; Kobayashi and Lü, 2003) where 14 skeletons were found in very close association, and a further expedition found another 13 in another association. In both of these groups there are a small number of sub-adults to adults with the majority being juvenile individuals and it has been suggested they got trapped in a drying mudhole.

Block containing  8 individuals of Sinornithomimus dongi (Kobayashi and Lü, 2003).

Since then, a description of three North American ornithomimids found together by Sternberg in 1926 has appeared in the literature (Cullen et al., 2013). In addition there is a large ornithomimosaur bone-bed from France that may suggest that more primitive ornithomimosaurs (and not just ornithomimids) also lived in groups (Néraudeau et al., 2012).

Since the discovery of the first ornithomimosaur foot, it was realised they were very bird like, and indeed with the naming of Struthiomimus the convergence with the ostrich (Struthio) was laid bare. All ornithomimosaurs, except the screwy Deinocheirus, have elongate legs and are believed to be relatively rapid runner. They have adapted so much to high speed running that their claws on their feet have all become very flat (very similar to that of the ostrich). Muscle reconstruction of the hindlimb in Dromeciomimus(Ornithomimus) suggests that the hindlimb muscle moment arms were comparable to those of the ostrich, the fastest of the terrestrial avians presently alive, whilst Struthiomimus had smaller muscle forces and might not have been quite as quick (Russell, 1972). The construction and articulation of the limb bones as well as ontogenetic scaling suggest that ornithomimosaurs of all sizes were probably using a combination of endurance and speed to outrun predators (Paul, 1998). Of importance seems to be the arctometatarsalian foot. Such morphology appears to have evolved to evenly transmit forces to the metatarsals, this hypothesis being tested using finite element modelling (Snively and Russell, 2002) principal component and thin-plate spline analyses (Snively et al., 2004).

Much has been made of ornithomimosaur brains, with all of the literature suggesting that they have relatively large brains:

REQ (reptile encephalisation quotient - a measure of relative brain size to body mass for reptiles) for various archosaur groups. Dromecieomimus (Ornithomimus) is shown here having brains comparabale to the low end of Troodon (considered the relatively biggest brained of non-avian dinosaurs). From Lautenschlager et al., 2012.

However, I have done some reconstructions on three taxa using the same metrics, and come up with far smaller REQs (comparable to Erlikosaurus [Lautenschlager et al., 2012), and indeed work presented at SVP this year showed that Deinocheirus has an REQ similar to that of Brachiosaurus in the above image. Wait for official publications to confirm these findings, but it perhaps shouldn't be surprising seeing that ornithomimosaurs have relatively small heads (probably associated with their relatively long necks). This isn't to say they weren't smart; group living does involve a reasonable level of social interaction and cognition

Ignoring their brains (which have limited description to date so I don't want to say much on inferring the sense of smell) the other major sense for ornithomimosaurs is their vision. Ornithomimosaurs have quite large orbits.

Ornithomimus edmontonicus (RTMP 95.110.1) showing off its large circular orbits. 

The large orbits of ornithomimosaurs (Makovicky et al., 2004) would likely have hosted large eyes, Within the eyes of many dinosaurs are sclerotic rings, which are rings of bones that all articulate much like a camera diaphragm, and help keep shape and potentially help adjust the amount of light coming in. Ornithomimosaurs indeed possess relatively large sclerotic rings too (Schmitz and Motani, 2011). This suggests that vision played a large, if not the largest factor in ornithomimosaurian habits. The enhanced visual abilities may have been for locating edible plants or flowers and undoubtedly played a vital role in escaping predators by early detection (Russell, 1972). It has also been suggested that their enlarged orbits allowed mesopic behaviour (in all light levels: Schmitz and Motani, 2011) at least in Ornithomimus, furthered by some potential ornithomimosaurian finds from polar regions where it is dark for many months in winter (Watanabe et al., 2013).

Hopefully that gives you all an insight into the group of dinosaurs that I spent 3 and a bit years of my life working on. They are truly a cool group of dinosaurs and it's not because of big teeth and curved sharp claws, but because they have beaks, massive eyes, crazy feathers, ran around in groups at high speed, and if you trust Jurassic Park they stop to watch their flock-mates get eaten.

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