Wednesday, October 4, 2017

Entering the Nth Dimension of Terra-Forming, Macro-Plant Scavenging, Hive-Minded Hadrosaurs

Do you have a drink handy? Down it. Or any inebriating substance, might be time to imbibe. I can assure you that I wrote this stone cold sober but I realize that a good portion of my audience, rationally minded, might be better served with a little bit of chemical disequilibrium to help foster nth dimensional Mesozoic astral projecting. 

Buckle up kiddos, hold on to your butts...

The following, although couched in as much science as I can drape it in, is not necessarily strictly science per se. I make a lot of assumptions that prove necessary to accept in order to come up with my grander ideas here. It requires a certain amount of suspension of disbelief... so be it.  I will at times wear a scientific hat, at others a more deductive, intuitive, detective methodology will supersede the limits of the available science.  It does therefore harken back to a largely eschewed paleontological tradition of detective work, coming up with the best possible scenario with the best possible evidence.  As such it is an adventure in story telling, but I think the story is a good one - quite possibly the best one - and worth sharing. I will gladly change my take on the subject with new or better data available. Until then I think the following ideas as I outline here best answers the question of how large nesting colonies of mega-herbivorous hadrosaurs operated ecologically and, as far as I have seen, sits comfortably in the available data. 

But be forewarned: things will get weird.

To bring you up to speed it might prove useful to go review these old posts I put forth on the subject way back in the day.

Following the Poop Trail: Can Dinosaurs Be Blamed For Termites?

Some of the finer points may have changed but here I argued for unusual and novel food chains in Mesozoic ecosystems and possibly a conserved gut flora from sauropods to termites!! This idea is certainly due for a revisit.

Dinosaur Alternative Food Stuffs

Lichen munching dinosaurs? Sauropods browsing foliose lichens in high latitude polar beech forests? How did these cool, temperate high latitude forests operate?

Rot N' Roll in the Mesozoic Muck

It is in this post, from way back from May 21, 2013, that I first ventured forth an idea that I was very smitten with at the time and which has stayed with me since.

In short format: that the consumption of decayed sequoia coniferous wood by large ornithischians documented via coprololite remains from the Two Medicine Formation and often attributed to Maisaura peeblesorum (Chin, 2007) document a novel ecological relationship. Namely Maisaura was consuming rotten wood and fungi in order to deposit and provision hatchling Maisaura with a ready supply of beneficial gut flora via coprophagy (consumption of dung) but also grow tremendous fungal gardens that further nourished and provishioned young hatchlings with a readily digestible and nutritious bumper crop of mushrooms. Like a giant version of a banana slug spores would pass through the parent's digestive system and find a nice dinosaur dung patty to set up shop in. Because fungi have a more complete and easily digestible protein profile than raw plants they serve as a nice energy and growth boost for the hatchling hadrosaurs. Basically the best analogy being to the large underground fungal gardens leaf cutter ants create. A very simple, diffuse, and elegant energy transfer from detrital food chains to parent hadrosaur to offspring. Because the environmental conditions could not be controlled as in the underground gardens of leaf cutter ants, some years would be better than others in terms of fungi production. Furthermore it is not an obligate food source, the dung and whatever foodstuffs come with it - insects, mollusks, crustaceans, fungi - merely augment the growing hatchlings needs. If there is no fungal bloom of sporocarps, the hatchlings feed on other stuff.

At the time I wrote it I didn't have much readership, nor was there any more compelling evidence at hand to argue such a hypothesis past the point of imaginative fantasy so I kinda let the idea sit fallow for a couple of years. Until I woke up the other day to check out my news feed to unexpectedly discover more evidence of rotten wood munching giant ornithischians has been unveiled via Karen Chin!! Only this time the coprololites hail from the Kaiparowits formation of Utah across multiple horizons and not only contained rotten conifer wood but sizable bits from crustaceans and mollusks!! Like mana from heaven these heavenly piles of shit keep dropping on my doorstep like flaming bags of shit!! But they are bags of poop I certainly don't want to stamp out!!

While the shellfish eating aspect of this study is getting the most attention, it really is the confirmation of rotten wood munching ornithischians - probably hadrosaurs - that has the most far reaching implications. This revelation confirms that dinosaurs tapping into detrital food chains was not some aberrant activity, or the result of some ecological catastrophe but really was just a pretty regular menu option. Let me restate the obvious - multiple instances of rotten wood riddled dung from multiple horizons confirming that this ingestion was not accidental or aberrant and that it was spread across at least two species of hadrosaurs!!

From Chin et. al., 2017:

"The sizes of the exposed deposits are variable, ranging from sub-decimeter sized fragments to masses that cover several square meters and appear to represent multiple defecation events. More than 15 discrete coprolite deposits were discovered within at least three stratigraphic levels in the lower half of the middle unit of the Kaiparowits formation (Eric Roberts personal communication) at sites up to 20 kim apart."

After reading Chin's latest study and especially after reading this piece I hope we are done with the hadrosaurs as the "cows of the Cretaceous" comparison. I mean, how can you be like a cow without giving milk and eating grass anyways? Aren't those two fundamentally cow like attributes that hadrosaurs lacked? When I'm done with you I hope you start seeing hadrosaurs as more analogous to giant kaiju hive forming colonial terra forming insects as opposed to dumb ol' cows.

Indeed a main thrust of the Chin paper is that modern large mammalian herbivores in general don't compare favorably to the large ornithischians that harvested rotten wood, fungi, mollusks, and crustaceans - because quite literally large mega-herbivorous mammals don't really go after these resources to any appreciable extent. This is not to suggest that the paucity of large mega-herbivorous mammals around today shows the full extent of dietary flexibility of the recently extinct gompotheres, ground sloths, notoungulates, and others. And sure everyone loves to point out the occasional deer eating field mice or eggs or whatever BUT no one is suggesting that natural wild populations of modern mammalian herbivores are stuffing their guts full of rotten wood and crabs, they simply don't go after detrital and invertebrate resources as a way of life… they are a lot more conservative dietarily compared at least to a subset of late Cretaceous ornithischians, most parsimoniously hadrosaurs according to Chin.

I think it is also important to make an important distinction that was perhaps not hammered in hard enough in the popular discussions of the most recent Chin paper. The Kaiparowits formation records a much wetter, lacustrine subtropical environment compared to the drier, more temperate Two Medicine. It therefore stands to reason that the exploitation of rotten wood was not due to severe environmental stress as has been stipulated for the Two Medicine coprolites - that there was some sort of drought or environmental degradation that forced these animals into an unusual food resource. In short the repeated documentation of rotten wood foraging across at least two species, from multiple occurrences across multiple horizons forces us to conclude that this is not an aberrant or unusual adaptation - that we should not feel compelled to "explain it away" - but that it was a systematic and deliberate exploitation of a food source not usually associated with mega-herbivores. But why?

The recent Chin paper is on the right track. They link wood rotten wood consumption to reproductive activities both in terms of sequestering necessary proteins and elements needed for egg laying and possible latrine behavior:

"There is no definitive evidence linking the Kaiparowits coprolites with reproductive activities. However the large, multi-deposit coprolite masses may provide some indirect support for correlation of reproductive activities with the woody coprolites because latrine behavior (repeated defecation in a confined area) can be indicative of animals that are spatially constrained. Nesting activities would have necessarily curtailed nomadic movements by breeding dinosaurs."

I think Chin et. al. is correct here to link wood consumption with reproductive actives, especially colonial nesting. Unfortunately due to the limiting nature of peer reviewed scientific product they can't or won't go far enough. They are not thinking BIG ENOUGH. The truth as I will lay out is entirely more interesting and answers some very pertinent questions regarding colonial nesting hadrosaurs. Not only is it time to stop thinking about hadrosaurs as "cows of the Cretaceous" we have to re-imagine what it means to be mega-colony forming mega-herbivore because such situations are not seen today except in artificial human feed lots of factory animal buffets. It is time to start opening up the speculative vistas wider and wider...

I believe the key line of inquiry lies in elucidating the unique nature and ecology of dinosaurian reproductive biology, long incubation periods, nesting ecology, and the flow of nutrients in Cretaceous Laramidia.

Before I do that I want to stipulate two assumptions that will color my interpretations and which I shall disclose now. Keep in mind that these assumptions are not settled science, there is still disagreement. However for simplicities sake they are assumptions I will be working from.  1) Nesting colonies of hadrosaurs were in fact, real, and they were large. The parent(s) did stick around to at least guard the nest from predators until hatching. Given the long incubation time (Erickson et. al. 2017) for Hypacrosaurus (6 months) such nest guarding activities were substantial investments in time and energy. 2) Hatchlings were at least semi-precocial (Geist & Jones, 1996). They could and did upon hatching forage and move around on their own. This does not preclude some amount of direct provisioning by parents, but it is no required. The young continued to use the nest for creche style groups, sleeping and protection especially huddling for warmth,  for some time after hatching (nidicolous i.e. nest bound vs nidifugous i.e. nest leaving). Parental provisioning may or may not have been negligible or non-existent, food stuffs gathered in the immediate vicinity of the nest by the hatchlings on foraging expeditions of the abundant invertebrates, fresh greens and fungi all of which were fostered by the parent hadrosaurs woody dung.

The Elephant(s) in the Room - Long Incubation Periods & Mega-herbivore Nesting Colony Environmental Dilemma

There is an elephant in the room that no one is talking about with regards to nesting colonies of hadrosaurs. For some reason(s) these issues never seem to crop up in discussions of hadrosaur nesting colonies. In fact there are two elephants in the room that need to be coupled and dealt with.

Elephant in the room #1: Large herbivores don't stay tethered to one spot very long, they will quickly strip it of vegetation. 

Elephant in the room #2: Dinosaurs are showing strong signs of long incubation periods.  

Notice that elephant #2 collides with elephant #1 and exponentially increases the dilemma of large colonially nesting hadrosaurs. If you have 6 months of incubation, that gives you a minimum of six months of nest guarding duties. Those six months will create an increasingly large radius of environmental devastation around the nesting colony. The hatchlings inherit a wasteland - not good!!

Although John  Horner does not mention the dilemma of feeding stationary colonies of nesting hadrosaurs in his influential book Digging Dinosaurs he does at least acknowledge the destructive powers in the 10, 000 strong mega-herds of Maisaura he claims are represented at site Camposaur (pg. 138) :

" I wonder a bit about how these kinds of herds affected the environment. Certainly the herds had to keep on the move. They must have stripped one area and then moved on to the next."

Probably the best treatment of the environmental carnage nesting hadrosaurs colonies would impose is taken from Dale Russel's epic tome An Odyssey in Time: The Dinosaurs of North America page 151:

"… Nearby sources of food would soon be depleted, and the parents would had to range ever more widely each day… The rains came, the brushlands greened, and the nests burgeoned with stumbling, chirping hatchlings. Lines of parents streamed along muddy, dung-filled paths to and from the colony. The air was rent by sounds of breaking branches, as trees and shrubs were stripped of their foliage in an ever-widening circle of environmental devastation… Carcasses of dead hadrosaurs began to dot the stripped shrublands, and the area began to acquire something of the appearance of a battlefield."

At least in the case of Maisaura, suggestions that not just one, but maybe several hundred or even thousand multi-ton herbivorous dinosaurs hunkered down for maybe half a year in a rather concentrated locale is mind boggling. There would be an ever increasing radius of devastation surrounding such colonies, making it increasingly hard for nesting hadrosaurs to guard their nest and giving hatchling hadrosaurs a veritable wasteland to inherit.

Overall, the prospect of hadrosaur nesting colonies does not paint a pretty picture but just wait, it gets even bleaker...

The pesky little problem that Dale Russel did not foresee is the long incubation period of non-avian dinosaurs, especially pronounced in hadrosaurs. While the recent Chin paper is correct in pointing its nose in the direction of reproduction (but not for ecological reasons for replenishing calcium and other nutrients) it does not go far enough in making the link between long incubation times, colonial nesting, and the unusual situation of many tons of mega-herbivores remaining in a relatively small area.

The recent work on dinosaur incubation times (Erickson et. al. 2017) is potentially every bit the game changer on the behavioral front as ornithoscelidia is on the phylogenetic front - I don't know why people have been sort of, "whatever" about it? Or just glossed right over it. Maybe they can't see the forest from the trees on this one, but, well, I'm not waiting for them… Hypacrosaurus was one of the dinosaurs in the Erickson study and analysis reveals it to have an incubation period of… wait for it… six months!! Which implies that other hadrosaurs - such as the ones that putatively laid those rotten wood coprolites in the Kaiparowits and Two Medicine formations respectively - had similarly long incubation periods. Not to mention that there is abundant evidence for large Hypacrosaurus nesting colonies (Horner & Currie, 1994). This chain of thought should get us thinking: how did such nesting colonies not become ecological wastelands, given the long incubation periods?

From Dinosaurs' Long Egg Hatching Times Might Have Led to Their Demise paleontologist David Varricchio said:

"These long incubation times likely restricted dinosaurs… If they had parental care, for example, parents would be bound to a specific spot for months (up to six months) of a given year. This would limit migration. Perhaps it would also hinder dinosaurs' response to environmental change."

No wonder so many people are dismissive of the long incubation periods of dinosaurs, it creates a scenario too weird and unstable to justify ecologically. That is until we stop thinking so narrowly about what it means to be a dinosaurian "herbivore" and how long incubation periods are not the "negatives" so many people ascribe them to be but absolute positives.

There is a way to flip this predicament of large mega-herbivorous mega-colonies, long incubation, and environmental degradation on its head. We can actually inverse this equation to come up with a spectacularly novel, productive, and intuitive outcome that better explains the success of these colonies and is not disharmonious with the evidence at hand.

Long incubation periods and nest guarding - potentially via bonded mates - set the stage not for evolutionary failure but forced hadrosaurs into a unique kaiju hive mind of ecological terra-forming. It was the long incubation periods that let hadrosaurs exquisitely render and shape their own environment. They were not victims of long incubation times as often posited from the Erickson study but beneficiaries of them.

The key to imagining long incubation periods as absolute positives is the woody coprolites - the rotten wood consumption that Karen Chin has been documenting for some time now. Hadrosaurs learned how to crack the code of exploiting old growth coniferous forests in a manner that no mammal has done. This allowed hadrosaurs to graft themselves into ecosystems that are today not notable for large abundant mega-herbivores but which dinosaurs, especially hadrosaurids, thrived in: large old growth coniferous forests. Even today large old growth coniferous forests are powerhouses of life, just not necessarily powerhouses of large vertebrate life like they were in the Mesozoic.

Imagining Hadrosaurs as Analogous to Sea Birds - "Hunting Down" Patches of Rotten Wood

Big Sur redwoods Duane Nash

Ever wander through the redwood cathedral forests of Northern California or the old growth coniferous forests of the Pacific Northwest? These temperate rain forests certainly inspire the mind, and if you are inclined towards Mesozoic musings, they certainly bend the mind towards thoughts of how dinosaurs slotted into these settings as forests very similar harken back to those days. An interesting dilemma occurs though in such mind questing: such forests today are not known for a particularly diverse or large mega-herbivorous bestiary. Sure you have your bears, deer, elk and what not. However during the Pleistocene there is not much suggestion that things were much different with the large megafauna preferring open grasslands, savanna, and hardwood forests. Dinosaurs were doing something very different from mammals. Sauropods answer the question in part by being large enough to push over the great conifers or simply reach up and bite their foliage. Hadrosaurs were doing something completely different from even sauropods. Hadrosaurs cracked the code, that mammals have so far failed at cracking, by consuming the entire tree itself after it died. They outsourced the initial digestion - the delignification - to fungal partners and then, in an exquisite transfer of nutrients, they shuffle the organic wealth of the old growth forest to their nesting colonies - in more open conditions - which benefits hatchling recruitment. They also eventually die, quite possibly in a forest, where they return the nutrients they stole. They thus speed up the "slow" transfer of nutrients in modern old growth coniferous forests.

Hypacrosaurus foraging on rotten wood resources by Duane Nash

The reason detrital wood resources are important is that they let us think about hadrosaur nesting colonies less like giant conglomerates of typical mega-herbivores and more like nesting colonies of seabirds. Seabirds provide a model for how large groups of active, high energy homeotherms can nest together for several months at one location. They venture to sea to forage for high quality, dependable food stuffs. They can follow other birds and marine life (like cetaceans) to predictably rich foraging grounds. Hadrosaurs actively nesting, I will venture, switched from a foraging strategy of mainly green, live growth towards detrital rotten wood resources not immediately prior to egg laying but subsequent to egg laying and during the nest guarding phase. Like seabirds that depart breeding colonies to locate highly discrete but dependable food sources parental nesting hadrosaurs departed the nesting colony to locate highly discrete but dependable rotten wood resources, completely or largely eschewing live plant resources en route. This ensured several critical aspects that ameliorated conditions for the hatchlings;

1) Vegetal resources were not demolished in the immediate vicinity of the nesting colony which ensured adequate forage and hiding for the hatchlings at a critical juncture;

2) Parent hadrosaurs could follow known paths to localized patches of dead and rotten wood falls. A type of hadrosaur "hive mind" comes into play here that as soon as some hadrosaurs know where to find the rotten trees, they all just follow along (like seabirds);

3) Because these dead trees might be several kilometers away from the nesting colony the net movement of rich partially degraded dung brings a net influx of nutrients into the immediate vicinity of the nesting colony;

4) This flips the notion of nesting grounds becoming "ecological wastelands" on its head, hadrosaurs were terraforming the area around nesting colonies to favor their own hatchlings - indeed the highest concentration of dung would be adjacent to the nesting areas;

5) Hatchling dinosaurs could forage on dung itself via coprophagy, invertebrates attracted to dung, fungi within the dung, spores and seeds sprouting from the dung. In short, imagining detrital wood foraging in this manner solves every problem that long incubating, high metabolism, nesting "mega-herbivorous" hadrosaurs presents. Nesting grounds don't become run down ecological wastelands but booming dinosaur baby powerhouses, the actions of the parents directly terra-forming the immediate vicinity of the nesting colony for the hatchlings benefit. The richest foraging grounds are actually those in the immediate vicinity of the colony!!

Karen Chin has been documenting a particularly nuanced and diverse invertebrate interactions with dinosaur dung, especially from the Two Medicine formation. No less than seven taxa of terrestrial and aquatic snail were found within coprolites from that formation by Chin (2009). Throw in the abundant evidence of back filled burrows of dung beetles and one has a good buffet for young, growing hadrosaurs. For me, this diet of dung based resources is a much more plausible, nutritious, and easily obtainable diet than predigested gruel or some sort of crop milk. One has to imagine that such rough, fibrous plant stuffs as adult hadrosaurs were consuming is not going to produce the richest, most protein laden gruel or crop milk for young and quickly growing hadrosaurs. However if young hadrosaurs can tap into the riches provided by six months or more of woody dung accumulating adjacent to their nest their you have a much more nutritious buffet of insects, gastropods, small invertebrates, fungi, young seedlings, fern fiddleheads, and coprophagous delights. Much more simpler and direct in my estimation.

Maisaura kindergarten by Duane Nash

A Transfer of Nutrients: Dinosaur Terraforming

Duane Nash w/fallen coastal redwood. Jedidiah Smith Park CA

As we start to flip off the switch in our brains of giant "Cretaceous cow "and on to giant Cretaceous macro-plant scavenging, detrital hunting seabird-banana slug,  the ecological scenario starts to switch. Large coniferous dead fall trees are predictable in old growth coniferous forests and they stick around for hundreds of years. A massive fallen old growth redwood Sequoia would provide adequate forage for possibly hundreds of Maisaura!! Indeed massive dead trees are the largest biological caloric windfalls of all, more than dead sauropods or whales. These are truly massive stockpiles of carbon, nitrogen, fungi, and invertebrates. Hadrosaurs would tap into the most productive and important trophic category in old growth forests - the detrital food chain - a niche largely unexploited by modern mega-herbivorous mammals. Once located or "hunted down" - hadrosaurs may have had to march several kilometers to find them - hadrosaurs would make periodic trips from the nesting colony, perhaps even switching egg guarding duties between mates, to the dead falls. We start to imagine these hadrosaurs less like typical herbivores but more like giant macro plant scavengers that let fungi do the hard work of digestion while they reap the rewards. All the while a transfer of nutrients is occurring as dino dung patties are being deposited away from the old growth forests and adjacent to the nesting colony. This energy transfer creates a very different sort of ecosystem from the old growth coniferous forests of the Pacific Northwest of North America. Here decaying trees take centuries to break down and the nutrients stay in the forest. In hadrosaur mediated old growth forests the transfer of nutrients may have been much more rapid and multifaceted. Nutrients being cycled out of the forest to the nesting colonies and possibly back into the forest as hadrosaurs died in secluded forest haunts or dragged into deep redwood groves by tyrannosaurids.

When the hatchling hadrosaurs emerge, precocial and ready to fend for themselves, hopefully coincident with rains they are met with a riot of new - dino dung fertilized - growth. The carbon and nitrogen rich chewed up rotten wood dino patties built up over six months of nest guarding and potentially several hundred years worth of colony fidelity is literally dinosaur terra-forming. They have ameliorated conditions for their progeny on a grand scale. Far from being an ecological wasteland the areas around hadrosaur colonies may have been almost preternaturally abundant with life!!

Today redwoods can thrive in small microclimates in borderline semi-arid conditions, as I documented in Redwood Grove Along the Big Sur Coast: Southern Limits where I explored some of the southernmost occurrences of redwoods along the California coast.

southern Redwood Big Sur coast credit Duane Nash

As the picture above attests you can actually see redwood groves growing on the fog enshrined cooler north facing slopes and ravines right adjacent to the traditional xeric chaparral of California. Similar micro habits may have allowed patches of large conifers and sequoia to thrive in semi-arid habits, like in Two Medicine, where colonies of hadrosaurs may have been in striking distance of such stands and their bounty of decaying wood.

A Diffuse Hive Mind Emerges: What is Good for the Colony is Good For the Individual

When we start to look at hadrosaur nesting colonies not as individuals in a collective but more as a meta-organism a "hive" if you will we can start to better make sense of how these giant amalgamations of bio-mass operated in a manner that did not ravage their environment but actually enhanced it. They were terraforming it. Remember that these were nesting operations that - at a minimum - may have required parental supervision of the nest for over half a year. When every rule of modern mega-herbivorous mammals is broken by remaining stationary in a locale for such an extended time it is time to invent new rules. It is actually not hard to imagine a certain evolutionary feedback loop coming into play.

Rule #1: Clutch-mates are the social basis. Your clutch mates are family. You share a genetic heritage. Whatever you do to augment their chance of survival also enhances your genetic legacy via shared genes.

Rule #2: The nesting colony is sacrosanct. Just as clutch mates form the basis for social groupings in dinosaur life, it is indeed clutch mates that establish and form the nucleus for new breeding colonies. Over time these breeding colonies grow in size and scope but their is still a shared genetic legacy.  This is part of the reason, I will venture, Laramidia was so diverse in giant herbivorous dinosaurs. Colonial nest site fidelity and shared genetic heritage decreed a more insular evolutionary swap stakes.

Rule #3: Large colony size is inevitable. Playing the numbers game against predators would push colonies towards the largest possible size. As colonies grew in size and scope they would reach an ecological imbalance. Food resources would become so impoverished and ravaged that the travel time to good patches of vegetation for adults would become untenably long. They could no longer make the treks to food and return to guard eggs in a timely and efficient manner. Likewise in such a scenario hatchlings would emerge to a world of utter devastation - all available vegetation was scoured to the ground in the vicinity of the colony ( I am going with the notion that hadrosaurs were fairly precocial, but may have still sheltered in the nest). In such scenarios we would see colony collapse. The increasingly long treks for vegetal resources by adults coupled with the increasing radius of environmental degradation would spell disaster for adults - increasingly exposed to predators, fatigue, disease, and stress - and hatchlings inheriting an impoverished ecosystem. Colonies would have to change their current ecological course OR FACE OBLITERATION. Unless such large colonies could find a way to skirt the issue of environmental devastation then small grouping or solitary egg laying would be more beneficial than colonies.

Corollary 1: Solution - for adults during nesting periods they eschew greenery and tap into detrital food chains. This solves the problem of environmental collapse within the vicinity of the colony. Such a diet, bolstered with fungi and animal protein, is evinced by the coprolite remains. Hadrosaurs now go on long distance foraging ventures to secure rotten wood, largely foregoing the greenery around the colony. It allows us to compare hadrosaur colonies to sea bird colonies: which instead of "hunting down" discrete patches of rich oceanic life, hadrosaurs "hunted down" discrete patches of dependable detrital resources. Large dead coniferous trees are, then as now, the largest organisms that have ever lived. The haul of carbon, nitrogen, fungal, and animal resources is immense. In processing these giants hadrosaurs circumnavigated the "stationary megaherbivore dilemma", avoided competition with their offspring, and actually transferred nutrients via dung to the immediate vicinity of the colony. They thus terraformed their environment for the benefit of their offspring and enhanced their genetic legacy. It was a WIN-WIN situation for the hadrosaurs.

Corollary 2: The colony becomes a semi-permanent home. Once the detrital-dung-fertilizer feedback loop is set in motion, the bigger the colony gets the more successful the colony becomes. Quite simply the more animals are spreading dung around the vicinity of the colony, the better quality forage is produced for the young i.e. fresh growth, invertebrates, fungi, the more juvenile recruitment succeeds. The colony becomes a sort of metropolis where the richness provided actually allows hadrosaurs to remain there well past the hatchling stage - a phenomena evinced by the multiple growth stages present at Maisaura nesting colonies and mass death assemblages. In fact I will venture a speculative guess that many or most hadrosaur mass death assemblages do not represent migrating herds but more of a time averaged death assemblage pulled from hadrosaurine colonial metropolises.

Corollary 3: The glass ceiling, a potential limit to colony growth at which stage colonies may shrink or experience collapse. It is not necessarily the availability of green growth but the availability of dead growth. Once colonies have cleared all of the old, dead, rotten logs out of the area adjacent to their colony it may in fact be time to move the colony. The hive relocates. It secures a territory with untapped detrital resources and sets in motion the terraforming process anew.

Migrating Mega-Herds or Hadrosaur Metropolises?

credit Metropolis © Friedrich-Wilhelm-Murnau-Foundation fair use

And here is where I want to really go for it - really blow your mind... that all of this was a warm up, I mean, why stop now? A fascinating topic in paleontological musings is the holes in the fossil record and how, slipping between the cracks, could be lost and vanished "higher" beings perhaps not unlike ourselves in some scope of magnitude, some ability to shape and manipulate their environment. Let me offer that such beings might not be entirely reminiscent of us in shape or appearance - but still are like us in some quintessential way set apart from other animals in the magnitude of their effect. And let me offer such beings have been staring at us in the face all along that colony forming terraforming hadrosaurs were modifying and shifting environments on a grand scale far beyond that of even typical mega-herbivores. That hadrosaurs were not only ameliorating conditions for their hatchlings but for all growth stages "ontogimorphs" of their  individual species as well. That their terraforming via dung actually created, over generations, prolific hadrosaur citadels or metropolises. Thriving with multiple generations and growth stages of hadrosaur as well as a panoply of commensal, symbiotic, predatory, and commensal species. That these metropolises, somewhat like our own, had their own cadence of birth, growth, decay, and renewal. That these hadrosaur modified environments - metropolises based on an economic driver of conifer dung - not only existed but that we might have tantalizing proof of them.

Although not as well known as the saga of Egg Mountain and the miraculous nesting colonies of Maisaura and Hypacrosaurs that Jack Horner is famed for it is his sight called Camposaur where, "conservatively", 10,000 animal strong Maisaura herd death assemblages are located. No one really knows how these death assemblages formed, if they were just one single event or multiple events. The story usually goes something like this: volcanic out gassings killed a migrating herd of at least 10,000 Maisaura. Volcanic soils partially preserved the bones as fossils, later redeposited in floods.

But let me paint another picture that this "herd" actually represents a primarily residential assemblage of Maisaura killed over time in one locale. An astonishingly productive Maisaura metropolis fertilized not just by conifer dung but by periodic volcanic ash falls. They were double dipping. But there was a price to be paid for this Maisaura heaven. Periodic out gassings of the volcano would kill off vast droves of Maisaura. Bones would be partially fossilized in the volcanic soils, seldom disturbed by scavengers due to the sheer volume of animal mass and volcanic killing fields and periodically swept away via floods for later deposition and preservation.

Altogether this scenario of multiple killing events of a primarily residential Maisaura metropolis is not at all incompatible with the data as we know it. As opposed to one giant migrating herd dying en masse in a single volcanic killing event, multiple killing episodes of perhaps a more modest residential colony might actually be a more parsimonious explanation for Camposaur.

All of this is very typical of our species in-born bias that we would not recognize such complex biological ecological systems from the past because we are looking for something that resembles our own tool mediated civilizations. Instead we should expect such systems more to resemble the most dominant, successful and ubiquitous colonial living arrangements known - social insect hives. That dinosaurs would evolve such systems that parallel insect societies should really not be that surprising in hind sight, they had more than enough time to evolve such complexity - even if it was not tool mediated or under the auspices of some great intellect.

Brigitte Helm Metropolis

There is perhaps a moral in all of this. Perhaps colonial hadrosaurs that tapped into detrital ecosystems, cracked the code of conifer forests, and broke all the rules of what it means to be a mega-herbivore… perhaps they offer a model of right living in hot house worlds. That using dung to form metropolises and outsourcing digestion to fungi - maybe these are tricks we can learn something from?

Anyways, something to chew on...




Chin, K., Feldmann, R.M., and Tashman, J.N. 2017. Consumption of crustaceans by megaherbivorous dinosaurs: dietary flexibility and dinosaur life history strategies. Scientific Reports 7, article number 111163. online

Chin, K., Hartman, J.H., and Roth, B. 2009. Opportunistic exploitation of dinosaur dung: fossil snails in coprolites from the Upper CretaceousTwo Medicine Formation of Montana. Lethaia 42: 185-198.

Chin, K. 2007. The paleobiological implications of herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana: why eat wood? Palaios 22: 554-566.

Erickson, G.M., Zelenitsky, D.K., Kay, D.I., and Norell, M.A. 2017. Dinosaur incubation periods directly determined from growth line counts in embryonic teeth show reptilian-grade development. Proceedings of the National Academy of Sciences. vol 114, no. 3. online

Geist, N.R., Jones, T.D. 1996 Juvenile Skeletal Structure and the Reproductive Habits of Dinosaurs. American Association for the Advancement of Sciences. online

Horner, J., and Gorman, J. 1990. Digging Dinosaurs

Russel, D.A. 1992. An Odyssey in Time: The Dinosaurs of North America


Beetle Boy said...

Wow . . . I don't even know what to say but that - that was incredible reading! You raise some good points, and you're really not afraid to take speculation to its extreme, something that most simply can't do. I applaud you sir!

Duane Nash said...

I'm glad you got something from it Beetle Boy.

Nick Fonseca said...

Very interesting idea! I appreciate the notion of seeing herding and long gestation not as hindrances to survival rather as being key to survival.

Another colony to consider would be the emperor penguin. Apparently after egg laying the females go off to sea and forage for 2 months to replenish their energy stores. In the mean time, the males stay with the egg balanced on their feet for approximately 64 days until the egg hatches. All told the males may fast for almost 115 days after arriving at the nesting site. That is almost 4 months with no food. "I learned that from reading a children's animal book to my 6 year old haha." I wonder if there is the possibility for half of the adult herd and part of the non mating herd to leave the nesting site for part of the nesting phase to alleviate food shortages?

Bk Jeong said...


If this was the case, ecosystems around hadrosaur colonies could have been insanely complex. We would have tyrannosaurs and dromaeosaurs taking up hunting sites around these colonies: small enantiornithines, small ornithischians, etc eating the fungi and vegetation that grew from the dung; etc...

Duane Nash said...

@Bl Jong If you read digging dinosaurs by Jack Horner that is exactly what he found.

William Bailey said...

Wow! One of your best articles in terms of argumentative strength and also in terms of creating a larger, believable picture. I also greatly appreciate how you see those "hindrances" as strengths, which is quite more plausible on an evolutionary standpoint. Of course, you still have your style but I find it better used in this article. The form serves the content. Bravo. ;)

PS: making hadrosaurs interesting is a most impressive feat and a necessary one. ;)

Nick Fonseca said...

I agree, seeing the "hindrances" as strengths makes more sense. By that logic any adaptation should be considered a hindrance. I like the idea of thinking about the ecosystem as a system not just a backdrop for dinosaurs to run around in.

Duane Nash said...

@Bk Jeong In his book Horner also spends a lot of time on Orodromeus which appears to have nested by Maisaura, also appears to be evidence of Troodon type eggs (?) & remains, some azhdarchid stuff, tyrannosaur teeth. ALso the invertebrates Karen Chin has been documenting for some time. It may have been a humming place.

@WilliamBailey Thanks. "Making hadrosaurs interesting is a most impressive feat and a necessary one" I'm flattered.

@Nick Fonseca Regarding emperor penguins and seabirds in general. It is interesting and something I've thought a lot about comparing hadrosaurs to seabirds. One of the challenges is that seabirds can go idle and fast for a while because their foodstuff is comparably richer than a herbivores, even hadrosaurs augmenting their diet w/fungi and crab (yummy) is not as rich as krill, squid, and oily fish. Another question I would poise with herbivores going for long spells without food (a month or two) is that this might have deleterious effects on their gut flora. Of course they could probably scarf down some dung piles to repopulate… If mates did trade off duties I am leaning to something on the order of a week or two not several months. Another possibility is that there was no partner trade off, the collective defense of enough adults in the colony at any one time might be good defense. This aspect gets into notions of shared genes among colony members.

Bk Jeong said...

I quite like how you look at the supposed "weaknesses" and "impossibilities" of animals and flip it into an evolutionary adaptation.

Because evolution only works if a species can actually survive.

"Carnosaurs/terror bird jaws were too weak to kill prey or eat bone" => shows that they were killing large prey and eating bone in methods that actually need low bite forces

"T. rex isn't agile" => not a problem if you are using straight-on charges to ambush prey

"Microraptor cannot be nocturnal because iridescence" => Microraptor wasn't analogous to iridescent birds anyways, but it's ecology was a lot closer to iridescent stealth-hunting snakes

"Elasmosaur necks were a weak point" => shows that such a huge neck is actually muscular and strong, and helps with stealth-hunting and tussling with prey

And now hadrosaurs....

The Eurypterid said...

I think the repeats of 'kaiju hive-mind' are a teensy bit sensational, but a colonial lifestyle for hadrosaurs seems pretty reasonable. A point in favour is that large modern herbivores already use hive intelligence- wildebeest herds cross rivers safely by probing the opposite bank with a growing number of animals, then using numbers to wear down the nearest possible exit into a safe, easy passage, all the while deterring predators with their numbers.

BrianL said...

I think this is probably the best post you have ever written even if I agree that it need not have been as sensationalist. Do you think this same strategy also applied to other large colony-nesting dinosaurs or do you suppose this was limited to hadrosaurs? What about, say, sauropods and ceratopsids? Also, do you still see (some) ankylosaurs as adapted to take advantage of mega-colonies such as the ones you propose?

Duane Nash said...

@The Eurypterid the "kaiju hive-mind" is a reference to Pacific Rim. For me personally I like the analogy, even if it is patently sensationalistic. It lets us think of dinosaurs more in a mindset of animals that had to sequester huge amounts of resources, go through multiple ontogimorphs, invest huge amounts in nesting resources, and were large and spectacularly adorned and weaponized - these same attributes are found in various kaiju. It is certainly a difference in degree but not necessarily type and I think an interesting example of nature imitating art - which the art was initially inspired by the nature (kaiju always having a strong dinosaurian influence). Yes it is intentionally sensational but I think it serves a greater purpose of thinking of these animals in new and different ways. Not merely as "cows of the Cretaceous" or whatever…

Point taken regarding wildebeests. Indeed there does seem to be a tipping point where herds of any type reach a size where a single leader or band of leaders falls away to more of a hive-like collective intelligence.

@BrianL On other colony-nesting dinosaurs? For simplicities sake I focused on hadrosaurs because I could at least point to "look this might be why they are chewing up rotten wood." I honestly have not put to much effort into looking at other colonial nesters if there are any clues pointing us in this direction. Protoceratops is not very massive and afaik the colonies of various par avian theropods are not truly massive… sauropods are interesting because the size difference between adult and hatchling is so profound. They may have fertilized the areas around colonies just because of the size of their "offerings" even if they were, as it looks now, more of the lay 'em and leave 'em style. I don't think we can say anything concrete about derived ceratopsid nesting habits… that seems to be wide open. Of course it is very possible some hadrosaurs were not colony forming...

Duane Nash said...

@Brian L Sorry I forgo to mention your query about ankylosaurs. Yes I do think ankylosaurs would have profited from hadrosaur colonies. I think they would have relished dung, eggs (esp rotten ones), maybe even scavenged and "cleaned up a bit" for the colony. More on ankylosaurs in the not too distance future.

khalil beiting said...

Sorry for not commenting in a while Duane. I just started college so I don't have too much free time (besides being lazy and sleeping), but I read your posts regardless.

But I'd like to say that this is by far my favorite post from you as of yet. It's not as sensationalist as you say it is, and although it's somewhat speculate it makes PERFECT sense and clears up soooo many questions that many if not all of us have had for years.

And this has so many implications for the ecology and evolution of the macro and micro fauna/flora of Hadrosaur bearing ecosystems. So much I don't even know where to begin.

And I believe Andrea Cau pointed this out a few posts ago, but can you please make a peer reviewed paper? I know you said how restricting and monotonous they are (which I also agree with), but that's the major way the scientific community works. You seriously need to rethink it, because it could really escalate you in the paleo community and bring a whole flock of people. And I know without a doubt that several other researchers and scientists would gladly help you with it. Your work is worth the fine print.

Beetle Boy said...

This makes me wonder if there were any hadrosaurids that practiced brood parasitism - not actually caring for their young at all, just laying their eggs in the nesting grounds of other hadrosaurids species.

Duane Nash said...

@Khaili Hey thanks. We can talk about peer review via email if you want. Let me know if you have any questions about open science.

@Beetle Boy I remember that suggestion somewhere else but for a theropod nest if I remember?

Beetle Boy said...

I think an oviraptorid nest was found containing oviraptorid eggs, but also 2 small troodontid skulls covered in scraps of egg material - so I think there was some suggestion that there may have been brood parasitism going on there. Honestly, I'm not convinced by that particular find, but I do think that the possibility of brood parasitism is a very likely one with dinosaurs.

Duane Nash said...

@Beetle Boy yeah that was the one, thanks. But I also seem to remember some mixed up nest? or collection of crocodilian eggs with dinosaur eggs? am I imagining that one?

Beetle Boy said...

Hm can’t say that I’ve heard of that one. It would be interesting if we found a nest which showed convincing evidence of brood parasitism in dinosaurs, and its not impossible either, so I’m keeping my fingers crossed.

Anthony Morris said...

Hi Duane, I'm a big fan of your writing and musings. This one is as good as anything I've seen written about dinosaurs behavior. I find myself thinking about dinosaur behavior a great deal more than I would have Without your work, I suspect. I have noticed the complex workings of dead trees from time to time and seen bears and pigs going after grubs and such and even seen a herd of cows chewing up the remains of a rotten stump that a neighbor had pulled up. When I read this I found myself nodding away because, of course, I'd seen this before. My biggest difficulty is conceiving the enormous amount of wood in an old-growth forest. I've never been in one myself. Something to put on my to-do list.

Duane Nash said...

Thanks for commenting Anthony. Comments like this make it all worthwhile. Hope you get into an old redwood forest, they are special places.

Lam Luong said...

I remember seeing in some of your older posts about sauropods and large ceratopsids knocking down trees while foraging - something that would benefits renewal of fresher greenery and other herbivores attracted to the new growth, including hadrosaurs when they seek rotting wood for their colonies. I don't remember if you've mentioned this before, but I'm pretty sure I've seen a lot of hadrosaur/iguanodont species coexisting with those of sauropods and/or ceratopsids in various formations. I was wondering if it might be possible for hadrosaur nesting behavior to be interrelated with "deforesting" behaviors of sauropods and ceratopsids? Or even their nesting behavior - perhaps colonies of hadrosaurs often associated (or even intermingled) with those of sauropods/ceratopsids ?

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