Following the Poop Trail: Can Dinosaurs be Blamed for Termites?

December has come around and that means here in southern California it is time to lay out some theropod crap underneath the citrus trees. What I am really talking about is chicken manure that, liberally applied underneath citrus trees, provides a great slow release fertilizer for lemons, mandarins, oranges etc etc. You see, citrus trees love and need lots of nitrogen- when you notice bleached out leaves where the older leaves lose color first- chances are your citrus is lacking nitrogen. But be careful - too much at once, especially from water soluble liquid fertilizers- will create nitrogen burn that can harm the plant (throws off the osmotic balance). The water uptake is too high in the hot season and the tree may just take in too much nitrogen all at once in the high heat as it sucks up more water.

Nitrogen burn

The trick with  putting out chicken manure during the winter season, as opposed to the summer,  is that the chicken manure gets to slowly compost over the cool, wet rainy season. The tree is not actively growing mid-winter, but come spring the chicken manure will have nicely decomposed into a nice, slow trickle of nitrogen. This nitrogen input will give your citrus a nice, green glossy sheen due to the nitrogen when the weather warms and the growing season commences. And that nitrogen, remember, comes from a dinosaur. And dinosaur poop, as I have discussed before here, is fundamentally different from mammal poop. And this is due to the inclusion of uric acid in the excrement- you know its that white chalky stuff in bird poop that does a number on your car's paint job? Reptiles produce uric acid, including crocodiles and birds. And by extension, invoking phylogenetic bracketing, dinosaurs did too.

Both uric acid (diapsids) and urea (anapsids) void the body of nitrogenous waste produced from digestion/metabolism. But uric acid is much less water soluble, and hence more chunky, than urea- which is mainly water (i.e. pee). Of course since organisms that produce urea lose a lot more water than those that produce uric acid this has potential repercussions for water balance/ behavior/adaptations towards aridity. But for our purposes here you can think of uric acid as a heavy bombardment of nitrogen while urea being less intense, but more readily available for uptake by plants (water soluble). Of course that doesn't mean you should start fertilizing your garden with your pee. Especially if you eat a lot of protein- protein means high nitrogenous waste, which means more nitrogen in the urine.
You should at least dilute it by 75% and maybe avoid spraying it on the edibles.

But now imagine yourself as a putative Mesozoic groundskeeper.

Because of the high carbon dioxide levels in the atmosphere growth rates are very high following the monsoonal rainy season. This leads to abundant, but not necessarily nutritious plant growth. There is loads of it, but it is composed of highly fibrous material- loads of cellulose and lignocellulose. Trying to shape this yearly riot of growth into nice looking well groomed parkscapes proved illogical on your machinery never mind your back. And the thin oxygen levels further diminish your resolve. Dense thickets of rhizomatous ferns, spiky stands of cycads, tenacious drifts of horsetails and tangles of cheirolepidiaceae are more than a match for you. And you quickly learn the futility of thwarting the massive gangs of megaherbivores. Nothing short of 50 foot high electricity fencing will keep the sauropods and iguanodontids out of your garden. Let's just say you become a more enlightened organically inclined gardener. You try to take advantage of the natural cycles in the system.

As you watch the movements and foraging behaviors of the dinosaur herds you notice a couple of patterns. Although their impact is immense and dramatic, it is short lived. The herbivores are very nomadic. This is due to a couple reasons. The animals don't want to eat themselves out of house and home. Parasites and predators zoom in and target the herds that stay in one spot. And the animals do not want to be late in getting to the nesting colonies. Significant selective forces have shaped this system and these large herbivores into this ecological arrangement. And as you get more and more in tune with what is going on with the system you start to notice that a central hub in the ecosystem is the dinosaur poop itself. These dung piles represent one of the predominant transfers of energy in the ecosystem- short of a 40 tonnne sauropod dying. And the shits are enormous. And common. And highly fibrous. These guys are sometimes biting off more stems than leaves it seems. And even though the herbivores themselves only visit your plot of 12 acres for about 2 weeks a year, when they are absent these dino dung piles still remain the central foci for many of the organisms in the environment. Spores and seeds that pass through the dinos digestive system umharmed sprout in the dung- so that it is a recruiting spot for ferns, fungi, moss, and cycads- a little miniecosystem. Amphibians hide in it, attracted to the moisture and invertebrate prey. Hatchling dinos eat it both for food, get a little protein boost from the infauna, and to inoculate themselves with the bevy of microbial endosymbionts necessary for a lifetime of plant consumption. Smaller mesopredators such as pterosaurs, predatory mammals, sphenodonts, and land crocs also frequent the dung piles to snatch up the small prey items.

Campos Arceiz. (c)

But ubiquitous in the dung are cockroaches. In the dung, under the dung, scurrying everywhere all over the place are cockroaches. But these roaches, although superficially similar in appearance to the ones that kleptoparasitize humans, are actually quite different. The mated pairs are monagamous, forming multigenerational families- over time colonies are formed in the largest piles of dung. Numerous antechambers and nurserygrounds crisscross the largely alive pile of excrement. In these cockroach nurseries the young gather, after molting their insides and outsides, to reinoculate their intestines with the microbial flora necessary to digest the fibrous dung. The cockroaches most resemble extant wood roaches of the genus Cryptocercus- a social roach that nestles closer to termites than other cockroaches. Indeed these Mesozoic decomposers represent the ancestral stock from which termites and wood roaches arose from- their common ancestor.

And a recent paper, Cockroaches Probably Cleaned Up After Dinosaurs available for free at PLOS ONE, suggests that there may be a degree of truth to the speculation I offered above. Blattulidae, an extinct but widespread, persistent and evolutionarily conservative family of Mesozoic cockroaches, are posited as important consumers of dinosaur dung. As evidence the paper sites a roach preserved in amber with fecal remains both inside the roach and in the act of voiding. Wood fragments are suggestive of being modified by digestion prior to the roach consuming them. Although PLOS ONE has received some flack from people I confer with regarding the integrity of the publication and some of the papers that this article sites are questionable in my opinion, the paper does make a compelling argument for Blattulidae roaches dining on dino crap.

The preserved poop is in pic d and e

And now lets revisit the earlier part of the post (dinos made high nitrogen crap), with the revelation of dino-dung eating roaches. Now remember large doses of nitrogen (uric acid laced dino dung) especially combined with high temperatures and hence high water uptake can be lethal for plants. Nitrogen is necessary for plants, but too much all at once is a bad combo. Enter the dung consuming roaches and they prove a useful link in the process. By eating up all that nitrogen (roaches love it) they lower the intensity of the nitrogen input into the system. The nitrogen does not go away, it simply trickles into the system at a slower pace via the roaches crapping or the dead bodies of the roaches (and whatever other decomposers you want to insert- snails, termites, worms, fungi, bacteria etc etc).

Campos-Arceiz. (c)

Another interesting idea that the paper promotes, and an idea that converges with some of my own speuculations, is that the intestinal flora that allow digestion of cellulose and lignocellulose has essentially been conserved from dinosaur to cockroach to termite. That is dino dung eating roaches picked up the dino intestinal flora from eating dino dung. These roaches, the Blattulidae, passed on the flora to their descendant the wood roaches (Cryptocercus) and termites. With the extinction of the dinosaurian herbivore megafauna, the Blattulidae bit the dust as well. Wood roaches and termites, not as intrinsically dependent on dino dung as the Blattulidae, carried on to their present state. So, by extension, if you have a problem with termite damage in your building- maybe you can blame dinosaurs.

Now for me this is an endlessly fascinating idea. As you may tell I am very interested in the grey space between things and drawing a seamless narrative from deep time into the present. This idea encapsulates both of those interests of mine. If you came to this blog hoping to hear about phylogenetic arguments, you came to the wrong blog.

On a related note, I have in the past suggested that many dinosaurs took advantage of detrital food chains. In a Mesozoic world full of highly fibrous plants with a slow release of energy it may have proved useful to utilize the external stomach of detritivory. Ankylosaurids I am especially suspicious of-  having dubbed them the "mobile compost bins of the Mesozoic". So I am going to leave you with a photo below of a nile crocodile eating elephant dung to consume all the dung beetles within it. It should not be too hard for you to follow my train of thought here.

Nile croc eating dung for insect prey. Here.

Cheers!!

Pertinencia

Campos-Arceiz, Ahimsa. (2009) Shit Happens (to be Useful)! Use of Elephant Dung as Habitat by Amphibians. Biotropica 41(4): 406-407

Vršanský P, van de Kamp T, Azar D, Prokin A, Vidlička L, et al. (2013) Cockroaches Probably Cleaned Up after Dinosaurs. PLoS ONE 8(12): e80560. doi:10.1371/journal.pone.0080560

Campos-Arceiz. (c)

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Urine For Some Trouble... Renal Wars

Anyone who has kept or lived with cats is probably all too familiar with one of the more unfortunate byproducts of Felis domesticus- cat piss. Yes it is bad enough that they always have to deposit it in the corner of the litter box, but the toxic aroma of all that nitrogenous waste gets really bad if it soaks into your carpet, wood, mattress, or favorite t-shirt. That cat piss smells so bad and that we are so overwhelmingly repulsed by it is a natural byproduct of a protein rich diet and that nitrogenous waste is toxic to life.

It goes something like this: Nitrogen derived from proteins and nucleic acids are continually being broken down during an animals life. Although some of this nitrogen is reused, the vast majority of it enters the bloodstream as waste where it is bound up with hydrogen to form ammonia which is filtered by the kidneys and formed into a less toxic but highly water soluble product urea. And finally what you see in your toilet or what your cat deposits (hopefully) in the litter box is urine. Urine is actually a mixture of water, urea, sugars, elements and various other compounds. But urine in mammals is generally mainly water due to the water solubility of urea. Here is a more in depth look at the physiology of mammal urinary systems.

I also keep a corn snake as a pet, and although it also ingests a high protein diet, I can't really say that I have the same problem with snake piss as I do with the cat piss. In fact I can't really say that my snake pisses at all. This is due to the fact that reptiles and birds have an entirely different urinary physiology than mammals. They do not produce urea or urine as a final product, instead they produce uric acid. 

In the pic above of some chicken crap you can see the uric acid as the white chalky material on the right. And of course bird crap is familiar to anyone who has looked under bird nests or bird roosts. And if you have had the misfortune of living in an area full of seagulls (like me) or other birds you likely have at least once in your life had the misfortune of getting crapped on by a bird. But, given that bird crap is a composite of ingested waste as well as uric acid it is probably more accurate to say that you were crapped and pissed on by a bird simultaneously- double jeopardy. Here is a good write up of the physiology.

Now anyone who has suffered from gout may know that uric acid is not very water soluble so when uric acid is voided by reptiles and birds it comes out as a viscous white, pasty semi-fluid or even solid "chalk". But while gout in humans is generally seen as a product of western diet/ with  

genetic input which results in high uric acid recrystallization and subsequent inflammatory response- reptiles and birds actively synthesize uric acid from urea. And although uric acid production is more energetically costly than urea production- animals that produce uric acid have the advantage of voiding less water with their waste. The water saving capacity of animals that produce uric acid, as well as the water losing nature of urea production, has profound affect on the ecology, behavior, and activity patterns of these animals. In water limited environments this is especially pronounced. Indeed, if one lives in an arid or semi-arid environment one need only take a short day hike to realize this. For every mammal that you spot out and about during the day, you will probably see several dozen birds and reptiles. Its not that the mammals are not there, as in most cases rodents are probably predominant vertebrates- they are just in hiding to avoid excessive water loss.

And if we invoke the en vogue methodology of phylogenetic bracketing it is immediately apparent that dinosaurs, nestled nicely between reptiles and birds, almost certainly produced uric acid and not urea as their chief method to void nitrogenous wastes. And just as the urinary physiology of modern animals affects their ecology, behavior, and habits so it must have been for dinosaurs during the Mesozoic.

The Mesozoic was characterized by higher temperatures and higher aridity on average than the Cenozoic. Many of the most notable dinosaur formations, such as the Nemegt and the Morrison,  are characterized best as semi-arid or desert. And even in wetter areas, a monsoonal regime with a pronounced dry season , seems to have been predominant. An ability to save and conserve water by producing uric acid would have provided pronounced benefits for dinosaurs, especially for the mega-herbivores who may have had to travel far and wide for vegetation. A sixty tonne Argentinosaurus that can recoup much of the water it ingests from its diet as well as go for several weeks without drinking, has a distinct advantage over African bush elephants (Loxodonta africana), which are dependent on the accumulated knowledge of matriarch females and must drink at least every 2-3 days up to 190 L/ 50 gal of water.

If we extend out this argument that dinosaurs were less be-holden to water resources than mammals I believe we can make some inferences about how this difference affected the entire ecosystem. 

While the negative effects of livestock on riparian habitats is well documented large mammalian herbivores can also rapidly deplete vegetation around water holes even in natural settings. Just look at any picture of an east African watering hole during the dry season. The large mammals can not travel very far from the water source and the surrounding vegetation is both consumed and trampled. There literally is not a a riparian zone or any vegetative zone to speak of adjacent to the water. 

Everything is either consumed or shredded by sharp mammalian hooves. East Africa with its bevy of large megafauna naturally invokes comparisons to dinosaur ecosystems and this idea of the dinosaur watering hole is well entrenched in dino art/culture. Just check out the pic below from Reign of Dinosaurs.

But such scenes were the exception not the rule in my opinion. In addition to their superior renal physiology which did not constrain dinosaurs to water to the extent of large mammalian herbivores, several other factors are suggestive of relatively lessened burden on riparian vegetation. Large herbivorous mammals in xeric habitats got it rough. Large herbivorous mammalian mothers in such habitats got it even rougher. They absolutely need to drink to keep lactating and feeding their offspring. Mother dinos were under no such constraint. 

Dino feet, unlike the sharp/hard hooves of the majority of mammal herbivores were relatively soft and padded. They would not dig into the substrate damaging biological soil crusts and roots. The affect of feet upon soil, often overlooked, can dramatically alter the landscape when the actions are repeated ad nauseum.  

Dinosaur reproductive biology differs dramatically from large mammalian. Unlike say a herd of wildebeest in which the majority of individuals are fully grown- in any dinosaur population the population would be skewed towards smaller more immature individuals. So although dinos were absolutely larger than mammals, they were on average a lot smaller than generally portrayed. A good window into this would be to look at the size range in a population of crocodiles. Very few large adults, numerous subadults, and abundant young/hatchling. Herds of fully mature dinosaurs were likely few and far between, the vast majority of dinos were immatures and their effect on water resources was likewise smaller.

All of this is not to suggest that dinosaurs did not consume and eat riparian vegetation. In fact quite the opposite in my opinion, there is much to suggest that dinosaurs, especially in arid regions, exploited riparian vegetation. But it is through their urinary physiology, repro strategy, foot design, and movement across the landscape that water resources were not stressed and therefore the water table itself was kept higher providing the riparian vegetation that benefited all.

Cheers!!!

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Cowboys and Dinosaurs

For some reason cowboys and dinosaurs always seem to mesh together well for me. Maybe it is their shared history of the American west. Indeed when Triceratops skulls started popping up in the west they were believed to be an extinct form of giant buffalo. Maybe it is an enduring boyhood coupling of two of the most iconic features of childhood. Maybe the self reliant, rough and tumble cowboy persona seems the best fit for dealing with the unruly archosaurs if humanity was suddenly thrown back in the mix with dinos. Whatever the case dinosaurs and cowboys are an exciting cultural mash-up and one that has been explored before such as in the Harryhausen vehicle The Valley of Gwangi.

Valley of Gwangi 1969

Certainly a modern  re-imagining of the dinosaur cowboy crossover would be a lot of fun-  it could not be any worse than the recent Cowboys v Aliens. And perhaps the coupling of dinosaurs and cowboys is more appropriate, ecologically speaking, than is apparent at first glance...

The American cowboy mythos, perpetuated as a truly "Americana" cultural hero, is a bit of a bastard of spanish vaquero culture and American westward frontiersmen. It is a construct maintained in the hearts and minds of Americans through iconographic films, a fetishization of cowboy style, and cultural inertia. But at the heart of cowboy celebration and appropriation is a weak link- the cowboys namesake itself, cows or Bos taurus- the domestic cattle. You see trying to run cattle in the arid west is, ecologically speaking, trying to fit the proverbial square peg into a round hole. They just don't fit.

Ask yourself this- which state runs the most cattle? If you answered Texas you are right- but this state is misleading for Texas is the largest state in the lower 48. Check out this if you wanna see the real breakdown of where Americas cattle are at. Although not the most up to date (2007) this chart does reveal some interesting trends. What you really want to look at is the number of beef cattle per square mile of farm. What you see here is that only three western states break into the top twenty for beef cattle per square mile of farm, Nebraska (16), Nevada (18) and Texas (19). Western states that have a long tradition of free range cattle (often on public lands) such as Arizona (48), New Mexico (43), Montana (27) and Oregon (21) score even lower. But what really should jump out at you are the states that lead in beef cattle per square mile. The top ten in order are Tennessee, Florida, Virginia, Kentucky, Hawaii, Alabama, Missouri, Arkansas, Louisiana, and Oklahoma. What you should notice is that for the top ten most productive cattle producing lands- 9 of them are southern states but all of them are wet!!!

Cattle in the desert? You are doing it wrong!!!

It may be a little discomforting to western states and to our cherished notion of cattle belonging in the arid west but the data do not lie. Cattle do not thrive in the arid west but do marvelously in the humid south. How to explain this data? We need to look at the species from which cattle are descended, the now extinct Auroch cattle- Bos primigenius.

The last wild auroch died in floodplain forests like the one to the left during the 18th century. As you can see this habitat is dramatically different from the Arizona desert scrub of the cattle in the picture above. How did these bovines behave? By all historical accounts they were more agile, taller, more aggressive, and possibly more stealthy than their domesticated descendants. I would not want to attempt cow tipping one of these brutes.

Auroch

Tracking the wild cattle of Sapelo Island may give us our best glimpse of how Auroch behaved and their preferred habitat. On this Georgia Barrier Island wild cattle have colonized the island and, like feral pigs, reverted back to the ways of their wild ancestors.  Paleontologist/ichnologist (footprints) Tony Martin paints a picture of a stealthy, cryptic, salt marsh foraging beast that is rarely seen and utilizes both maritime forest and salt marsh habitats fully. When encountered on open ground these cattle quietly slip back into the cover of the forest. This image of cattle repatriated to their rightful habitat contrasts sharply with our traditional view of cattle as slow, insipid, docile and dumb. Instead they are robust, vibrant Pleistocene survivors most at home in wet/well covered habitats- not open arid plains. Below is a picture of a lone male bull on Sapelo Island that captures the wild nature of these cattle.

(c) Anthony Martin

Of course when you put cattle on arid lands they do survive- but at what costs to the land?

Well one could go on and on on this tangent but   I would rather outsource my arguments. Look at this photoessay on the degradation of riparian habitats, loss of biodiversity and ultimately loss of cattle to drought when cattle are raised in arid climates. If you want to go further read Welfare Ranching. If you come across some pseudo-scientific arguments that heavy grazing is good for the land, usually it goes under the name of holistic management don't buy it. Cattle raised on arid lands are not suitable replacements for buffalo because cattle are not mobile like buffalo. Buffalo, or bison, are constantly on the move and do not overgraze- ideally they are also in the presence of wolves and bears which keep 'em moving as well. And in southwestern deserts bison did not occur in any great numbers or were absent entirely.

Redband trout stream degraded by cattle. Magpie creek ID

Left is private land right is public overgrazed land Sequoia Nat Park CA

It is a little ironic that many of the traits we admire in "cowboy mythos" persona stem from the very difficult conditions of raising cattle in arid lands. Perseverance, hard work, grit determination and gumption would be traits absolutely necessary when trying to raise a water dependent domesticate in an arid land full of predators, droughts, and generally hostile conditions. Right now the wolf and grizzly bear are making inroads on lands they were extirpated from. Americans must decide if they want healthy intact ecosystems with top predators or repeat our mistakes if they want to maintain the cherished cowboy mythos which is more and more apparent as a mirage in the desert.

Oh yeah and what was that I mentioned earlier about dinosaurs being better suited ecologically for ranching by cowboys on arid lands than cattle? Well like reptiles and birds dinosaurs produced uric acid    which is less water soluble than the urine which mammals produce. This does not dehydrate organisms who produce uric acid as readily as those who produce urine. Essentially this is why you can go for a hike on a relatively warm day in an arid area and see plenty of reptiles and birds- but all the mammals are outta sight. Dinosaurs were essentially preadapted to desert/arid conditions. Having to drink less than large mammals large dinosaurs would not have had to make as many trips to drink and therefore had less of an impact on riparian vegetation communities. Another argument of mine in support of my theory that dino herbivores were less harsh on vegetative communities than herbivorous mammals...but I digress.

What type of dinosaur would be the ideal type for ranching in the arid west? Well it would be able to survive on poor quality fodder, reproduce quickly, have an efficient food to flesh turnover, not have sharp feet that erode thin soils, and be able to either defend itself or escape from large predators. And this dinosaur still exists- its called an ostrich.

Cheers!!!