Time For the Giant Heron Spinosaurid Analogy to Bite the Dust Part 2: Getting More Than Just Your Feet Wet

Now first things first I want to clarify why I am writing these posts and how dealing with spinosaurid paleobiology is a veritable land mine. In my last post I echoed the sentiment of a poster for #2014SVP (Anduza, Danny Museum of the Rockies) that the wading "heron" analogy is not the best analogy of all possible tetrapod analogies for spinosaurids. As Dr. Thomas Holtz pointed out in the comments this is a "strawman argument" because no one has seriously suggested that spinosaurids hunted in the manner of herons which rely on a rapid acceleration of a sigmoidally orientated head and neck after visual detection of prey. And this is true: it is well known that spinosaurids had overall straighter and stiffer necks than let's say a fairly basal theropod like Coelophysis. Where the heron analogy works is that it provides a blueprint for an animal that wades in after prey. 

However, and this is a big however, spinosaurids - perhaps more than any other group of extinct animals save tyrannosaurids - lie squarely at the intersection of public and professional interest and interpretation. We saw this very interesting phenomena just occur - #Spinogate2014 - that illuminated an interesting socio-scientific-internet experiment. A radical new stance for Spinosaurus asserted, questions raised and doubt sown on the internet, and a very loud and vocal internet cadre of anti-Ibrahim et. al.  proponents sallied forth. In fairness some of the Ibrahim et. al. critics are acting in good accordance with sound scientific scepticism and legitimate concerns. But one can not deny that a social aspect - bipedal loyalists, "bipedalists" if you will - are really hellbent on keeping the JP3 Spinosaur alive and well. And continue to do so even after Ibrahim et. al. discounted the criticisms.

But this post is not about Spinosaurus or Spinogate it is about how an analogy for the group as a whole - the "giant heron" model - has been seized upon by the lay public as a point of reference for the group to the point where, I will argue, the "heron" model is providing a disservice. And the manner in which the "heron analogy' has gone too far is not wholly from paleontologists - but from how these animals are portrayed in artistic and especially video reconstructions. Unfortunately even though the "heron" analogy was useful in the broadest sense, popular reconstructions have run rampant with the "giant heron" model dominating. This is a hindrance.

Spinosaurus (c) BBC

Spinosaurus (c) Todd Marshall

Baronyx (c) Mark Witton

 Suchomimus w/Kryptops (c) Julius Csotonyi

As you can see in these assorted and very beautifully rendered pictures of various spinosaurids there are echoes of the "giant heron" analogy in all of them. In none of the pictures do we see the spinosaurid body largely submerged. The BBC pic and the Witton pic (btw check out Witton's great deal on art, remember the holidays are coming and we should support independent paleo projects) suggest an emphasis on visual detection a la herons.  But the lack of strong adaptation for binocular vision and emphasis on sensory pits speak against this. Unfortunately even the new NOVA doc on Spinosaurus depicts it fishing heron style.  The Marshall pic shows the Spinosaurus taking advantage of its sensory pits but the choice of putting the animal on the bank of the water instead of in the water speak to a larger problem in spinosaurid reconstruction - A PERVASIVE FEAR OF GETTING WET. The Csotonyi illustration is awesome, but again conservative in term of water depth (book here).

Seriously, for a group of animals that we can say exploited aquatic resources with strong confidence, there is a strong reluctance to actually put the whole animal in the water (where its sensory adaptations and foraging options make much more sense I will argue anyways). The above images have a profound impact on how these animals are imagined by laypeople and professionals alike. These representations that invoke the heron model to various degrees need to be dialed back a few degrees in my estimation. And here I want to provide some nuance to my argument. It is not that I am suggesting scenes like the above did not ever happen. That a spinosaurid, at one time during the long span of time such creatures existed, happened to be standing on a river bank and coincidentally a large fish just happened to swim by in shallow water and that the spinosaurid was able to lunge at it in a heron like manner and catch it - this most certainly happened. What I am suggesting is that the representation of such a moment and style of predation is disproportionate to the amount of time such a style of predation actually occurred. 

More Than One Way To Be A Wading Aquatic Predator

Grey Heron & Painted Stork Feeding (c) Lau SY

What do they say?... a picture is worth a thousand words. Above you see the very obvious difference in feeding between a wading grey heron and a painted stork. The heron relying on slow, cryptic stalking and the stork employing tactile feeding - sweeping its sensitive bill through the water and feeling out prey. Below is another video this time of a wood stork and various heron illuminating the different feeding strategies - also note that the wood stork will intentionally disturb the substrate to flush out prey.

                                     

Hopefully you probably figured out where I am going with this. Various wading storks that employ a tactile method of foraging with their sensitive jaws immersed in the water offer more utility as an analogy to spinosaurid foraging behavior than visually dependent herons. A slight shift in perspective is all I am suggesting. In light of their downward sloping profile, sensory pit equipped snout, and lack of binocular vision this view of spinosaurids highlighting a foraging behavior dependent on tactile foraging is of some use and should be taken into consideration for popular reconstructions of these animals. You can see that Marshall almost got it right in the picture above where he at least got the snout in the water, but he did not want to get the spinos feet wet. Sheesh just put the whole animal in water and it makes much more sense.

Which brings me to my next point, how deep of water to put these animals in. Anywhere on the spectrum of foraging in drying up pools all the way to completely submerged in my opinion. In light of the preponderance of illustrations/animations depicting them as seemingly limited to the shallows I am definitely for more depictions of them in deeper water. Even the new spino doc used this depiction of shallow water heron foraging. Furthermore, all things being equal, big fish are going to seek refuge in deeper water especially in predator rich environments. All these illustrations of spinosaurids in ankle deep water and a 200 pound lungfish or coelecanth just sort of nonchalantly offering itself up for grabs in the shallows.... yeah, I don't really buy it. Its insulting to fishdom and seemingly outside the bounds of Darwinian evolution. If fish are being predated upon fish will seek refuge in deeper water. And if you are a big spinosaurid trying to stalk a fish visually - if you can see it chances are it can see you and evade you. 

So how did a spinosaurid go about hunting in deeper waters? Well first and foremost for spinosaurids to be successful they want as many things in their advantage as possible. And that means murkiness. If you can't see me that means you might not swim away in time before my snout detects you. This might mean foraging at night, when fish although not "sleeping" do go into sort of rest phase and of course vision is limiting. Also many waters are naturally murky or turbid, especially in the tropics. Illustrations of normally murky tidal and tropical rivers intentionally clean things up a bit but that is simply an artifact of allowing the viewer to see what is under the water. Tides and storms can also disturb the water creating turbidity. Other animals, dinosaurs frolicking, and the large fish themselves rooting around in the mud will create turbid conditions. Remember the "river of giants" and also that lungfish are known feeders of benthic prey and are rooting around all the time. And then finally the spinosaurid itself; if conditions are just a little too clear; if prey is hiding in vegetation or in the mud; just use your big feet to stir things up a bit. Like a 10, 000 pound wood stork. After that it is just a matter of slowly cruising the water until your sensitive snout and/or gaff clawed hands bump into something tasty.  

Which is pretty much what I am trying to illustrate above where a Suchomimus tenerensis is intentionally stirring up the sediment in order to flush prey ( a poor lungfish of course) and cloak itself as well. Notice I draped loads of barbs or "whisker" type integument over the animal in order to make the whole animal sensitive all over. Hey why not, we are in a bit of an integument free for all for better or worse.

But I should also add that when fish were concentrated in drying, shallow pools or pushed through topographic bottlenecks due to tides, migrations, spawings any self respecting spinosaurid in the area is going to hone in on that area to forage as opposed to the method depicted above. But such confluences of events that make fish so easily scooped up are usually few and far between.

Marabou Stork w/trapped catfish (c) Jonathan & Angela Scott

I should also note that there are some artists that have begun depicting spinosaurids mostly or even completely submerged. Brian Engh's depiction comes to mind (in fairness this Spinosaurus was made before the new reconstruction). He also has got the arms and claws outstretched feeling for possible prey and increasing its feeding envelope. His depiction is suggestive of an animal dependent on tactile senses. Check out his awesome blog Don't Mess With Dinosaurs and remember it for the holidays because he has some pretty cool merch and we all need to support independent paleo/monster creations if we want more of 'em!!

*Update 11/8/14 Andrea Cau of Theropoda has also been beating the drum for putting more of spinosaurid anatomy immersed in the water. Anatomically citing a body design pitching forward and down - an anteroventrally inclined head partially immersed in the water has depicted in the below schematic. In the comments below Andrea informed me that Simone Maganuco brought up the "stork analogy" 5 years ago in a conversation - so credit where credit is due.

Why the "heron" model has dominated the view of these animals over others such as "storks" is a question worth asking. Maybe many view herons and storks as sort of "samey" while in fact their foraging method is quite dissimilar. It is worth speculating that dinosaur thought and concept is still largely dominated by North American and European peoples. Where, especially in America, there are a lot more visual orientated heron type aquatic waders than tactile stork type aquatic waders. I believe we pretty much just have the wood stork. Europe is known for storks but primarily they have white storks which often feed in wet meadows and fields as opposed to in the water. But go down to an east African river or spend a week on the Pantanal and you will see relatively more of all types of waders - but especially stork style tactile waders.

So in conclusion more "stork" and less "heron". But actually I like "storkodile" the best because it gives reference to the tooth studded jaws, large size, and opportunistic nature of these beasties. Eat your heart out Crocosaurus.

Cheers!!

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Surf OR Turf: Can A Terrestrial/Aquatic Switchitter Really Exist?

Following on the heels of my Spinosaurus post I have been thinking about the aquatic/amphibious thing a lot lately and looking at the various levels of engagement in either realm animals partake in and have come to the conclusion that, at least for a terrestrial limbed tetrapod, being equally competent in the land and water at a high level is pretty darn hard to do if not impossible. (In the title I used the word switchitter to refer primarily to baseball players that are equally competent as hitters with both arms leading in case you miss the translation)

As a qualifier I stated this animal has to have limbs as numerous snakes are great in both realms - what comes to mind are garter snakes. I also stated that the animal has to be terrestrial i.e. not a bird as the dipper bird makes for a good case of being equally competent in both realms.

CA red-sided garter snake

I should also qualify my statement by clarifying what I mean when I say "equally competent". I would consider a predator that can both run down prey and swim down prey at high levels "equally competent".  Not just any prey though - this animals must outpace the fastest prey items in both respective realms. Or a prey species that could elude the fastest predators in either realm. Such an animal would be able to approach the cursorial capacity of canids or felids on land and in the water be on par with the fastest scromboids and dolphins.

For sure there are plenty of animals that hunt down and kill prey in both realms - bears, lots of mustelids, monitors, crocs, frogs - but when you really look at these animals they generally are geared primarily to one environment or the other. And in many cases they are utilizing stealth or elements of topography to gain an advantage. Remember I am looking for that animal that can capture - not by stealth but by speed - the quickest and most elusive prey in either environment with equal ease and efficiency.

I am starting to think there is very good reason such an animal does not exist nor could it ever.

The reason being that aquatic versus terrestrial movement asks the limbs to do very different things. Aquatic propulsion will tend to select for shorter and stouter limbs. Shorter and stouter limbs, being closer to the body, will create less drag than longer limbs during the recovery stroke. So if speed is something of importance in amphibious tetrapods shorter and stouter limbs will be selected for. This is seen very well in skeletons of otters. In addition a long torso works a bit like a hull of a boat to help cut through and displace water. Just look at the unique proportions of Michael Phelps - long arms and torso, but relatively short (read powerful) legs and flexible flipper like feet.

Now contrast this with what it takes to be an excellent cursor ; long legs, especially the lower leg elements; a deep thoracic cavity to house large heart and lungs; and compact torso for quick maneuvering. Seen against what is called for to be an efficient amphibious tetrapod it quickly becomes apparent that the two forms of locomotion are diametrically opposed. And also why Phelps, although a world class swimmer, would make a relatively poor runner. Compare and contrast the dimensions of Phelps with the ideal body dimensions of a runner and you will see that humans mirror the pattern seen in animals.

So although we have lots of examples of animals that dominate the aquatic and terrestrial realm in fiction - for example Dinocroc, Godzilla, Cloverfield Monster, Beast From 20,000 Fathoms, and the semi-mythologized JP 3 Spinosaurus - being equally proficient at a high level in both realms I don't see happening for a tetrapod.

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Planet Predator II: Kem Kem

(C) Troco. One of my fave recent images, read more about it here

Sometimes specific geologic formations take on a special, almost mythical, aspect to them. The Kem Kem beds of Morocco and Algeria definitely have that "special" quality to them. Documenting a roughly 10 million year span of the Cenomanian in the early Late Cretaceous, the formation represents several transgressive and regressive cycles of a coastal system on the southern Tethys Sea. Generally referred to as a "deltaic" environment, the Kem Kem contained a variety of environments within it- including offshore rudistid reefs, mangrove type forests, tidal rivers, floodplains and estuaries. But everywhere water was a dominant feature. The water was not necessarily all from precipitation- climatic models for this time predict arid conditions towards the equator- and rainfall when it occurred was episodic and monsonal. Picture the Kem Kem more like Arnhem land in the Northern Territory of Australia than the Amazon delta. No, the large amount of water was due to a very low gradient of land approaching high epicontintental seas.

Arnhem Land, Northern Australia

Ocean bound water slowed down and intermingled with waters from the Tethys creating a massive land sea interface on a continental scale- the magnitude of which we do not see today- perhaps the Sunderbans of India come close.

Sunderbans, India

Of course given this immense land/sea mosaic we would expect mangrove forests to dominate- except mangrove trees, which are angiosperms, had not yet evolved. Instead we had forests of Weichselia reticulata, a halophytic (salt tolerant) tree fern that was likely the dominant member of the intertidal floral community. Also common would have been various halophytic members of cheirolepidaceae- an extinct order of diverse coniferous plants- such as Frenelopsis. Herbaceous ferns, horsetails, and charyphytic algae rounded out the plant community.

Weichselia reticulata

Frenelopsis sp.

Thriving amongst these various watery microhabitats was a diverse assortment of fish. Amongst the more notable fish were a variety of medium to large sharks, coelecanths, bowfins, elasmobranchs, tarpons, lungfish, and even planktivorous paddlefish types. Adding to the diversity was a variety of marine reptiles, pterosaurs, turtles, crocodiles, snakes, and lizards. In a rather extensive review of the Kem Kem beds distinct faunas emerged relating to sea level and showing several biogeographic affinities to other marine/terrestrial provinces.

(c) Brian Engh

Of course what attracts the most attention, rightfully so, is the dinosaur remains from the Kem Kem. While several sauropods were present and ornithopod remains are sporadic as well what the Kem Kem does offer is a rather large assortment of multi-tonne theropods. And these guys were literally "whale sized". 

Kem Kem Theropods

So let's disregard the two smaller grey theropods- they are "problematic" to say the least. That still leaves you with the respectably sized Deltadromeus (green), gargantuan Spinosaurus (black), massive Carcharodontosaurus (red) and... who is that guy in blue? The guy in blue represents a potentially even more massive, as yet unnamed, basal carcharodontosaurid represented by basically a chunk of its braincase. Read about it here.

So whaaaa... how, ecologically speaking, did this suite of massive predators coexist? Especially in light of the fact that herbivorous dinos in this ecosystem appear to be less diverse/abundant?  (I don't buy completely the idea of "sampling bias" that some argue- although some truth to it is probable it does not erase the fact of four very large predators) Especially seen from the perspective of large mammal dominated ecosystems the idea of four contemporaneous multi-tonne predaceous dinos seem preposterous...

Well let's look at the ecosystem from the perspective of the most dominant feature present- water. Modern mangrove ecosystems are very productive places. But even in modern mangroves we don't really see anything analogous to say a seven tonne spinosaur tromping about. What was different in these areas from the present? Let's look a little lower on the food chain.

What really sets the Kem Kem apart is the abundance, diversity, and wealth of relatively large fish. If we look at modern large coastal fish we can draw some rough parallels. Many coastal fish do not typically stay in a single microhabitat. Barramundi, able to tolerate a variety of salinities (euryhaline), is known to descend downstream from its inland rivers into estuaries/tidal flats to breed.

Barramundi sp.

Tarpons (genus Megalops) are also euryhaline except that the fry are born in open ocean and then move in progressively closer to shore and sometimes into fresh water as they mature.

Tarpon (Megalops)

And then you have monstrous guys like the sawfish:

Some species can reach 23 feet in length and several tonnes in weight and all species can easily go between salt and freshwater habitats. And the Kem Kem is well known for having its own giant sawfish, the 8 meter long Onchopristis, often posited as a fave prey of spinos.

So what can be gleened from all this? Well all of the above fish are large, tropical, and are known for travelling between, among, and throughout various nearshore/offshore, estuarine, upriver habits in all kinds of combinations- seasonally or at will. The Kem Kem, with an even greater variety of large- most probably euryhaline- fish, was likely the focal point of energy transfer from both offshore, local, and inland water ecosystems. Combined minimal to nonexistent seasonal change, high epicontinental seas, and abundant primary productivity and you have a fish monger's dream.

Russel (2009) suggested fish were the basis for the food chain- culminating in the large predaceous theropods. I tend to agree with this statement. Although the case for spinos being sushi lovers is pretty iron-clad at this point- what about the several large carch species as well as Deltadromeus? Well, just because they have no clear adaptations towards piscivory like spinos do does not mean that fish- or aquatic organisms in general- were not a mainstay of their diets. Plenty of large typically terrestrial predators habitually take advantage of water based prey- jaguars, various bears, wolves to name a few- especially during seasonal migrations or when drought/geography funnel water prey into easily caught situations. Various crocodile species are well known for predicting fish movements and taking advantage of specific sites where fish are easily caught. Theropods, by no means brainiacs, were at least as cunning as crocodiles at predicting spots to opportunistically take advantage of certain hard to get prey items.

And even if the large theropods aside from spinosaurs were clumsy fishers don't forget about stealing your fish dinner!!! Indeed this form of piscivorous kleptoparasitism is rampant in extant theropods.

Maybe carchs habitually followed spinos and stole their fish...Maybe the efficient fish hunting spinos were key to the success of the other theropods- much as wolves indirectly benefit bears and other scavengers through their kills.

Remember the first Planet Predator I did a while back about the seeming paradox of a predator dominated ecosystem? One of the keys to the sustainability of such an ecosystem with 80% predator biomass  is that the prey base rapidly replenishes itself to sustain the losses. Well in the Kem Kem we may actually have a parallel situation with numerous fish species- many possibly rapid reproducers- supporting a top heavy cast of large predatory fish, pterosaurs, crocodiles, and theropods. Did the biomass of theropods and other top predators approach 80%- probably not, especially in light of the fact that the dinos were probably endothermic- but there is every possibility that it was higher than you might expect in a typical terrestrial plant>herbivore>carnivore pyramid. Here's why.

1) Abundant, rapidly replenishing fish prey base. The low continental relief combined with high consistent temps, epicontinental seas and productive mangrove/deltaic environment doubtless supported large nurseries of small fish and an unparalleled diversity of large, euryhaline fish. These fish likely moved between all environments distributing nutrients accordingly.

2) The predators themselves. Take a look at the pic in the beginning of the post. What it depicts is a large adult carch that has used a natural rock formation on abeach to pin down three immature spinos and kill one of them. A very realistic example of intraguild predation. Then go review this article and read about intraguild predation in raptors and monitors. Its quite possible intraguild predation was par for the course in these theropods. The theropods, especially given their reproductive capacity and size envelop from hatchling to adult, were partially acting as their own prey base!

3) Nary a herbivorous dino around. Its quite possible that the theropod diversity and numbers in the Kem Kem, buoyed up by abundant aquatic resources, were an active deterrent to abundant herbivorous dinos- despite the abundance of plants in these mangrove/deltaic conditions. Much like we saw in Shark Bay where the mere presence of tiger sharks was inhibiting foraging efforts by dugongs, dolphins, and cormorants- the abundance of theropods in the Kem Kem was simply not worth the effort for dino herbivores. So how did the lack of dino herbivores bolster theropod numbers? Well if the dinos were not eating the plants that means inverts were taking up the slack- insects, millipedes, snails, crabs, shrimp. It also means more decaying vegetation- which enriches the waters further- more food for the fish. In either case- more bugs or more nutrient rich water- the fish are happier. The fish are happier- the theropods are happier as well. The exclusion of abundant dino herbivores allowed a feedback loop which benefitted fish and theropod.

4) Spinosaurus was key. Of the four large theropods Spinosaurus was the only one specialized towards piscivory (crocodile jaws, pressure sensing snout, gaff like hand claws). The other theropods were likely not as efficient fishers. They may have benefitted from the presence of spinosaurs by either stealing or scavenging their catches or using spinos directly as prey in lieu of herbivorous dinos. In either case spinos were the keystone species- the presence of which affected all others.

Of course I could be wrong. Perhaps carcharodontosaurids were true landlubbers and hated sushi. Perhaps the predator bias is actual and herbivorous dinos were more common and diverse than thought.   Perhaps spinosaurids didn't eat much fish at all- but swam offshore to feast on rudistid clams...perhaps

Nah, I like the idea of a theropod dino dominated habit thriving on fish/aquatic animals. Herbivorous dinosaurs need not even be involved in this food web. Aspects of this type of ecosystem are not unparalleled. If we look to pristine reefs with tremendous top predator biomass we see potentially analogous characteristics. And if we look at the type of exclusionary actions abundant large predators impose on other animals (tiger sharks in Shark Bay, wolves in Yellowstone) this situation in the Kem Kem is not without precedent.

And now just for fun...

I imagine the "food journal" of an adult Carcharodontosaurus in the Kem Kem would go something like this...

Today I woke up and sniffed out the remains of a 5 meter sawfish hauled to shore by some spinosaurs from the night before. Left a little hungry but slept the rest of the warm day. Woke up at 3 am to the sound of several large stomatosuchid crocodiles scooping up schools of fish in some shallow water.

I knew the bay they were in and I knew if I could approach from deeper water I could probably snag one. I did it! - several of the larger 6 meter guys got away but I snatched up a good 4 meter croc which slid down my gullet in one swallow... Slept for three days in a row, I have not seen a carch larger than myself (about 13 m and 4 tonnes) in several months so I feel pretty confident in napping whenever and wherever. Several years ago  at about 10 meters I had to watch out for larger carchs as well as large adult spinos- my left arm was pulled off by a big bruiser of a spino at about 16 meters, Now even these guys leave me alone and I routinely steal fish from spinos up to about 14 meters. I start making my way upriver as it was about this time last year that large schools of coelecanth congregated to breed in some pretty dense horsetail beds where the water is more fresh tasting (I do have salt glands to deal with drinking salty water). Even if I don't catch a fish chances are I could steal one from the abundant spinos that congregate for the fish- or even kill a small spino if given the chance. Along the way I cross some open dry terrain and smell my old nemesis Deltadromeus. Until I was about 8 meters long these guys really gave me a hard time.

Pretty much all of my clutch-mates were gobbled up by these annoying speed demons. They were always faster than me and I would generally back myself into a corner to avoid their annoying but not very strong bites. Now I hunt them whenever I get the chance. They tend to avoid the deeper waters and denser forests where their speed is neutralized but I have got them before- usually in the chaotic monsoon when I can sneak up on 'em undetected. I finally reach the horsetail beds after several days travel. Over the years I have refined my hunting technique for these strong, maneuverable fish. I know of one spring fed pool, clear and deep where they like to breed. Leading to it is a small channel, only about 3 meters across and surrounded by dense cycads on either side. I set up my ambush site here and catch the 50 kg fish pretty consistently over several days. On the third day I notice a sound I had not heard for several months- a herd of rebbachisaurus sauropods.

I used to hunt these guys all the time before I discovered its a lot easier to be a hooligan in the delta. Even a midsize one can pack a whallop with its tail and I've been injured by these brutes before. Never the less I launch into an attack on one about the same size as me and deliver a good slashing bite to its left front leg- making it limp. Soon it will fall behind the herd and I will have a good feed of sauropod- a meal I have not enjoyed in several months!!!

Cheers!!!

Mawsonia libyca, 3 meters long, next to modern coelecanth & girl

Pertinencia

Cau, A.; Dalla Vecchia, F.M.; Fabbri, M. (2011). "Evidence of a new carcharodontosaurid from the Upper Cretaceous of Morocco". Acta Palaeontologica Polonica, in press. doi: 10.4202/app.2011.0043.

Loeuff, et al. Vertebrate assemblages from the early Late Cretaceous of Morocco: an Overview. Journal of African Earth Sciences. 2010.

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