Model organisms are a staple of biology. They are taxa that are used to answer larger questions about that group as a whole, or some general biological problem. Model organisms are chosen for their ease of handling, cheap acquisition, generally “generic” structures, or all of the above. Every major class has a model organism to represent it. Just among vertebrates we have:
Mammals with mice (Mus musculus), dogs (Canis familiaris [or Canis lupus familiaris if you lean that way]), cats (Felis catus [or Felis sylvestris catus for the same reason as dogs]), guinea pigs (Cavia porcellus) and rhesus monkeys (Macaca mulatta).
Birds with chickens (Gallus gallus), pigeons (Columba livia), and zebrafinch (Taeniopygia guttata).
Ray finned fish with zebrafish (Danio rerio), swordtails (Xiphophorous) and cichlids (Cichlidae).
Amphibians with the African clawed frog (Xenopus laevis), and axolotol (Ambystoma mexicanum).
Reptiles with anoles (Anolis), fence lizards (Sceloporous), painted turtles (Chrysemys picta) and finally, the American Alligator (Alligator mississippiensis).
Alligators are relatively new to the model organism realm, but they have proven to be extremely informative. They seem to the be most even tempered of extant crocodylians, making them “more safe” for researchers to work with. Hatchlings start off as miniscule 68 gram (0.15 lbs) animals that later can grow to 363 kg (800 lbs) adults, passing through an enormous size range throughout ontogeny. This growth rate is very food dependent, making it possible to raise alligators almost as bonsai trees. Also, with their unique position on the organismal family tree, alligators are one of the closest living relatives to dinosaurs. Along with birds, they have the potential to help constrain our assumptions about dinosaurs; thus making them very popular subjects for paleontological research as well.
Today, alligators get to make one more stamp on human knowledge with the release of the 3D alligator project from the Holliday and Witmer labs.
Researchers from both labs went through the painstaking process of digitizing the skulls of an adult and a hatchling American alligator, and then digitally separated each bone. The result is a 3D model that can have each bone turned on and off at will. The neat thing is that both labs have made these data freely available for anyone to look at, and download as 3D pdfs, wirefusion models, and multiple movies.
So if one every wanted to know just how many bones make up a crocodylian skull, or how each bone aligns to each other, I highly recommend downloading the 3D pdfs of the adult and hatchling. Not only will one learn all the different bones that compose the skull, but by comparing hatchling to adult, one can see just how radically these bones change throughout ontogeny.
It’s neat, free, informative and reptilian. What more can one ask for. 🙂
Continuing my trend of “catching up,” an article in the November issue of Natural History magazine, talks about a new study in the Quarterly Review of Biology, that finds group nesting to be very common place among extant reptiles.
That study would be:
Doody, J.S., Freedberg, S., Keogh, J.S.? 2009. Communal Egg-Laying in Reptiles and Amphibians: Evolutionary Patterns and Hypotheses. Quart. Rev. Biol. Vol.84(3):229-252.
In the paper, Doody et al (no laughing) did a massive search through the herpetological literature (both technical journals, and hobbyist magazines) to look at instances of communal egg laying in reptiles and amphibians (herps). I’m not being hyperbolic here either, as the paper states:
In total, our assembled database was gathered from 290 different sources, including 176 different scientific journals, 72 books or book chapters, 29 unpublished reports, and 13 unpublished theses. We also have included a number of reliable personal communications from herpetologists.
What the authors found was that group gatherings of herps are vastly more common than previously believed. Group egg laying was found to be present in 345 reptile species. Now you might be thinking 345 really isn’t all that much for a group composed of some 8700 species.
Well then aren’t you a Debbie Downer?
Seriously though, the authors address this by mentioning:
Although the difficulty in locating nests hampers our ability to determine the actual frequency of communal egg-laying among species, we can better estimate this proportion by dividing the number of known communally egg-laying species by the total number of species, excluding those for which eggs have not been found. We conducted such a calculation for the three families of Australian lizards known to include multiple communally egg-laying speciesâ€”Gekkonidae, Pygopodidae, and Scincidaeâ€”as gleaned from the Encyclopedia of Australian Reptiles database (Greer 2004). Proportions of these lizard families known to lay communally were 4â€“9%, but, when we exclude species for which nests are not known, these values rise dramatically to 73â€“100%
The biggest take home message to get from Doody et al’s review, is just how much we don’t know about extant reptiles.
…the present review highlights our inadequate knowledge of the nests and/or eggs of reptiles. For instance, the eggs or nests are known in only 7% of Australian lizards of the three families that commonly lay communally (N = 411 oviparous spp.) (Greer 2004).The extent of this knowledge for Australian lizards is probably similar to that for reptile eggs on other continents, particularly South America, Africa, and Asia, where the reproductive habits of reptiles are poorly known. This is in stark contrast to other vertebrates such as birds, for which complete field guides to the eggs and nests are available for several continents
Indeed, just by doing the brief research run needed to compile this blog post, it was apparent that communalism is much more common in reptiles than anyone ever thought. However, because so many of these reports are either anecdotal, or buried in obscure journals, it is easy to miss all the many cases where it is known.
This discovery lead the authors to the inevitable follow up question of: “why?” What benefit do mothers gain by nesting communally?
Numerous hypotheses for why animals nest communally, have been proposed.
Saturated habitat (only so many suitable nest sites)
Sexual selection (choice of males that live in a particular area)
Artifact of grouping for other reasons
Attack abatement (easier to hide a bunch of eggs in one site, than in multiple sites. Less chance that your eggs will be the ones that are eaten).
Maternal Benefits (save time and energy finding a suitable nest site by “freeloading”)
Reproductive success (if the nest site worked once before…)
The authors reviewed all of these possible reasons for communal egg laying in herps. In the end, they found evidence for both the maternal benefits hypothesis, and the reproductive success hypothesis, though they felt a mixed model better explained things.
Sadly, though the authors mentioned how a lack of information on the natural history of most reptiles is largely responsible for this sudden revelation about their nesting behavior, they nevertheless make repeated mentions of how “social interactions are generally less complex in reptiles and amphibians than in other tetrapods” or how herp sociality forms “relatively simple systems“.
The reality is that the old view of simplistic “loner” reptiles that only come together to mate, is not accurate. This is especially true for parental care in reptiles.
The popular view (among the public, and the scientific community) is that reptiles are? “lay’em and leave’em” types when it comes to reproduction. Despite all the herpetological knowledge to the contrary that has been acquired in the past 50 years, it is still popular to spout the party line about reptiles being “uncaring parents.”
Zoologist Louis Somma took issue with this view of reptilian (in particular, chelonian and lepidosaurian) parenting. He conducted a literature search to see how often mentions of parental care in reptiles are recorded. In the end he wound up finding 1400 references to parental care in reptiles (Somma 2003)!
Somma’s survey covered various aspects of parental care. He found reported evidence of nest building and / or guarding in tortoises like Manouria emys (McKeown 1999), Gopherus agassizii (Barrett & Humphreys 1986) and 4 other species of chelonian.
Turning to lepidosaurs, Somma found parental behaviour to be present in 133 species of lizards and 102 species of snakes. Even a species of tuatara (Sphenodon punctatus) is known to guard its nests (Refsnider et al. 2009). Though these numbers appear small compared to the total amount of species that have been described; much like the Doody et al. paper, this is just based off of species whose nesting behaviours we do know. That these taxa all span a wide phylogenetic range, suggests that parental care is more commonplace than initially thought.
Nest guarding is usually a maternal trait, but some squamates exhibit nest guarding behaviour in both parents, such as some cobra and crotaline snakes (Manthey and Grossman 1997) , as well as tokay geckos (Zaworski 1987).
Not only active guarding of the nest, but actual brooding of the eggs is also commonly reported in squamates such as various python species (Harlow & Grigg 1984, Lourdais et al. 2007), and skinks (Hasegawa 1985, Somma & Fawcett 1989). Some species are even known to groom their newly hatched young (Somma 1987).
More interesting still are various reports and observations of parental feeding in some reptile species, such as the skink Eumeces obsoletus (Evans 1959), and the cordylid lizard Cordylus cataphractus (Branch 1998). Not to mention recent evidence of parental feeding in captive crocodylians.
This leads me to the only reptile group where parental care is well publicized: that of the 23 extant crocodylian species. I could, at this point, list references for parental care in crocodylians. However because this behaviour is so well documented for this group, it would seem unnecessary. It is? better to shed light on the many (MANY) examples of parental care in other reptile species. I also didn’t include related examples like placental evolution in the skink genus Mabuya, or instances of egg binding in captive reptile mothers; due to a lack of appropriate substrate to lay their eggs.
In the end, the paper by Doody et al. adds to a growing body of evidence which suggests that the “lay’em and leave’em” reptile species of the world, are the exceptions? and not the rule.
Next time: Biomechanics of running suggest “warm-blooded” dinosaurs. Or: why the aerobic capacity model needs to die already.
Barrett, S.L. & Humphrey, J.A. 1986. Agonistic Interactions Between Gopherus agassizii (Testudinidae)
and Heloderma suspectum (Helodermatidae). Southwestern Naturalist, 31: 261-263.
Branch, B.. 1998. Field Guide to Snakes and Other Reptiles of Southern Africa. Third revised edition. Sanibel Island: Ralph Curtis Books Publishing.
Doody, J.S., Freedberg, S., Keogh, J.S.? 2009. Communal Egg-Laying in Reptiles and Amphibians: Evolutionary Patterns and Hypotheses. Quart. Rev. Biol. Vol.84(3):229-252.
Evans, L.T. 1959. A Motion Picture Study of Maternal Behavior of the Lizard, Eumeces obsoletus Baird and Girard. Copeia, 1959: 103-110.
Harlow, P and Grigg, G. 1984. Shivering Thermogenesis in a Brooding Python, Python spilotes spilotes. Copeia. Vol.4:959?965.
Hasegawa, M. 1985. Effect of Brooding on Egg Mortality in the Lizard Eumeces okadae on Miyake-jima, Izu Islands, Japan. Copeia, 1985: 497-500.
Lourdais, O., Hoffman, T.C.M., DeNardo, D.F. 2007. Maternal Brooding in the Children’s Python (Antaresia childreni) Promotes Egg Water Balance. J. Comp. Physiol. B. Vol.177:560-577.
Manthey, U. and W. Grossman. 1997. Amphibein & Reptilien S?dostasiens. Natur und Tier Verlag, M?nster.
Mckeown, S. 1999. Nest Mounding and Egg Guarding of the Asian Forest Tortoise (Manouria emys). Reptiles, 7(9): 70-83.
Refsnider, J.M., Keall, S.N., Daugherty, C.H., & Nelson, N.J. 2009. Does nest-guarding in Female Tuatara (Sphenodon punctatus) Reduce Nest Destruction by Conspecific Females? Journal of Herpetology. vol.43(2):294-299.
Somma, L.A. 1987. Maternal Care of Neonates in the Prairie Skink, Eumeces septentrionalis. Great Basin Naturalist, 47: 536-537.
Somma, L.A. & Fawcett, J.D. 1989. Brooding Behaviour of the Prairie Skink, Eumeces septentrionalis, and its Relationship to the Hydric Environment of the Nest. Zoological Journal of the Linnean Society. Vol.95: 245-256.
Somma, L. 2003. parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey. Krieger Publishing Company. 174pgs. ISBN: 157524201X
Zaworksi, J.P. 1987. Egg Guarding Behavior by Male Gekko gecko. Bulletin of the Chicago Herpetological Society, 22: 193.
Two new papers have recently hit the journal circuit. Both of them involve using living crocodylians to gain a better understanding of paleo-life.
The first one comes from Denver Museum of Natural History paleontologist, Dr. Kenneth Carpenter:
Carpenter, K. 2009. Role of Lateral Body Bending in Crocodylian Track Making. Ichnos. Vol.16:202-207. doi:10.1080/10420940802686137.
The study used an adult Caiman sclerops (first use of a large adult reptile for a locomotion study; at least as far as I know) placed in a small room with two 30cm walls placed on either side of it. This restricted any lateral movement, and “funneled” the animal out the singular opening. At this opening, a camera was placed. It would photograph the animal as it left the room. The room itself, had a smoothed mud covering. This muddy floor would record the tracks of the C.sclerops as it walked by.? Several runs were done, and photographs were taken for each run.
This is the first study I have seen that gave a front view shot of an adult crocodylian as it walked along. As Carpenter mentioned in the paper:
This front view is in contrast to most photographic studies which only capture pro?le and top views….
Carpenter also mentioned the potential of there being an ontogenetic change in limb stance as animals move from hatchling to adult. This is something that I have hinted at previously Hatchling crocodylians seem to have weaker femoral adductors than adults. This is understandable given the greater weight that adult femora need to bear. This can result in a skewed view of crocodylian erect stance; with most authors tending to underestimate the degree of “parasagittality.”
That said, I was surprised to read that Carpenter had found the adult Caiman sclerops to have a hip adduction angle of approximately 65? from the horizontal. Judging from figure1B, the hindlimb appears to be much closer to the midline than the forelimb. Fig1D seems even closer to, if not 90?. It is important to point out that much of the hindlimb is blocked by the body in this shot, as the animal is fully laterally extended. A concurrent shot from behind would have been very useful here; as would an x-ray series of shots throughout the walk phase (for instance: see this long video of a Crocodylus acutus walk cycle. Pay special attention to the position of the femur).
Alas, that is not what the paper is about.
The paper is about how lateral movements during locomotion, have substantial effect on trackways. Dr. Carpenter points out how, despite the semi-erect stance of the forelimbs, the track evidence would suggest an animal with a much narrower (parasagittal?) stance. This has bearing on how prehistoric reptiles, in particular: quadrupedal dinosaurs, may have stood.
One might rightfully ask if we should expect dinosaurs to have had any lateral movement to their walking cycle at all. Carpenter points out that lateral body bending, though not quite as exaggerated as that of crocs, is present in most tetrapods. Birds seem to be the sole exception, with their extremely stiff thorax. However birds are also obligate bipeds, and the avian thorax is much shorter and stiffer than that of dinosaurs.
So it would seem to be a likely bet that quadrupedal dinosaurs likely exhibited some degree of lateral body bending.
Carpenter’s work rightfully asks us to caution reconstructions of stance based largely off of trackway evidence. A fine case study that the paper brings up, is ceratopians. This group, more than any other, has received considerable attention for how the forelimbs were oriented. Early work on ceratopians, favoured a hefty sprawl to the forelimbs (e.g.? Gilmore 1905, or Lull 1933). This was critically evaluated during the heyday of the dinosaur renaissance. Authors such as Bakker (1986), Paul and Christiansen (2000), instead favoured a fully erect stance. A large portion of the data supporting this assertion, was trackway based. The results of this study call into question that view. However this was not the first paper to have done so. Thompson and Holmes (2007) also questioned the “erect ceratopid” view, using a half scale model of a Chasmosaurus irvinensis forelimb. Their results come closer to the results from this paper. Though Thompson and Holmes felt that there was no real modern analogue to ceratopian forelimb mechanics.
In the end, Dr. Carpenter reminds future researchers of the importance in incorporating the entire animal when analyzing trackways.
The second paper comes from the Journal of Experimental Biology.
Owerkowicz, T., elsey, R.M. and Hicks, J.W. 2009. Atmopsheric Oxygen Level Affects Growth Trajectory, Cardiopulmonary Allometery and Metabolic Rate in the American Alligator (Alligator mississippiensis). J.Exp.Biol. Vol.212:1237-1247. doi:10.1242jeb.023945.
The authors embarked on a study of how previous paleo-atmospheric oxygen levels might have affected the lives of animals that would have been alive through these times. According to Owerkowicz et al, crocodylians were chosen because:
Given their phylogenetic position and highly conserved morphology throughout their evolutionary history, crocodilians are often thought to retain many characteristics of basal archosaurs.
I do take some issue with this, as prior reviews on crocodylomorph diversity (Naish 2001) coupled with many new discoveries ( Buckley et al 2000,? Clark et al 2004, Nobre & Carvalho 2006)? continually cast doubt on the old view that crocodylians have survived “unchanged” for some 200 million years. Nevertheless, the results of the study are both interesting, and relevant to reconstructions of how paleo-life would have adapted to these wildly different paleo-atmospheres.
Owerkowicz et al raised groups of hatchling American alligators (Alligator mississippiensis) under three different atmospheric conditions. A hypoxic (12% O2) condition reminiscent of paleo-atmospheric models for the late Triassic/Early Jurassic periods. Current atmospheric conditions (21% O2), and a hyperoxic (30% O2) condition reminiscent of paleo-atmospheric models for the Carboniferous and Permian periods.
The results were interesting, though not too surprising. As expected, hypoxic alligator hatchlings were smaller than their normal and hyperoxic counterparts. However, the degree of growth stunting is pretty surprising. Hypoxic hatchlings were about 12% shorter and 17% smaller than normal hatchlings.
Surprisingly, hatching time did not change under any conditions. This suggests a degree of “hard wired” embryological development inside the egg. In the case of the hypoxic hatchlings, they came out “almost done.” While all three groups had remnants of a yolk sac upon hatching, the hypoxic hatchlings actually had the yolk sac still protruding (normal and hyperoxic hatchlings just showed distended bellies). In some cases, the yolk sac was larger around than the hind legs, thus making movement clumsy and cumbersome.
Other interesting results from this study, included notable changes to the cardiopulmonary system. Hypoxic hatchling lungs were actually smaller than the lungs of normal hatchlings; which appears counterintuitive. The heart, meanwhile, showed distinct hypertrophy in hypoxic animals. The authors believe that lack of lung growth in hatchlings may have been due to the fact that lung function does not start until after hatchlings have hatched.? The heart, on the other hand, is hard at work circulating blood just as soon as it is formed; so it would have experienced the challenges of hypoxia at a very early stage.? Bolstering this hypothesis from the authors was the fact that three months after hatching, hypoxic alligators showed a distinct increase in lung growth rate (the lungs appeared to be “catching up” to the heart).? Hypoxic alligators showed shrunk livers as well. No real explanation for this was given, but it was mentioned that reduced liver mass seems to be a common trait in animals raised in hypoxic conditions. It appears to have some bearing on overall metabolic rate.
Hyperoxic hatchlings exhibited “typical” organ growth rates.? Where hyperoxic animals excelled was in breathing and metabolic rate.
Breathing rates were smaller in this group, while metabolism and growth rate were all larger. The explanation by the authors was that these hyperoxic animals were receiving such high amounts of oxygen in each breath, that they were actually hitting saturation at much shallower breaths; hence the shallow breathing. The higher metabolic rate is believed? due to a lack of right-left shunting in the crocodylian heart. This shunting is usually caused by low oxygen levels (like that experienced in diving), and tends to result in metabolic depression to conserve available oxygen stores.? Since these alligators lungs were constantly saturated with oxygen, right-left shunting never occurred, resulting in an elevated metabolism.
Incidentally, Owerkowicz et el give mention of a cardiac shunt known in embryological birds (via the ductus arteriosis). Though only analogous, one can’t help but wonder what this might have meant for all those dinosaurs that lie between these two groups.
Interestingly, hypoxic alligator hatchlings also showed a higher standard metabolic rate. Though these animals would voluntarily eat less than their normal and hyperoxic counterparts, their metabolism was more like hyperoxic hatchlings than they were normal hatchlings.? Owerkowicz et al believe the reason for the increased metabolism was due to the higher cost of breathing in these animals. Despite taking “normal” breaths, hypoxic hatchlings were taking in a larger tidal volume than their normal and hyperoxic siblings. The heart was also working harder to deliver enough oxygen to tissues.
Finally the authors give mention of growth rates in hyperoxic animals. Basically, it is faster. The authors mention that this might be caused by the persistently elevated metabolic rate, or perhaps from channeling saved energy from breathing (which is one of the main energetic costs in reptiles) into biomass.? It could be a mix of both, but I’m more inclined to think that it comes more from channeling energy reserves into other parts of the body. A high metabolism means nothing, if there is not enough free energy to go around. Just look at the hypoxic gators from this study. Despite their high metabolism, they grew slower than their peers.
The results of this study showed how modern animals can acclimate to different atmospheric conditions. They don’t show how animals would adapt and evolve in these conditions, but they do hint at the general directions, and help give us a clearer picture of what life was like millions of years ago.
Bakker, R. 1986. The Dinosaur Heresies. William Morrow. New York. ISBN: 0821756087, 978-0821756089 pps: 209-212.Buckley, G.A., Brochus, C.A., Krause, D.W., Pol.D. 2000. A Pug-Nosed Crocodyliform from the late Cretaceous of Madagascar. Nature. vol.405:941-944.
Clark.J.M., Xu, X., Forster, C.A., Wang, Y. 2004. A Middle Jurassic ‘Sphenosuchian’ from china and the Origin fo the Crocodylian Skull. Nature. Vol.430:1021-1024.
Gilmore, C.W. 1905. The Mounted Skeleton of Triceratops porosus.? Proceedings United States National Museum. Vol.29:433-435.
Lull, R.S. 1933. A Revision of the Ceratopsia, or Horned Dinosaurs. Memoirs of the Peabody Museum of Natural History. Vol.3:1-175.
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Nobre, P.N. and Carvalho, I.S. 2006. Adamantinasuchus navae: A New Gondwanan Crocodylomorpha (Mesoeucrocodylia) from the Late cretaceous of Brazil. Gondwana Research. Vol.10:370-378.
Paul, G.S., and Christiansen, P. 2000. Forelimb Posture in Neoceratopsian Dinosaurs: Implications for Gait and Locomotion. Paleobiology, 26(3):450-465.
Back in 2002, a short paper came out that commented on the observation that captive caimans would eat fruit left in their cage. When I initially read the paper, I found it interesting. In the end, though, I assumed this to just be a fairly anomalous incident.
As one can see, this observation has been filmed at least once.
So does this mean that crocodylians are not as completely carnivorous as once thought? It’s hard to say. All observations made so far have been from alligatorids (alligators and caimans). This might be an apomorphic trait to this group. Only more observations will say for sure.
Another option that Darren pointed out, is that this was a learned trait of these captive animals. In each case, observed animals were found to be sharing their enclosures with herbivorous animals (usually tortoises). This type of operant learning is rather rare, and would be amazing if found to be true.
However, as evidenced by the comments of St. Augustine Alligator Farm park director, John Brueggen, fruit eating has been observed in wild animals too; so this is not simply a case of bored captives.
Whatever the case, these observations do illustrate just how adaptaptable crocodylians are as a group.
The Telegraph had a report today on a unique event in Africa. Hal Brindley, an American Wildlife photographer, happened to be in the right place in the right time for this to happen. The shots come from Kruger national park.
Photo taken by Hal Brindley and destributed by Solo Syndication
The photos show a leopard (Panther pardus) taking out a Nile crocodile (Crocodylus niloticus). The croc is smaller than the leopard, but the results are no less shocking.
Leopards, and most big cats in general, don’t go after crocodiles as a normal routine. Not only do the felines run the risk of becoming the hunted, but even if they do win (like this leopard did), there is really not a lot of meat to go around. Much of it is locked away by the bony osteoderms.
The only time I have seen a leopard take out a crocodile, was in a case where both animals were slowly starving and dehydrating. The crocodile died first, and the leopard took advantage of the free meal.
As one can see in the photo, this leopard wasn’t exactly starving, so it is hard to imagine desperation being the cause of this attack. The article also mentioned that the leopard attacked the croc on its own turf. Both situations are very unique, and make the fact that the leopard won, all the more amazing.
There was no run time given for the battle, so I’m not sure how long it lasted. Judging from some of the shots though, it must have been intense.
Recently published in Proceedings of the Royal Society B, scientists in Brazil have found the remains of a prehistoric crocodyliforme that used to roam the oceans of the Paleocene.
The critter has been given the name: Guarinisuchus munizi, which translates out to: Muniz’s warrior crocodile. Despite the “crocodile” in its name, G.munizi was not that closely related to true crocodylians. It was more closely related to the giant pholidosaur Sarcosuchus imperator.
Close relative of Guarinisuchus munizi [left] was Sarcosuchus imperator [right]. Not true crocodiles.
The neat thing about the paper, was not so much the crocodyliforme itself. At 3 meters, G.munizi was small for a dyrosaurid. Rather, it is the implications of this find that are intriguing.
Dyrosaurids first appeared in the Late Cretaceous Period (Maastrichtian age) . During this time they were very scarce, and hard to find. They were shallow marine predators, and in the Cretaceous that niche was already filled by another group of animals: the mosasaurs.
These ancient sea lizards had one of the fastest diversification rates of any vertebrate group studied. They went from nothing to dozens of species with a cosmopolitan distribution and domination of many ecological niches. All of this occurred in the space of only 25 million years! That’s faster than mammal diversification, and faster than dinosaur diversification.
Mosasaurs were showing no signs of slowing down right up to the K/T asteroid event. After that, they disappeared.
That’s when the dyrosaurids started taking over.
Analysis of Guarinisuchus munizi material has found that it is more closely related to African taxa than its geographically closer relatives in North America. This suggests that dyrosaurids had crossed the Atlantic ocean from Africa sometime before the K/T event. After said event, the vacant niches left by the mosasaurs were quickly snatched up by these dyrosaurids, as they moved up North, and eventually, worldwide.
It is interesting to see how this group of animals was apparently held back during their earlier evolution. Yet, if they hadn’t been held back; if they had out-competed mosasaurs for the top spot in the food chain, then they wouldn’t have survived the K/T event.
It’s funny – and completely make believe – but it almost appears as if dyrosaurids were already setting themselves up to take over. It’s almost as if they knew…
They didn’t of course, but it’s fun to pretend that they did. >:)
They suggested that as the fish moved from polluted rivers into the Chambal, they ingested chemicals in their tissues. When the gharials eat the fish, these harmful substances pass into their systems.
One of the international vets who has been working on the case, Paolo Martelli, explained to the publication: “When cold temperatures came, the uric acid precipitated [separated into a fine suspension of solid particles] and began causing problems.
“So winter coupled with excess food could have made the gharials more susceptible to the toxin.”
We used electromyography on juvenile American alligators to test the hypothesis that the following muscles, which are known to play a role in respiration, are recruited for aquatic locomotion: M. diaphragmaticus, M. ischiopubis, M. rectus abdominis, M. intercostalis internus, and the M. transversus abdominis. We found no activity with locomotion in the transversus. The diaphragmaticus, ischiopubis, rectus abdominis and internal intercostals were active when the animals executed a head-down dive from a horizontal posture. Weights attached to the base of the tail resulted in greater electrical activity of diaphragmaticus, ischiopubis and rectus muscles than when weights were attached to the head, supporting a role of this musculature in locomotion. The diaphragmaticus and rectus abdominis were active unilaterally with rolling maneuvers. Although the function of these muscles in locomotion has previously been unrecognized, these data raise the possibility that the locomotor function arose when Crocodylomorpha assumed a semi-aquatic existence and that the musculoskeletal complex was secondarily recruited to supplement ventilation.
Scientists at the University of Utah have discovered the unique internal subtleties that allow crocodylians to sink, rise, pitch and roll; all without disturbing the water (much). It turns out that the main muscles used for breathing, are also used to actually shift the lungs within the body!
That’s just crazy awesome. Uriona & Farmer’s work raises the question of how prevalent this ability is in other semi-aquatic animals (e.g. seals, terrapins, manatees). By shifting the lungs further back in the body, crocodylians are able to change their local density. This allows the front, or back of the animal to rise and sink separately from the rest of the body. So too does moving the lungs from side to side allow for rolling in the water. All of this can occur without the need to move any external body parts. This means no extra turbulence gets created in the water, thus allowing crocodylians to better sneak up on their fishy, or fleshy prey.
Baby crocodiles exhibiting their unique pulmonary powers.
If anything, it sure speaks to why crocodyliformes have held dominion over the semi-aquatic niche for over 200 million years. Uriona and Farmer do suggest that the ability of these respiratory muscles to do this might not be an exaptation. Rather, this might have been the initial impetus behind the evolution of these muscles. Only later would they have been exapted to help with breathing on land. Though the authours provide some good parsimonious reasons for why this may be (basically it would take less evolutionary steps to accomplish than the other way around), it doesn’t really jive with the fossil evidence. Part of the reason why the crocodylian diaphragm works, is because the pubis (the forepart of the hip bone in most animals, and the part that juts out so prominently in theropod dinosaurs), is mobile. This mobility occurred early on in crocodyliforme evolution, with the crocodylomorph Protosuchus having a pubis that was almost mobile. The problem arises when one looks at this early crocodylomorph. Protosuchus was obviously terrestrial. If Protosuchus was evolving a mobile pubis already, then it was doubtful that it was being used to allow lung shifting in the body (an ability that is helpful when underwater, but pretty pointless on land). Furthermore, Crocodylia proper is the umpteenth time that crocodyliformes have returned to a semi-aquatic existence. It is doubtful that all the numerous land outings that occurred during crocodyliforme evolution, would have retained the ability to move the lungs to and fro. It seems far more likely that this was an ability that evolved in Crocodylia, or somewhere close by on the evolutionary tree, in some taxa that was still semi-aquatic.
Protosuchus richardsoni. An example of an early crocodylomorph.
Of course it is also possible that crocodyliforme phylogeny is just all f-ed up. With the amount of convergence rampant in that lot, this remains a distinct possibility.
Foraging mode has molded the evolution of many aspects of lizard biology. From a basic sit-and-wait sprinting feeding strategy, several lizard groups have evolved a wide foraging strategy, slowly moving through the environment using their highly developed chemosensory systems to locate prey. We studied locomotor performance, whole-body mechanics and gaits in a phylogenetic array of lizards that use sit-and-wait and wide-foraging strategies to contrast the functional differences associated with the need for speed vs slow continuous movement during foraging. Using multivariate and phylogenetic comparative analyses we tested for patterns of covariation in gaits and locomotor mechanics in relation to foraging mode. Sit-and-wait species used only fast speeds and trotting gaits coupled with running (bouncing) mechanics. Different wide-foraging species independently evolved slower locomotion with walking (vaulting) mechanics coupled with several different walking gaits, some of which have evolved several times. Most wide foragers retain the running mechanics with trotting gaits observed in sit-and-wait lizards, but some wide foragers have evolved very slow (high duty factor) running mechanics. In addition, three evolutionary reversals back to sit-and-wait foraging are coupled with the loss of walking mechanics. These findings provide strong evidence that foraging mode drives the evolution of biomechanics and gaits in lizards and that there are several ways to evolve slower locomotion. In addition, the different gaits used to walk slowly appear to match the ecological and behavioral challenges of the species that use them. Trotting appears to be a functionally stable strategy in lizards not necessarily related to whole-body mechanics or speed.
I haven’t had a chance to read much more than what was written already. I do take a bit of offense to the authours referring to scleroglossan foraging technique as “slow,” but what are you going to do?
I do find it interesting that lizards seem to have lost the ability to “walk” numerous times. That almost seems bizarre. The study points out that ecology produces heavy pressures on lizards in terms of their locomotion type. This is extremely pertinent given how often one hears the old (and wrong!) adage about “reptiles” being incapable of intense aerobic activity.
According to the above study (among others), it all depends on the animals being tested.
There we go. Two really cool papers on reptiles, being released in one day.
Yesterday a new study was released in the journal of Physiological and Biochemical Zoology. Researchers from the University of Utah, studied the effects of the well documented right-to-left shunt in crocodylians.Okay, let’s get the exposition out of the way first.
Mammals and birds are both characterized by a 4 chambered heart. This heart allows the complete separation of oxygenated and deoxygenated blood streams. Less publicized, but equally as important, this separation also allows for a pressure differential to exist between the two ventricular chambers. That way the right – pulmonary side – of the heart can pump deoxygenated blood at low pressure to the delicate walls of the alveoli in the lungs, while the left – systemic side – of the heart, can pump oxygenated blood at much higher pressure (~7 times higher) to the entire body.
Reptiles and amphibians differ from mammals and birds, in that they have a heart divided into 3 chambers (two atria, one ventricle). This allows for mixing of oxygenated and deoxygenated blood, which reduces aerobic efficiency.
Please note the qualifier: aerobic.
Now, as is often the case with herps, this is a rather broad generalization. The hearts of all reptiles, show various degrees of ventricular separation. Also, for all extant reptiles, there are physiological/haemodynamic mechanisms in place that reduce the amount of blood mixing. Meanwhile, some lizards (e.g. varanids), and snakes (e.g. pythons) have such a large muscular septum near the middle of their ventricle, that it actually completely separates the ventricle during the contractile phase (ventricular systole). Thus making varanids and various snakes, functionally four chambered. These reptiles are capable of producing pressures on their systemic side, that are 7 times higher than the pressures in their pulmonary side. In other words, their functional four chambered hearts allow for pressure differentials that are on par with mammals.
Then there are the crocodylians. Crocs have the most complicated heart of any vertebrate. They are the only reptiles that have evolved a complete seperation of their ventricles. They are anatomically four chambered. Yet, they also retain the ability to mix their oxygenated and deoxygenated blood supplies. This is accomplished through a small connection between the right and left aortic arches (which come out of each respective ventricle). This connection is referred to as the foramen of Panizza. Making things more interesting still, croc hearts also feature a cog toothed valve that can completely block the flow of blood to the lungs, thus turning their hearts into a double pump systemic circuit.
So now we know the how it works, the question we want answered next is: why did it evolve in the first place? The classic “orthodox” explanation has been that all of these traits evolved to allow formerly land dwelling crocodyliformes stay underwater for long periods of time. A four chambered heart is great for aerobic endurance, but pretty darn useless for an animal that spends most of its time holding its breath. In that arena, a three chambered heart is a more efficient system. By mixing oxygenated and deoxygenated blood together, crocodylians and other reptiles are able to siphon as much oxygen as possible from their blood, and thus stay underwater longer.
As I said, that was the old explanation. Now there is a new one:
Farmer, C.G., Uriona, T.J., Olsen, D.B., Steenblick, M., Sanders, K. The Right-to-left Shunt of Crocodilians Serves Digestion. Physiological and Biochemical Zoology. Vol. 81(2): 125-137. doi: 10.1086/524150
Farmer et al studied several groups of juvenile American alligators (Alligator mississippiensis). Each group underwent surgeries of various sorts to measure, and/or block the right to left shunt. The working hypothesis was that crocodylians use their right to left shunt, to serve digestion, by providing a greater reservoir of hydrogen ions (left over from the retention of CO2) for stomach acid secretion. It was suspected that if this was true, then one should see a greater degree of right to left shunting in animals that have just eaten.
So what did they find?
Well, for one, they found that juvenile alligators have a preferred postprandial body temperature of ~30?C, and will maintain that temperature to within .03?C. That’s a degree of temperature control worthy of any mammal, or bird.
Another thing they learned was that alligators that were allowed to stay at that temperature, were a real bugger to keep under control. So they had to drop the temp down 3 degrees, to 27?C instead.
Farmer et al learned that gastric acid secretion is temperature sensitive. Alligators produced greater quantities of gastric acid at 27?C, than at 19?C.
Oh yeah, they also learned that crocodylians produced a tonne of acid. At maximum secretion, acid production was an order of magnitude greater than that measured in any mammal, or bird. For those keeping tally at home; that’s 10 times greater.
The authours final observations warrant some thoughts.
That the left aorta, which arises from the right – pulmonary – ventricle, is the main blood delivery route for the digestive system. During right to left shunting, oxygenated blood from the left ventricle, gets shoved to the left aorta, and down to the digestive system. That this coincides with increased gastric acid secretion is telling, and strongly suggestive as to the role of the R-L shunt.
Yet R-L shunting also occurs during dives, and this is still the best explanation for the cog toothed valve. If the crocodylian heart really was specifically developed to increase digestion, then why block the path to the lungs at all? This study shows that the gastrointestinal system benefits from increased oxygen to these tissues. So why block the lungs, if one is trying to keep them oxygenated. Unfortunately the paper doesn’t really mention whether, or not the cog toothed valve was activated during this process. Personally, I don’t remember reading any case of the R-L shunt being used in crocs, without incorporating the cog tooth valve, so…
I felt that the authours put too much emphasis on endothermy vs. ectothermy. Their final observations involved a blanket statement regarding the R-L shunt in all reptiles. As I mentioned above, crocodylians are unique in their cardiovascular anatomy and physiology. They are also renowned for their very acidic stomach acid. It would seem more parsimonious to say that the R-L shunt in crocodylians, plays a large role in gastric acid secretion for these animals only; and wait for subsequent studies in other reptiles before saying this is true for the whole class.
Okay, so maybe their acid isn’t quite this strong, but you get the point.
Lastly (I know, I know, this just keeps going), I found it interesting that they studied the effects of gastric acid secretion on the vertebra of a cow. This vert took over 2 weeks to digest! While I can accept that this was partly due to the size of the object, and it’s material (bone is tough, after all.), but 2 weeks! Even at the lower temperature that the experimental group was kept at, it seems hard to believe. The authours gave no mention of gizzard usage in these animals, which suggests that the animals were never given access to gastroliths, which should have sped up the digestive process considerably.
Either way, the study was interesting. I just think that the authours took their final results a little too far.
I have been meaning to talk a bit about this one for a few days now. Getting around to doing this blog has been a bit of a hassle for me. I’m still not quite used to the whole “update often” format.Anywho, it appears that a group of gharials in a river sanctuary in India were dying from some enigmatic disease. Whatever the pathogen was/is, it appears to affect the liver and kidneys. So, the pathogen seems blood based then. Current thoughts seem to focus on pollution from the nearby Yamuna river. The animals may have eaten some badly contaminated fish.The end result has been the unfortunate demise of 50 animals so far. Given the critically endangered status of Gavialis gangeticus (they only live in India), setbacks like these are always a big deal. Hopefully this will turn out to be a localized event, and not a contagion.